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The Project Gutenberg eBook of Vegetable Teratology
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Title: Vegetable Teratology
Author: Maxwell T. Masters
Release date: November 6, 2007 [eBook #23354]
Language: English
Other information and formats: www.gutenberg.org/ebooks/23354
Credits: E-text prepared by Charlene Taylor, Leonard Johnson, and the
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*** START OF THE PROJECT GUTENBERG EBOOK VEGETABLE
TERATOLOGY ***
This eBook is for the use of anyone anywhere in the United States and
most other parts of the world at no cost and with almost no restrictions
whatsoever. You may copy it, give it away or re-use it under the terms
of the Project Gutenberg License included with this eBook or online at
www.gutenberg.org. If you are not located in the United States, you
will have to check the laws of the country where you are located
before using this eBook.
Title: Vegetable Teratology
Author: Maxwell T. Masters
Release date: November 6, 2007 [eBook #23354]
Language: English
Other information and formats: www.gutenberg.org/ebooks/23354
Credits: E-text prepared by Charlene Taylor, Leonard Johnson, and the
Project Gutenberg Online Distributed Proofreading Team
(https://www.pgdp.net)
*** START OF THE PROJECT GUTENBERG EBOOK VEGETABLE
TERATOLOGY ***
Page 4
E-text prepared by Charlene Taylor, Leonard Johnson,
and the Project Gutenberg Online Distributed Proofreading Team
(http://www.pgdp.net)
THE
RAY SOCIETY.
INSTITUTED MDCCCXLIV.
and the Project Gutenberg Online Distributed Proofreading Team
(http://www.pgdp.net)
THE
RAY SOCIETY.
INSTITUTED MDCCCXLIV.
Page 5
This volume is issued to the Subscribers to the Ray Society for the
Year 1868.
LONDON:
MDCCCLXIX.
VEGETABLE TERATOLOGY,
AN ACCOUNT
OF THE
PRINCIPAL DEVIATIONS FROM THE USUAL
CONSTRUCTION OF PLANTS
Year 1868.
LONDON:
MDCCCLXIX.
VEGETABLE TERATOLOGY,
AN ACCOUNT
OF THE
PRINCIPAL DEVIATIONS FROM THE USUAL
CONSTRUCTION OF PLANTS
Page 6
BY
MAXWELL T. MASTERS, M.D., F.L.S.
WITH NUMEROUS ILLUSTRATIONS BY
E. M. WILLIAMS.
LONDON:
PUBLISHED FOR THE RAY SOCIETY BY
ROBERT HARDWICKE, 192. PICCADILLY.
MDCCCLXIX.
TO
JOSEPH DALTON HOOKER, M.D.,
D.C.L., LL.D., F.R.S., F.L.S., ETC. ETC.
DIRECTOR OF THE ROYAL GARDENS, KEW,
This Volume
IS
GRATEFULLY INSCRIBED.
MAXWELL T. MASTERS, M.D., F.L.S.
WITH NUMEROUS ILLUSTRATIONS BY
E. M. WILLIAMS.
LONDON:
PUBLISHED FOR THE RAY SOCIETY BY
ROBERT HARDWICKE, 192. PICCADILLY.
MDCCCLXIX.
TO
JOSEPH DALTON HOOKER, M.D.,
D.C.L., LL.D., F.R.S., F.L.S., ETC. ETC.
DIRECTOR OF THE ROYAL GARDENS, KEW,
This Volume
IS
GRATEFULLY INSCRIBED.
Page 7
TABLE OF CONTENTS.
PAGE
INTRODUCTION xxi
BOOK I.
DEVIATIONS FROM ORDINARY ARRANGEMENT 1
PART I.—Union of Organs 8
CHAPTER I.
Cohesion 9
Cohesion between axes of same plant, 9—Fasciation, 11—Cohesion of foliar
organs, 21—Of margins of single organs, 21—Tubular petals, 23—Cohesion
of several organs by their margins, 25—Of the sepals, 27—Of the petals, 28—
Of the stamens, 29—Of the pistils, 29—Ascidia or pitchers, 30.
CHAPTER II.
Adhesion 32
Adhesion of foliar organs, 32—Of leaves by their surfaces, 33—Of foliar to
axile organs, 34—Of sepals to petals, 34—Of stamens to petals, 34—Of
stamens to pistils, 35—Miscellaneous adhesions, 35—Of fruit to branch, 36—
PAGE
INTRODUCTION xxi
BOOK I.
DEVIATIONS FROM ORDINARY ARRANGEMENT 1
PART I.—Union of Organs 8
CHAPTER I.
Cohesion 9
Cohesion between axes of same plant, 9—Fasciation, 11—Cohesion of foliar
organs, 21—Of margins of single organs, 21—Tubular petals, 23—Cohesion
of several organs by their margins, 25—Of the sepals, 27—Of the petals, 28—
Of the stamens, 29—Of the pistils, 29—Ascidia or pitchers, 30.
CHAPTER II.
Adhesion 32
Adhesion of foliar organs, 32—Of leaves by their surfaces, 33—Of foliar to
axile organs, 34—Of sepals to petals, 34—Of stamens to petals, 34—Of
stamens to pistils, 35—Miscellaneous adhesions, 35—Of fruit to branch, 36—
Page 8
Synanthy, 36—Syncarpy, 45—Synspermy, 50—Between axes of different
plants of same species, 50—And of different species, 55—Synophty.
PART II.—Independence of Organs 58
CHAPTER I.
Fission 59
Fission of axile organs, 60—Of foliar organs, 61—Of petals, 66—Of stamens,
68—Of carpels, 68.
CHAPTER II.
Dialysis 69
Dialysis of margins of individual parts, 70—Of margins of parts of same
whorl-calyx, 70—Of corolla, 71—Of stamens, 73—Of carpels, 73.
CHAPTER III.
Solution 76
Solution of calyx from ovary, 77—Of stamens from petals, 82.
PART III.—Alterations of Position 83
CHAPTER I.
Displacement 84
Displacement of bulbs, 84—Of inflorescence, 85—Of leaves, 87—Of parts of
flowers, 91—Of carpels, 96—Of placentas and ovules, 96.
CHAPTER II.
Prolification 100
Prolification of the inflorescence, 102—Median foliar, 103—Median floral,
105—Lateral foliar, 106—Lateral floral, 107—Prolification of the flower, 115
plants of same species, 50—And of different species, 55—Synophty.
PART II.—Independence of Organs 58
CHAPTER I.
Fission 59
Fission of axile organs, 60—Of foliar organs, 61—Of petals, 66—Of stamens,
68—Of carpels, 68.
CHAPTER II.
Dialysis 69
Dialysis of margins of individual parts, 70—Of margins of parts of same
whorl-calyx, 70—Of corolla, 71—Of stamens, 73—Of carpels, 73.
CHAPTER III.
Solution 76
Solution of calyx from ovary, 77—Of stamens from petals, 82.
PART III.—Alterations of Position 83
CHAPTER I.
Displacement 84
Displacement of bulbs, 84—Of inflorescence, 85—Of leaves, 87—Of parts of
flowers, 91—Of carpels, 96—Of placentas and ovules, 96.
CHAPTER II.
Prolification 100
Prolification of the inflorescence, 102—Median foliar, 103—Median floral,
105—Lateral foliar, 106—Lateral floral, 107—Prolification of the flower, 115
Page 9
—Median foliar, 116—Median floral, 119—Axillary prolification, 138—
Foliar, 141—Floral, 142—Complicated prolification, 151—Of embryo, 155.
CHAPTER III.
Heterotaxy 156
Formation of adventitious roots, 156—Of shoots below the cotyledons, 161—
Adventitious leaves, 162—On scapes, 163—Production of leaves or scales in
place of flower-buds, 164—Viviparous plants, 168—Formation of buds on
leaves, 170—In pith, 171—On bulbs, 172—Production of gemmæ in place of
spores, 173—Of flowers on leaves, 174—Of flower-buds in place of leaf-
buds, 176—Of flowers on spines, 177—Of flower-buds on petals, 177—On
fruits, 178—In ovaries, 180—Of stamens in ovaries, 182—Of pollen in
ovules, 185—Homomorphic flowers of Compositæ, 188—Heterotaxy
affecting the inflorescence, 188—Supra-soriferous ferns, 190.
CHAPTER IV.
Heterogamy 190
Change in the position of male and female flowers, 191—From monœcious to
diœcious condition, 193—From diœcious to monœcious, 193—From
hermaphroditism to unisexuality, 195—From unisexuality to
hermaphroditism, 197—Pollen replaced by ovules, 201.
CHAPTER V.
Alterations in the direction of organs 201
Fastigiation, 202—Eversion, 204—Altered direction of leaves, 205—Altered
direction of flower, 206—Reflexion, 209—Gymnaxony, 211.
BOOK II.
DEVIATIONS FROM ORDINARY FORM 213
PART I.—Stasimorphy 216
CHAPTER I.
Foliar, 141—Floral, 142—Complicated prolification, 151—Of embryo, 155.
CHAPTER III.
Heterotaxy 156
Formation of adventitious roots, 156—Of shoots below the cotyledons, 161—
Adventitious leaves, 162—On scapes, 163—Production of leaves or scales in
place of flower-buds, 164—Viviparous plants, 168—Formation of buds on
leaves, 170—In pith, 171—On bulbs, 172—Production of gemmæ in place of
spores, 173—Of flowers on leaves, 174—Of flower-buds in place of leaf-
buds, 176—Of flowers on spines, 177—Of flower-buds on petals, 177—On
fruits, 178—In ovaries, 180—Of stamens in ovaries, 182—Of pollen in
ovules, 185—Homomorphic flowers of Compositæ, 188—Heterotaxy
affecting the inflorescence, 188—Supra-soriferous ferns, 190.
CHAPTER IV.
Heterogamy 190
Change in the position of male and female flowers, 191—From monœcious to
diœcious condition, 193—From diœcious to monœcious, 193—From
hermaphroditism to unisexuality, 195—From unisexuality to
hermaphroditism, 197—Pollen replaced by ovules, 201.
CHAPTER V.
Alterations in the direction of organs 201
Fastigiation, 202—Eversion, 204—Altered direction of leaves, 205—Altered
direction of flower, 206—Reflexion, 209—Gymnaxony, 211.
BOOK II.
DEVIATIONS FROM ORDINARY FORM 213
PART I.—Stasimorphy 216
CHAPTER I.
Page 10
Persistence of juvenile forms 217
Stasimorphy in leaves of Conifers, 217—Regular peloria, 219.
PART II.—Pleiomorphy 228
CHAPTER I.
Irregular peloria 228
PART III.—Metamorphy 240
CHAPTER I.
Phyllody 241
Phyllody of bracts, 242—In inflorescence of Conifers, 245—Of calyx, 245—
Of corolla, 251—Of stamens, 253—Of pistils, 256—Of ovules, 262—
Changes in nucleus of ovule, 269—Phyllody of accessory organs, 272—
Chloranthy, 273—General remarks on, 278.
CHAPTER II.
Metamorphy of floral organs 281
Sepalody of petals, 282—Petalody of calyx, calycanthemy, 283—Petalody of
stamens, 285—Of anther, 291—Of connective, 293—Compound stamens,
294—Petalody of pistils, 296—Of ovules, 297—Of accessory organs, 297—
Staminody of the bracts, 298—Of sepals and petals, 298—Of pistils, 299—Of
accessory organs, 301—Pistillody of perianth, 302—Of sepals, 302—Of
stamens, 303—Of ovules, 310.
PART IV.—Heteromorphy 311
CHAPTER I.
Deformities 311
Formation of tubes, 312—In flower, 314—Spurs, 315—Contortion, 316—
Spiral torsion, 319—Of leaf, 326—Adventitious tendrils, 326—Interrupted
Stasimorphy in leaves of Conifers, 217—Regular peloria, 219.
PART II.—Pleiomorphy 228
CHAPTER I.
Irregular peloria 228
PART III.—Metamorphy 240
CHAPTER I.
Phyllody 241
Phyllody of bracts, 242—In inflorescence of Conifers, 245—Of calyx, 245—
Of corolla, 251—Of stamens, 253—Of pistils, 256—Of ovules, 262—
Changes in nucleus of ovule, 269—Phyllody of accessory organs, 272—
Chloranthy, 273—General remarks on, 278.
CHAPTER II.
Metamorphy of floral organs 281
Sepalody of petals, 282—Petalody of calyx, calycanthemy, 283—Petalody of
stamens, 285—Of anther, 291—Of connective, 293—Compound stamens,
294—Petalody of pistils, 296—Of ovules, 297—Of accessory organs, 297—
Staminody of the bracts, 298—Of sepals and petals, 298—Of pistils, 299—Of
accessory organs, 301—Pistillody of perianth, 302—Of sepals, 302—Of
stamens, 303—Of ovules, 310.
PART IV.—Heteromorphy 311
CHAPTER I.
Deformities 311
Formation of tubes, 312—In flower, 314—Spurs, 315—Contortion, 316—
Spiral torsion, 319—Of leaf, 326—Adventitious tendrils, 326—Interrupted
Page 11
growth, 327—Cornute leaves, 328—Flattening, 328.
CHAPTER II.
Polymorphy 329
Heterophylly, 330—Dimorphism, 333—Sports or bud-variations, 336.
CHAPTER III.
Alterations of colour 337
Albinism, 337—Virescence, 338—Chromatism, 339.
BOOK III.
DEVIATIONS FROM ORDINARY NUMBER 340
PART I.—Increased Number of Organs 343
CHAPTER I.
Multiplication of axile organs—inflorescence 346
Multiplication of branches, 346—Plica, 346—Polyclady, 347—Multiplication
of branches of inflorescence, 348—Of bulbs, 350—Of florets, 351.
CHAPTER II.
Multiplication of foliar organs 352
Pleiophylly, 353—Multiplication of stipules, bracts, &c., 357—Polyphylly,
358—Increased number of leaves in a whorl, 358—Polyphylly of bracts, 358
—Of calyx, 358—Of corolla, 359—Of andrœcium, 361—Of gynœcium, 363
—Of flower in general, 365—Increased number of ovules and seeds, 367—Of
embryos, 369—Of cotyledons, 370—Pleiotaxy, or multiplication of whorls,
371—Pleiotaxy of bracts, 371—Of calyx, 374—Of perianth, 375—Of corolla
(hose in hose), 376—Andrœcium, 379—Andrœcium of Orchids, 380—
Pleiotaxy of gynœcium, 388—Increased number of flowers in an
inflorescence, 391.
CHAPTER II.
Polymorphy 329
Heterophylly, 330—Dimorphism, 333—Sports or bud-variations, 336.
CHAPTER III.
Alterations of colour 337
Albinism, 337—Virescence, 338—Chromatism, 339.
BOOK III.
DEVIATIONS FROM ORDINARY NUMBER 340
PART I.—Increased Number of Organs 343
CHAPTER I.
Multiplication of axile organs—inflorescence 346
Multiplication of branches, 346—Plica, 346—Polyclady, 347—Multiplication
of branches of inflorescence, 348—Of bulbs, 350—Of florets, 351.
CHAPTER II.
Multiplication of foliar organs 352
Pleiophylly, 353—Multiplication of stipules, bracts, &c., 357—Polyphylly,
358—Increased number of leaves in a whorl, 358—Polyphylly of bracts, 358
—Of calyx, 358—Of corolla, 359—Of andrœcium, 361—Of gynœcium, 363
—Of flower in general, 365—Increased number of ovules and seeds, 367—Of
embryos, 369—Of cotyledons, 370—Pleiotaxy, or multiplication of whorls,
371—Pleiotaxy of bracts, 371—Of calyx, 374—Of perianth, 375—Of corolla
(hose in hose), 376—Andrœcium, 379—Andrœcium of Orchids, 380—
Pleiotaxy of gynœcium, 388—Increased number of flowers in an
inflorescence, 391.
Page 12
PART II.—Diminished Number of Organs 392
CHAPTER I.
Suppression of Axile Organs 393
Acaulosia, 393—Non-development of peduncle, 393—Nature of calyx-tube
(Casimir de Candolle), 394—Suppression of columella, 395.
CHAPTER II.
Suppression of Foliar Organs 395
Aphylly, 395—Meiophylly, 396—Of calyx or perianth, 396—Of corolla, 397
—Of andrœcium, 398—Of gynœcium, 399—Of flower, 400—Meiotaxy, 403
—Of calyx, 403—Of corolla, 403—Of andrœcium, 405—Of gynœcium, 406
—Of ovules or seeds, 407—Of flower, 408—General remarks on suppression,
409.
BOOK IV.
DEVIATIONS FROM ORDINARY SIZE AND
CONSISTENCE 413
PART I—Hypertrophy 416
CHAPTER I.
Enlargement 417
Of axile organs, 418—Knaurs, 419—Enlargement of buds, 420—Of flower-
stalk, 421—In pears, 423—Of placenta, 424—Of leaves, 426—Development
of parts usually abortive, 427—Enlargement of perianth, 428—Of andrœcium,
430—Of gynœcium, 430—Of fruit, 431—Alterations of consistence, 432.
CHAPTER II.
Elongation 488
CHAPTER I.
Suppression of Axile Organs 393
Acaulosia, 393—Non-development of peduncle, 393—Nature of calyx-tube
(Casimir de Candolle), 394—Suppression of columella, 395.
CHAPTER II.
Suppression of Foliar Organs 395
Aphylly, 395—Meiophylly, 396—Of calyx or perianth, 396—Of corolla, 397
—Of andrœcium, 398—Of gynœcium, 399—Of flower, 400—Meiotaxy, 403
—Of calyx, 403—Of corolla, 403—Of andrœcium, 405—Of gynœcium, 406
—Of ovules or seeds, 407—Of flower, 408—General remarks on suppression,
409.
BOOK IV.
DEVIATIONS FROM ORDINARY SIZE AND
CONSISTENCE 413
PART I—Hypertrophy 416
CHAPTER I.
Enlargement 417
Of axile organs, 418—Knaurs, 419—Enlargement of buds, 420—Of flower-
stalk, 421—In pears, 423—Of placenta, 424—Of leaves, 426—Development
of parts usually abortive, 427—Enlargement of perianth, 428—Of andrœcium,
430—Of gynœcium, 430—Of fruit, 431—Alterations of consistence, 432.
CHAPTER II.
Elongation 488
Page 13
Elongation of root, 434—Of inflorescence, 434—Of flower-stalks, 435—Of
leaves, 437—Of parts of flower, 438—Of thalamus and placenta, 440—
Apostasis, 440.
CHAPTER III.
Enation 443
Excrescences from axile organs, warts, 444—Enation from leaves, 445—
From sepals, 448—From petals, 448—Catacorolla, 451—Enation from
stamens, 453—From carpels, 453.
PART II.—Atrophy 454
CHAPTER I.
Abortion 455
Abortion of axile organs, 455—Of receptacle, 457—Of leaves, 458—Of
perianth, calyx, and corolla, 460—Of stamens, 463—Of pollen, 463—Of
pistil, fruit, &c., 464—Of ovules, 466—Depauperated ferns, 466—General
remarks, 467.
CHAPTER II.
Degeneration 470
Formation of scales, 470—Of hairs, 472—Of glands, 473—Of tendrils, 473.
GENERAL CONCLUSIONS.
General morphology of the leaf and axis, 476.—Homology, 476—Special
morphology, 479—Calyx-tube, 480—Andrœcium, 482—Inferior ovary, 482
—Placentation, 483—Structure of the ovule, 484—Leaves of Conifers, 484—
Relative position of organs, 484—Law of alternation, 485—Co-relation, 486
—Compensation, 488—Teratology and classification, 488.
APPENDIX.
leaves, 437—Of parts of flower, 438—Of thalamus and placenta, 440—
Apostasis, 440.
CHAPTER III.
Enation 443
Excrescences from axile organs, warts, 444—Enation from leaves, 445—
From sepals, 448—From petals, 448—Catacorolla, 451—Enation from
stamens, 453—From carpels, 453.
PART II.—Atrophy 454
CHAPTER I.
Abortion 455
Abortion of axile organs, 455—Of receptacle, 457—Of leaves, 458—Of
perianth, calyx, and corolla, 460—Of stamens, 463—Of pollen, 463—Of
pistil, fruit, &c., 464—Of ovules, 466—Depauperated ferns, 466—General
remarks, 467.
CHAPTER II.
Degeneration 470
Formation of scales, 470—Of hairs, 472—Of glands, 473—Of tendrils, 473.
GENERAL CONCLUSIONS.
General morphology of the leaf and axis, 476.—Homology, 476—Special
morphology, 479—Calyx-tube, 480—Andrœcium, 482—Inferior ovary, 482
—Placentation, 483—Structure of the ovule, 484—Leaves of Conifers, 484—
Relative position of organs, 484—Law of alternation, 485—Co-relation, 486
—Compensation, 488—Teratology and classification, 488.
APPENDIX.
Page 14
Double flowers, varieties of, 491—Causes of production, 491—Relation to
variegated foliage, 497—List of plants producing, 499.
NOTE 508
INDEX TO SUBJECTS 511
INDEX OF NAMES OF PLANTS 517
variegated foliage, 497—List of plants producing, 499.
NOTE 508
INDEX TO SUBJECTS 511
INDEX OF NAMES OF PLANTS 517
Page 15
LIST OF ILLUSTRATIONS.
FIG. PAGE
1. Diagram of regular pentamerous flower 4
2. Cohesion of branch Dipsacus sylvestris 10
3. Fasciated lettuce ('Gard. Chron.') 11
4. Fasciation in Asparagus 12
5. Fasciation in Pinus Pinaster 13
6. Fasciation and spiral torsion in Asparagus
('Gard. Chron.') 14
7. Fasciation in scape of dandelion 16
8. Pitcher on leaf of Pelargonium 22
9. Transition from flat to tubular segments of
the perianth in Eranthis 24
10. Pitcher of Crassula arborescens (C. Morren)
26
11. Gamopetalous corolla, Papaver bracteatum
28
12. Adhesion of petaloid stamen to segment of
perianth, Crocus 35
13. Adhesion of petal, stamen and carpel,
Cheiranthus Cheiri 36
14. Synanthy. Campanula Medium ('Gard.
Chron.') 37
15, 16. Synanthic flowers of Calanthe vestita 39
17. Synanthy in Digitalis purpurea ('Gard.
Chron.') 40
FIG. PAGE
1. Diagram of regular pentamerous flower 4
2. Cohesion of branch Dipsacus sylvestris 10
3. Fasciated lettuce ('Gard. Chron.') 11
4. Fasciation in Asparagus 12
5. Fasciation in Pinus Pinaster 13
6. Fasciation and spiral torsion in Asparagus
('Gard. Chron.') 14
7. Fasciation in scape of dandelion 16
8. Pitcher on leaf of Pelargonium 22
9. Transition from flat to tubular segments of
the perianth in Eranthis 24
10. Pitcher of Crassula arborescens (C. Morren)
26
11. Gamopetalous corolla, Papaver bracteatum
28
12. Adhesion of petaloid stamen to segment of
perianth, Crocus 35
13. Adhesion of petal, stamen and carpel,
Cheiranthus Cheiri 36
14. Synanthy. Campanula Medium ('Gard.
Chron.') 37
15, 16. Synanthic flowers of Calanthe vestita 39
17. Synanthy in Digitalis purpurea ('Gard.
Chron.') 40
Page 16
18. Synanthy in Calceolaria 41
19, 20. Syncarpic apples 47
21. Adhesion of two stems of oak ('Gard.
Chron.') 51
22. Adhesion of branches of elm ('Gard. Chron.')
52
23. Adhesion of two roots of carrot, one white,
the other red ('Gard. Chron.') 53
24. Section through inverted and adherent
mushrooms 54
25. Bifurcated male catkin, Cedrus Libani 61
26. Bifurcated leaf, Lamium album 62
27. Bifurcated leaf, Pelargonium 63
28. Bifurcated frond, Scolopendrium 64
29. Three-lobed lip of Oncidium 68
30. Dialysis of corolla in Correa 71
31. Dialysis of corolla in Campanula (De
Candolle) 72
32. Anomalous form of orange 74
33, 34. Disjoined carpels of orange (Maout) 75
35. Proliferous rose (Bell Salter) 78
36, 37. Apple flower, with detached calyx, &c. 79
38. Flower of Œnanthe crocata, with detached
calyx, &c. 80
39. Anomalous bulbs of tulip 85
40. Displaced leaf of Gesnera (C. Morren) 88
41. Leaves of Pinus pinea 89
42. Deranged leaves of yew 90
43. Cohesion of sepals and displacement of parts
of Oncidium cucullatum 92
44. Malformed flower of Cypripedium 93
45. Diagram of the same 93
46. Diagram of natural structure in Cypripedium
93
47. Diagram of malformed flower of Lycaste
Skinneri 93
19, 20. Syncarpic apples 47
21. Adhesion of two stems of oak ('Gard.
Chron.') 51
22. Adhesion of branches of elm ('Gard. Chron.')
52
23. Adhesion of two roots of carrot, one white,
the other red ('Gard. Chron.') 53
24. Section through inverted and adherent
mushrooms 54
25. Bifurcated male catkin, Cedrus Libani 61
26. Bifurcated leaf, Lamium album 62
27. Bifurcated leaf, Pelargonium 63
28. Bifurcated frond, Scolopendrium 64
29. Three-lobed lip of Oncidium 68
30. Dialysis of corolla in Correa 71
31. Dialysis of corolla in Campanula (De
Candolle) 72
32. Anomalous form of orange 74
33, 34. Disjoined carpels of orange (Maout) 75
35. Proliferous rose (Bell Salter) 78
36, 37. Apple flower, with detached calyx, &c. 79
38. Flower of Œnanthe crocata, with detached
calyx, &c. 80
39. Anomalous bulbs of tulip 85
40. Displaced leaf of Gesnera (C. Morren) 88
41. Leaves of Pinus pinea 89
42. Deranged leaves of yew 90
43. Cohesion of sepals and displacement of parts
of Oncidium cucullatum 92
44. Malformed flower of Cypripedium 93
45. Diagram of the same 93
46. Diagram of natural structure in Cypripedium
93
47. Diagram of malformed flower of Lycaste
Skinneri 93
Page 17
48. Diagram of malformed flower of
Dendrobium nobile 94
49. Natural arrangement in same flower 94
50. Diagram of malformed violet 94
51. Monstrous flower of Cerastium ('Gard.
Chron.') 97
52. Inflorescence of Polyanthus, with tufts of
leaves at the summit 105
53. Lateral prolification in inflorescence of
Pelargonium 108
54. Paniculate inflorescence of Plantago major
109
55. Branched spike and leafy bracts of the same
110
56. Inflorescence of Plantago lanceolata, with
leaves and secondary flower-stalks at its summit
111
57. Branched inflorescence of Reseda Luteola
112
58. Thalamus of strawberry prolonged into a
leafy branch ('American Agriculturist') 116
59. Flower of Verbascum, with dialysis of calyx
and corolla and prolonged thalamus 116
60. Median floral prolification of Dianthus 120
61. Leafy carpels and prolification of Daucus
Carota 123
62. Median floral prolification of Delphinium
126
63. Median prolification, &c., of Orchis
pyramidalis 128
64. Proliferous rose (Bell Salter) 130
65. Axillary floral prolification of Nymphæa
Lotus ('Gard. Chron.') 144
66. Axillary floral prolification of Dianthus 146
67. Proliferous rose 151
68. Proliferous rose ('Gard. Chron.') 152
Dendrobium nobile 94
49. Natural arrangement in same flower 94
50. Diagram of malformed violet 94
51. Monstrous flower of Cerastium ('Gard.
Chron.') 97
52. Inflorescence of Polyanthus, with tufts of
leaves at the summit 105
53. Lateral prolification in inflorescence of
Pelargonium 108
54. Paniculate inflorescence of Plantago major
109
55. Branched spike and leafy bracts of the same
110
56. Inflorescence of Plantago lanceolata, with
leaves and secondary flower-stalks at its summit
111
57. Branched inflorescence of Reseda Luteola
112
58. Thalamus of strawberry prolonged into a
leafy branch ('American Agriculturist') 116
59. Flower of Verbascum, with dialysis of calyx
and corolla and prolonged thalamus 116
60. Median floral prolification of Dianthus 120
61. Leafy carpels and prolification of Daucus
Carota 123
62. Median floral prolification of Delphinium
126
63. Median prolification, &c., of Orchis
pyramidalis 128
64. Proliferous rose (Bell Salter) 130
65. Axillary floral prolification of Nymphæa
Lotus ('Gard. Chron.') 144
66. Axillary floral prolification of Dianthus 146
67. Proliferous rose 151
68. Proliferous rose ('Gard. Chron.') 152
Page 18
69. Diagram of prolified orchid 153
70. Diagram of prolified orchis 154
71. Adventitious roots from petiole of celery 158
72. Germinating plant of mango 159
73, 74. Adventitious roots from leaves 160
75. Hip of rose bearing leaf 162
76. Leaves proceeding from the ovary of
Nymphæa sp. 162
77. Flower-stalk of dandelion, with leaves 163
78. Tuft of leaves in place of flowers in Valeriana
sp. 165
79. Scale-bearing spikelets of Willdenovia 167
80. "Rose Willow" 167
81. Viviparous flowers of Aira vivipara 169
82. Formation of shoot on leaf of Episcia bicolor
171
83. Adventitious buds on root of sea-kale 172
84, 85. Production of adventitious bulbs in
hyacinth ('Gard. Chron.') 172
86. Adventitious buds on hyacinth ('Gard. Chron
') 173
87–92. Nepaul barley 174, 175
93. Formation of buds on fruit of Opuntia 179
94. Flower-bud in the pod of Sinapis 181
94*, 95. Adventitious pod in silique of
Cheiranthus 182
96, 97. Grapes, with adventitious fruits in interior
183
98. Stamens in ovary of Bæckea diosmifolia 184
99. Pollen in ovule of passion flower (S. J.
Salter) 185
100. Female flowers at the summit of the
inflorescence of Carex acuta 192
101. Monœcious hop ('Gard. Chron.') 193
102. Superior ovary, &c., of fuchsia 198
103. Hermaphrodite flower of Carica 199
70. Diagram of prolified orchis 154
71. Adventitious roots from petiole of celery 158
72. Germinating plant of mango 159
73, 74. Adventitious roots from leaves 160
75. Hip of rose bearing leaf 162
76. Leaves proceeding from the ovary of
Nymphæa sp. 162
77. Flower-stalk of dandelion, with leaves 163
78. Tuft of leaves in place of flowers in Valeriana
sp. 165
79. Scale-bearing spikelets of Willdenovia 167
80. "Rose Willow" 167
81. Viviparous flowers of Aira vivipara 169
82. Formation of shoot on leaf of Episcia bicolor
171
83. Adventitious buds on root of sea-kale 172
84, 85. Production of adventitious bulbs in
hyacinth ('Gard. Chron.') 172
86. Adventitious buds on hyacinth ('Gard. Chron
') 173
87–92. Nepaul barley 174, 175
93. Formation of buds on fruit of Opuntia 179
94. Flower-bud in the pod of Sinapis 181
94*, 95. Adventitious pod in silique of
Cheiranthus 182
96, 97. Grapes, with adventitious fruits in interior
183
98. Stamens in ovary of Bæckea diosmifolia 184
99. Pollen in ovule of passion flower (S. J.
Salter) 185
100. Female flowers at the summit of the
inflorescence of Carex acuta 192
101. Monœcious hop ('Gard. Chron.') 193
102. Superior ovary, &c., of fuchsia 198
103. Hermaphrodite flower of Carica 199
Page 19
104. Ovuliferous anthers of Cucurbita sp. 200
105, 106. Prolonged inflorescence of fig 205
107. Hollow turnip, with inverted leaves 206
108. Normal flower of Gloxinia 207
109. Regular peloria of Gloxinia 207
110, 111. Structural details of erect Gloxinia 208
112. Reflected corolla of azalea 209
113, 114. Enlarged and erect placenta of Cuphea
miniata (C. Morren) 210
115. Dimorphic leaves of Juniperus sinensis 217
116. Regular peloria of Delphinium 219
117. Structural details of the preceding 219
118. Regular peloria of violet 220
119. Double-flowered regular violet 220
120. Regular peloria of Eccremocarpus scaber
222
121. Regular peloria of Cattleya 223
122. Peloria of Calceolaria 230
123. Peloric flower of Aristolochia 232
124, 125. Peloric flowers of Corydalis 236
126. Rose plantain 242
127. Leafy bracts in Plantago major 243
128. Leafy scales of Dahlia 244
129. Leafy sepals of rose 246
130. Leafy sepals of Fuchsia ('Gard. Chron.') 247
131. Leafy calyx of primrose 248
132. Leafy calyx of melon 248
133. Leafy sepals and petals of Geranium 251
134. Leafy stamens of Petunia 254
135, 136. Leafy anthers of Jatropha (Müller) 255
137. Proliferous rose, with leafy carpels, &c. (Bell
Salter) 257
138. Cucumber, with adventitious leaf attached (S.
J. Salter) 258
139. Leafy carpels in flower of Triumfetta 260
140. Leafy ovules of Sinapis 264
105, 106. Prolonged inflorescence of fig 205
107. Hollow turnip, with inverted leaves 206
108. Normal flower of Gloxinia 207
109. Regular peloria of Gloxinia 207
110, 111. Structural details of erect Gloxinia 208
112. Reflected corolla of azalea 209
113, 114. Enlarged and erect placenta of Cuphea
miniata (C. Morren) 210
115. Dimorphic leaves of Juniperus sinensis 217
116. Regular peloria of Delphinium 219
117. Structural details of the preceding 219
118. Regular peloria of violet 220
119. Double-flowered regular violet 220
120. Regular peloria of Eccremocarpus scaber
222
121. Regular peloria of Cattleya 223
122. Peloria of Calceolaria 230
123. Peloric flower of Aristolochia 232
124, 125. Peloric flowers of Corydalis 236
126. Rose plantain 242
127. Leafy bracts in Plantago major 243
128. Leafy scales of Dahlia 244
129. Leafy sepals of rose 246
130. Leafy sepals of Fuchsia ('Gard. Chron.') 247
131. Leafy calyx of primrose 248
132. Leafy calyx of melon 248
133. Leafy sepals and petals of Geranium 251
134. Leafy stamens of Petunia 254
135, 136. Leafy anthers of Jatropha (Müller) 255
137. Proliferous rose, with leafy carpels, &c. (Bell
Salter) 257
138. Cucumber, with adventitious leaf attached (S.
J. Salter) 258
139. Leafy carpels in flower of Triumfetta 260
140. Leafy ovules of Sinapis 264
Page 20
141, 142. Leafy ovules of Trifolium repens
(Caspary) 265
143. Portion of leafy carpel of Delphinium, with
ovules (Cramer) 266
144, 145. Enlarged view of section of leafy carpel,
&c., of Delphinium (Cramer) 267
146. Placentæ of Dianthus, bearing ovules and
carpels ('Gard. Chron.') 268
147. Ovules passing into carpels, Dianthus ('Gard.
Chron.') 268
148. Leafy shoot in place of ovule of Gaillardia
270
149, 150. Leafy sepals, petals, &c., of Epilobium
273
151. Leafy carpel of rose, with deformed ovules
274
152. Flower of St. Valery apple 282
153. Petaloid calyx of Mimulus 284
154. Double stellate columbine 287
155. Four-winged filaments of Rhododendron 290
156. Diagram of malformed flower of Catasetum
291
157. Petaloid stamen of Viola 292
158. Double columbine 293
159. Petaloid stamens of Hibiscus 293
160. Displaced coloured leaf, &c., of tulip 302
161. Supernumerary carpels in orange (Maout)
303
162. Pistilloid stamens of poppy 304
163. Pistilloid stamens of wallflower, &c. 306
164. Passage of stamen to carpel in lily 307
165. Transition of stamens to carpels,
Sempervivum tectorum 309
166. Ascidia of cabbage 312
167. Stalked pitcher on lettuce leaf 313
168. Tubular petal of Primula sinensis 315
(Caspary) 265
143. Portion of leafy carpel of Delphinium, with
ovules (Cramer) 266
144, 145. Enlarged view of section of leafy carpel,
&c., of Delphinium (Cramer) 267
146. Placentæ of Dianthus, bearing ovules and
carpels ('Gard. Chron.') 268
147. Ovules passing into carpels, Dianthus ('Gard.
Chron.') 268
148. Leafy shoot in place of ovule of Gaillardia
270
149, 150. Leafy sepals, petals, &c., of Epilobium
273
151. Leafy carpel of rose, with deformed ovules
274
152. Flower of St. Valery apple 282
153. Petaloid calyx of Mimulus 284
154. Double stellate columbine 287
155. Four-winged filaments of Rhododendron 290
156. Diagram of malformed flower of Catasetum
291
157. Petaloid stamen of Viola 292
158. Double columbine 293
159. Petaloid stamens of Hibiscus 293
160. Displaced coloured leaf, &c., of tulip 302
161. Supernumerary carpels in orange (Maout)
303
162. Pistilloid stamens of poppy 304
163. Pistilloid stamens of wallflower, &c. 306
164. Passage of stamen to carpel in lily 307
165. Transition of stamens to carpels,
Sempervivum tectorum 309
166. Ascidia of cabbage 312
167. Stalked pitcher on lettuce leaf 313
168. Tubular petal of Primula sinensis 315
Page 21
169. Spurs on flower of Calceolaria 316
170. Contorted stem of Juncus 317
171. Contorted branch of Cratægus 317
172. Spirally-twisted stem of teazel 321
173. Spirally-twisted stem Galium 323
174. Spirally-twisted root ('Gard. Chron.') 324
175. Interrupted growth of radish (American
Agriculturist) 327
176. Interrupted growth in apple 327
177. Polymorphous leaves of lilac 331
178. Adventitious growth on frond of Pteris
quadriaurita 333
179, 180. Coloured flower-stalks of feather
hyacinth (C. Morren) 347, 348
181. Multiplication of catkins, Corylus 349
182. Branched inflorescence of broccoli ('Gard.
Chron.') 351
183, 184. Supernumerary leaf of elm 353, 354
185. Supernumerary leaf of hazel 355
186. Multiplication of parts of flower in a plum
('Gard. Chron.') 366
187. Wheat-ear carnation 372
188. Multiplication of bracts in Delphinium
Consolida 373
189. Multiplication of bracts in Pelargonium 373
190. Double white lily 376
191. Double flower of Campanula rotundifolia
378
192. Diagram of usual arrangement of parts in
Orchis (Darwin) 381
193. Diagram of malformed flower of Ophrys
aranifera 385
194. Malformed flower of Ophrys aranifera 385
195. Diagram of malformed flower of Orchis
mascula (Cramer) 386
196. Multiplication of carpels, Tulip 388
170. Contorted stem of Juncus 317
171. Contorted branch of Cratægus 317
172. Spirally-twisted stem of teazel 321
173. Spirally-twisted stem Galium 323
174. Spirally-twisted root ('Gard. Chron.') 324
175. Interrupted growth of radish (American
Agriculturist) 327
176. Interrupted growth in apple 327
177. Polymorphous leaves of lilac 331
178. Adventitious growth on frond of Pteris
quadriaurita 333
179, 180. Coloured flower-stalks of feather
hyacinth (C. Morren) 347, 348
181. Multiplication of catkins, Corylus 349
182. Branched inflorescence of broccoli ('Gard.
Chron.') 351
183, 184. Supernumerary leaf of elm 353, 354
185. Supernumerary leaf of hazel 355
186. Multiplication of parts of flower in a plum
('Gard. Chron.') 366
187. Wheat-ear carnation 372
188. Multiplication of bracts in Delphinium
Consolida 373
189. Multiplication of bracts in Pelargonium 373
190. Double white lily 376
191. Double flower of Campanula rotundifolia
378
192. Diagram of usual arrangement of parts in
Orchis (Darwin) 381
193. Diagram of malformed flower of Ophrys
aranifera 385
194. Malformed flower of Ophrys aranifera 385
195. Diagram of malformed flower of Orchis
mascula (Cramer) 386
196. Multiplication of carpels, Tulip 388
Page 22
197. Section of St. Valery apple 388
198. Regular dimerous flower of Calanthe vestita
402
199. Regular dimerous flower of Odontoglossum
Alexandræ 402
200. Hypertrophied branch of Pelargonium 418
201. Tubers in the axils of leaves of the potato 420
202. Hypertrophied pedicels of ash 421
203, 204. Hypertrophy and elongation of flower-
stalk, &c., in pears 422, 423
205. Hypertrophied perianth, Cocos nucifera 428
206. Elongation of flower-stalk, Ranunculus acris
436
207. Linear leaf-lobes of parsley 438
208. Passage of pinnate to palmate leaves in horse-
chestnut 439
209. Elongation of thalamus, apostasis, &c., in
flower of Delphinium (Cramer) 441
210. Adventitious growths from cabbage leaf 445
211. Crested fronds of Nephrodium molle 447
212. Supernumerary petals, &c., Datura fastuosa
450
213. Supernumerary petaloid segments in flower
of Gloxinia 451
214. Catacorolla of Gloxinia (E. Morren) 452
215. Atrophied leaves of cabbage 460
216. Abortion of petals, pansy 461
217. Flower of Oncidium abortivum 462
218. Bladder plum 464
198. Regular dimerous flower of Calanthe vestita
402
199. Regular dimerous flower of Odontoglossum
Alexandræ 402
200. Hypertrophied branch of Pelargonium 418
201. Tubers in the axils of leaves of the potato 420
202. Hypertrophied pedicels of ash 421
203, 204. Hypertrophy and elongation of flower-
stalk, &c., in pears 422, 423
205. Hypertrophied perianth, Cocos nucifera 428
206. Elongation of flower-stalk, Ranunculus acris
436
207. Linear leaf-lobes of parsley 438
208. Passage of pinnate to palmate leaves in horse-
chestnut 439
209. Elongation of thalamus, apostasis, &c., in
flower of Delphinium (Cramer) 441
210. Adventitious growths from cabbage leaf 445
211. Crested fronds of Nephrodium molle 447
212. Supernumerary petals, &c., Datura fastuosa
450
213. Supernumerary petaloid segments in flower
of Gloxinia 451
214. Catacorolla of Gloxinia (E. Morren) 452
215. Atrophied leaves of cabbage 460
216. Abortion of petals, pansy 461
217. Flower of Oncidium abortivum 462
218. Bladder plum 464
Page 23
INTRODUCTION.
Till within a comparatively recent period but little study was given to
exceptional formations. They were considered as monsters to be shunned,
as lawless deviations from the ordinary rule, unworthy the attention of
botanists, or at best as objects of mere curiosity. By those whose notions of
structure and conformation did not extend beyond the details necessary to
distinguish one species from another, or to describe the salient features of a
plant in technical language; whose acquaintance with botanical science
might almost be said to consist in the conventional application of a number
of arbitrary terms, or in the recollection of a number of names, teratology
was regarded as a chaos whose meaningless confusion it were vain to
attempt to render intelligible,—as a barren field not worth the labour of
tillage.
The older botanists, it is true, often made them the basis of satirical
allusions to the political or religious questions of the day, especially about
the time of the Reformation, and the artists drew largely upon their
polemical sympathies in their representations of these anomalies. Linnæus
treated of them to some extent in his 'Philosophia,' but it is mainly to
Angustin Pyramus De Candolle that the credit is due of calling attention to
the importance of vegetable teratology. This great botanist, not only
indirectly, but from his personal research into the nature of monstrosities,
Till within a comparatively recent period but little study was given to
exceptional formations. They were considered as monsters to be shunned,
as lawless deviations from the ordinary rule, unworthy the attention of
botanists, or at best as objects of mere curiosity. By those whose notions of
structure and conformation did not extend beyond the details necessary to
distinguish one species from another, or to describe the salient features of a
plant in technical language; whose acquaintance with botanical science
might almost be said to consist in the conventional application of a number
of arbitrary terms, or in the recollection of a number of names, teratology
was regarded as a chaos whose meaningless confusion it were vain to
attempt to render intelligible,—as a barren field not worth the labour of
tillage.
The older botanists, it is true, often made them the basis of satirical
allusions to the political or religious questions of the day, especially about
the time of the Reformation, and the artists drew largely upon their
polemical sympathies in their representations of these anomalies. Linnæus
treated of them to some extent in his 'Philosophia,' but it is mainly to
Angustin Pyramus De Candolle that the credit is due of calling attention to
the importance of vegetable teratology. This great botanist, not only
indirectly, but from his personal research into the nature of monstrosities,
Page 24
did more than any of his predecessors to rescue them from the utter
disregard, or at best the contemptuous indifference, of the majority of
botanists. De Candolle gave a special impetus to morphology in general by
giving in his adhesion to the morphological hypotheses of Goethe. These
were no mere figments of the poet's imagination, as they were to a large
extent based on the actual investigation of normal and abnormal
organisation by Goethe both alone, and also in conjunction with Batsch and
Jaeger.
De Candolle's example was contagious. Scarcely a botanist of any eminence
since his time but has contributed his quota to the records of vegetable
teratology, in proof of which the names of Humboldt, Robert Brown, the De
Jussieus, the Saint Hilaires, of Moquin-Tandon, of Lindley, and many
others, not to mention botanists still living, may be cited. To students and
amateurs the subject seems always to have presented special attractions,
probably from the singularity of the appearances presented, and from the
fact that in many cases the examination of individual instances of
malformation can be carried on, to a large extent, without the lengthened or
continuous investigation and critical comparative study required by other
departments of botanical science. Be this as it may, teratology owes a very
large number of its records to this class of observers.
While the number of scattered papers on vegetable teratology in various
European languages is so great as to preclude the possibility of collating
them all, there is no general treatise on the subject in the English language,
with the exception of Hopkirk's 'Flora Anomala,' a book now rarely met
with, and withal very imperfect; and this notwithstanding that Robert
Brown early lent his sanction to the doctrines of Goethe, and himself
illustrated them by teratological observations. In France, besides important
papers of Turpin, Geoffroy de Saint Hilaire, Brongniart, Kirschleger and
others, to which frequent allusion is made in the following pages, there is
disregard, or at best the contemptuous indifference, of the majority of
botanists. De Candolle gave a special impetus to morphology in general by
giving in his adhesion to the morphological hypotheses of Goethe. These
were no mere figments of the poet's imagination, as they were to a large
extent based on the actual investigation of normal and abnormal
organisation by Goethe both alone, and also in conjunction with Batsch and
Jaeger.
De Candolle's example was contagious. Scarcely a botanist of any eminence
since his time but has contributed his quota to the records of vegetable
teratology, in proof of which the names of Humboldt, Robert Brown, the De
Jussieus, the Saint Hilaires, of Moquin-Tandon, of Lindley, and many
others, not to mention botanists still living, may be cited. To students and
amateurs the subject seems always to have presented special attractions,
probably from the singularity of the appearances presented, and from the
fact that in many cases the examination of individual instances of
malformation can be carried on, to a large extent, without the lengthened or
continuous investigation and critical comparative study required by other
departments of botanical science. Be this as it may, teratology owes a very
large number of its records to this class of observers.
While the number of scattered papers on vegetable teratology in various
European languages is so great as to preclude the possibility of collating
them all, there is no general treatise on the subject in the English language,
with the exception of Hopkirk's 'Flora Anomala,' a book now rarely met
with, and withal very imperfect; and this notwithstanding that Robert
Brown early lent his sanction to the doctrines of Goethe, and himself
illustrated them by teratological observations. In France, besides important
papers of Turpin, Geoffroy de Saint Hilaire, Brongniart, Kirschleger and
others, to which frequent allusion is made in the following pages, there is
Page 25
the classic work of Moquin-Tandon, which was translated into German by
Schauer. Germany has also given us the monographs of Batsch, Jæger,
Rœper, Engelmann, Schimper, Braun, Fleischer, Wigand, and many others.
Switzerland has furnished the treatises of the De Candolles, and of Cramer;
Belgium, those of Morren, &c., all of which, as well as many others that
might be mentioned, are, with the exception of Moquin-Tandon's
'Eléments,' to be considered as referring to limited portions only and not to
the whole subject.[1]
In the compilation of the present volume great use has been made of the
facts recorded in the works just cited, and especially in those of Moquin-
Tandon, Engelmann, and Morren. A very large number of communications
on teratological subjects in the various European scientific publications
have also been laid under contribution. In most cases reference has been
given to, and due acknowledgment made of, the sources whence
information has been gathered. Should any such reference be omitted, the
neglect must be attributed to inadvertence, not to design. In selecting
illustrations from the immense number of recorded facts, the principle
followed has been to choose those which seemed either intrinsically the
most important, or those which are recorded with the most care. In addition
to these public sources of information, the author has availed himself of
every opportunity that has offered itself of examining cases of unusual
conformation in plants. For many such opportunities the author has to thank
his friends and correspondents. Nor has he less reason to be grateful for the
suggestions that they have made, and the information they have supplied. In
particular the writer is desirous of acknowledging his obligations to the
Society, under whose auspices this work is published, and to Mr. S. J.
Salter, to whom the book in some degree owes its origin.
The drawings, where not otherwise stated, have been executed either from
the author's own rough sketches, or from the actual specimens, by Mr. E. M.
Schauer. Germany has also given us the monographs of Batsch, Jæger,
Rœper, Engelmann, Schimper, Braun, Fleischer, Wigand, and many others.
Switzerland has furnished the treatises of the De Candolles, and of Cramer;
Belgium, those of Morren, &c., all of which, as well as many others that
might be mentioned, are, with the exception of Moquin-Tandon's
'Eléments,' to be considered as referring to limited portions only and not to
the whole subject.[1]
In the compilation of the present volume great use has been made of the
facts recorded in the works just cited, and especially in those of Moquin-
Tandon, Engelmann, and Morren. A very large number of communications
on teratological subjects in the various European scientific publications
have also been laid under contribution. In most cases reference has been
given to, and due acknowledgment made of, the sources whence
information has been gathered. Should any such reference be omitted, the
neglect must be attributed to inadvertence, not to design. In selecting
illustrations from the immense number of recorded facts, the principle
followed has been to choose those which seemed either intrinsically the
most important, or those which are recorded with the most care. In addition
to these public sources of information, the author has availed himself of
every opportunity that has offered itself of examining cases of unusual
conformation in plants. For many such opportunities the author has to thank
his friends and correspondents. Nor has he less reason to be grateful for the
suggestions that they have made, and the information they have supplied. In
particular the writer is desirous of acknowledging his obligations to the
Society, under whose auspices this work is published, and to Mr. S. J.
Salter, to whom the book in some degree owes its origin.
The drawings, where not otherwise stated, have been executed either from
the author's own rough sketches, or from the actual specimens, by Mr. E. M.
Page 26
Williams. A large number of woodcuts have also been kindly placed at the
disposal of the author by the proprietors of the 'Gardeners' Chronicle.'[2]
As it is impossible to frame any but a purely arbitrary definition of
teratology or to trace the limits between variation and malformation, it may
suffice to say that vegetable teratology comprises the history of the
irregularities of growth and development in plants, and of the causes
producing them. These irregularities differ from variations mainly in their
wider deviation from the customary structure, in their more frequent and
more obvious dependence on external causes rather than on inherent
tendency, in their more sudden appearance, and lastly in their smaller
liability to be transmitted by inheritance.
What may be termed normal morphology includes the study of the form,
arrangement, size and other characteristic attributes of the several parts of
plants, their internal structure, and the precise relation one form bears to
another. In order the more thoroughly to investigate these matters it is
necessary to consider the mode of growth, and specially the plan of
evolution or development of each organ. This is the more needful owing to
the common origin of things ultimately very different one from the other,
and to the presence of organs which, in the adult state, are identical or
nearly so in aspect, but which nevertheless are very unlike in the early
stages of their existence.[3] Following Goethe, these changes in the course
of development are sometimes called metamorphoses. In this way Agardh[4]
admits three kinds of metamorphosis, which he characterises as: 1st.
Successive metamorphoses, or those changes in the course of evolution
which each individual organ undergoes in its passage from the embryonic to
the adult condition, or from the simple and incomplete to the complex and
perfect. 2. Ascending metamorphoses, including those changes of form
manifested in the same adult organism by the several parts of which it
disposal of the author by the proprietors of the 'Gardeners' Chronicle.'[2]
As it is impossible to frame any but a purely arbitrary definition of
teratology or to trace the limits between variation and malformation, it may
suffice to say that vegetable teratology comprises the history of the
irregularities of growth and development in plants, and of the causes
producing them. These irregularities differ from variations mainly in their
wider deviation from the customary structure, in their more frequent and
more obvious dependence on external causes rather than on inherent
tendency, in their more sudden appearance, and lastly in their smaller
liability to be transmitted by inheritance.
What may be termed normal morphology includes the study of the form,
arrangement, size and other characteristic attributes of the several parts of
plants, their internal structure, and the precise relation one form bears to
another. In order the more thoroughly to investigate these matters it is
necessary to consider the mode of growth, and specially the plan of
evolution or development of each organ. This is the more needful owing to
the common origin of things ultimately very different one from the other,
and to the presence of organs which, in the adult state, are identical or
nearly so in aspect, but which nevertheless are very unlike in the early
stages of their existence.[3] Following Goethe, these changes in the course
of development are sometimes called metamorphoses. In this way Agardh[4]
admits three kinds of metamorphosis, which he characterises as: 1st.
Successive metamorphoses, or those changes in the course of evolution
which each individual organ undergoes in its passage from the embryonic to
the adult condition, or from the simple and incomplete to the complex and
perfect. 2. Ascending metamorphoses, including those changes of form
manifested in the same adult organism by the several parts of which it
Page 27
consists—those parts being typically identical or homologous, such as the
parts of the flower, or, in animals, the vertebræ, &c. 3. Collateral
metamorphoses, comprising those permutations of form and function
manifested in homologous organs in the different groups of organisms,
classes, orders, genera, species, &c.
Thus, in the first instance, we have a comparative examination of the form
of each or any separate part of the same individual at different epochs in its
life-history; in the second we have a similar comparison instituted between
the several parts of the same organism which originally were identical in
appearance, but which have in course of evolution altered in character. In
the third form we have the comparative view not of one organ at different
times, nor of the several parts of one organism, but of the constituent
elements pertaining to those aggregates of individuals to which naturalists
apply the terms classes, orders, &c.
In successive metamorphosis we have a measure of the amount of change
and of the perfection of structure to which each separate organ attains.
In ascending metamorphosis we have a gauge of the extent of alteration that
may take place in the several homologous organs under existing
circumstances.
In collateral metamorphosis, in the same way, we have an illustration of the
degree of change possible in aggregates of organisms under existing
circumstances.
Now it is clear that from an investigation of all three classes just mentioned,
we shall be able to gain an idea of those points which are common to all
parts, to all individuals or to all aggregates, and those that are peculiar to
some of them, and, by eliminating the one from the other, we shall arrive at
conclusions which will be more or less generally accurate or applicable,
parts of the flower, or, in animals, the vertebræ, &c. 3. Collateral
metamorphoses, comprising those permutations of form and function
manifested in homologous organs in the different groups of organisms,
classes, orders, genera, species, &c.
Thus, in the first instance, we have a comparative examination of the form
of each or any separate part of the same individual at different epochs in its
life-history; in the second we have a similar comparison instituted between
the several parts of the same organism which originally were identical in
appearance, but which have in course of evolution altered in character. In
the third form we have the comparative view not of one organ at different
times, nor of the several parts of one organism, but of the constituent
elements pertaining to those aggregates of individuals to which naturalists
apply the terms classes, orders, &c.
In successive metamorphosis we have a measure of the amount of change
and of the perfection of structure to which each separate organ attains.
In ascending metamorphosis we have a gauge of the extent of alteration that
may take place in the several homologous organs under existing
circumstances.
In collateral metamorphosis, in the same way, we have an illustration of the
degree of change possible in aggregates of organisms under existing
circumstances.
Now it is clear that from an investigation of all three classes just mentioned,
we shall be able to gain an idea of those points which are common to all
parts, to all individuals or to all aggregates, and those that are peculiar to
some of them, and, by eliminating the one from the other, we shall arrive at
conclusions which will be more or less generally accurate or applicable,
Page 28
according to the ability of the student and the extent to which the
comparative analysis is earned. It is thus that morphologists have been
enabled to frame types or standards of reference, and systematists to
collocate the organisms they deal with into groups. These standards and
groups are more or less artificial (none can be entirely natural) in proportion
to the amount of knowledge possessed by their framers, and the use they
make of it.
From this point of view teratological metamorphosis of all three kinds
demands as much attention as that which is called normal. We can have no
thorough knowledge of an organ, of an individual which is an aggregate of
organs, or of an aggregate of individuals of whatever degree, unless we
know approximately, at least, what are the limits of each. It is not possible
to trace these limits accurately in the case of natural science, but the larger
our knowledge and the wider our generalisations, the closer will be our
approach to the truth.
The most satisfactory classification of malformations would be one founded
upon the nature of the causes inducing the several changes. Thus, in all
organised beings, there is a process of growth, mere increase in bulk as it
were, and a process of evolution or metamorphosis, in accordance with
which certain parts assume a different form from the rest, in order the better
to fit them for the performance of different offices. Should growth and
development be uniform and regular, that is in accordance with what is
habitual in any particular species, there is no monstrosity, but if either
growth or development be in any way irregular, malformation results.
Hence, theoretically, the best way of grouping cases of malformation would
be according as they are the consequences of:—1st. Arrest of Growth;
2ndly. Excessive Growth; 3rdly. Arrest of Development; 4thly, of Excessive
or Irregular Development.
comparative analysis is earned. It is thus that morphologists have been
enabled to frame types or standards of reference, and systematists to
collocate the organisms they deal with into groups. These standards and
groups are more or less artificial (none can be entirely natural) in proportion
to the amount of knowledge possessed by their framers, and the use they
make of it.
From this point of view teratological metamorphosis of all three kinds
demands as much attention as that which is called normal. We can have no
thorough knowledge of an organ, of an individual which is an aggregate of
organs, or of an aggregate of individuals of whatever degree, unless we
know approximately, at least, what are the limits of each. It is not possible
to trace these limits accurately in the case of natural science, but the larger
our knowledge and the wider our generalisations, the closer will be our
approach to the truth.
The most satisfactory classification of malformations would be one founded
upon the nature of the causes inducing the several changes. Thus, in all
organised beings, there is a process of growth, mere increase in bulk as it
were, and a process of evolution or metamorphosis, in accordance with
which certain parts assume a different form from the rest, in order the better
to fit them for the performance of different offices. Should growth and
development be uniform and regular, that is in accordance with what is
habitual in any particular species, there is no monstrosity, but if either
growth or development be in any way irregular, malformation results.
Hence, theoretically, the best way of grouping cases of malformation would
be according as they are the consequences of:—1st. Arrest of Growth;
2ndly. Excessive Growth; 3rdly. Arrest of Development; 4thly, of Excessive
or Irregular Development.
Page 29
In practice, however, there are so many objections to this plan that it has not
been found practicable to carry it out. The inability arises to a great extent
from our ignorance of what should be attributed to arrest of growth, what to
excess of development, and so on. Moreover, a student with a malformed
plant before him must necessarily ascertain in what way it is malformed
before he can understand how it became so, and for this purpose any
scheme that will enable him readily to detect the kind of monstrosity he is
examining, even though it be confessedly artificial and imperfect will be
better than a more philosophical arrangement which circumstances prevent
him from employing.
The plan followed in this volume is a slight modification of that adopted by
Moquin-Tandon, and with several additions. In it the aim is to place before
the student certain salient and easily recognisable points by reference to
which the desired information can readily be found. Under each subdivision
will be found general explanatory remarks, illustrative details, and usually a
summary of the more important facts and the inferences to be derived from
them. Bibliographical references and lists of the plants most frequently
affected with particular malformations are also given. In reference to both
these points it must be remembered that absolute completeness is not aimed
at; had such fullness of detail been possible of attainment it would have
necessitated for its publication a much larger volume than the present.[5] It
is hoped that both the lists of books and of plants are sufficiently full for all
general purposes.[6]
In the enumeration of plants affected with various malformations the !
denotes that the writer has himself seen examples of the deviation in
question in the particular plant named, while the prefix of the * indicates
that the malformation occurs with special frequency in the particular plant
to which the sign is attached.
been found practicable to carry it out. The inability arises to a great extent
from our ignorance of what should be attributed to arrest of growth, what to
excess of development, and so on. Moreover, a student with a malformed
plant before him must necessarily ascertain in what way it is malformed
before he can understand how it became so, and for this purpose any
scheme that will enable him readily to detect the kind of monstrosity he is
examining, even though it be confessedly artificial and imperfect will be
better than a more philosophical arrangement which circumstances prevent
him from employing.
The plan followed in this volume is a slight modification of that adopted by
Moquin-Tandon, and with several additions. In it the aim is to place before
the student certain salient and easily recognisable points by reference to
which the desired information can readily be found. Under each subdivision
will be found general explanatory remarks, illustrative details, and usually a
summary of the more important facts and the inferences to be derived from
them. Bibliographical references and lists of the plants most frequently
affected with particular malformations are also given. In reference to both
these points it must be remembered that absolute completeness is not aimed
at; had such fullness of detail been possible of attainment it would have
necessitated for its publication a much larger volume than the present.[5] It
is hoped that both the lists of books and of plants are sufficiently full for all
general purposes.[6]
In the enumeration of plants affected with various malformations the !
denotes that the writer has himself seen examples of the deviation in
question in the particular plant named, while the prefix of the * indicates
that the malformation occurs with special frequency in the particular plant
to which the sign is attached.
Page 30
Teratological alterations are rarely isolated phenomena, far more generally
they are associated with other and often compensatory changes. Hence it is
often necessary, in studying any given malformation, to refer to two or
more subdivisions, and in this way a certain amount of repetition becomes
unavoidable. The details of the several cases of malformation given in these
pages are generally arranged according to their apparent degree of
importance. Thus, in a case of prolification associated with multiplication of
the petals, the former change is a greater deviation from the customary form
than the latter, hence reference should be made, in the first instance, to the
sections treating on prolification, and afterwards to those on multiplication.
To facilitate such research, numerous cross references are supplied.
In the investigation of teratological phenomena constant reference must be
made to the normal condition, and vice versâ, else neither the one nor the
other can be thoroughly understood. It cannot, however, be overlooked that
the form and arrangement called normal are often merely those which are
the most common, while the abnormal or unusual arrangement is often
more in consonance with that considered to be typical than the ordinary
one. Thus, too, it is often found that the structural arrangements, which in
one flower are normal, are in another abnormal, in so far that they are not
usual in that particular instance.
For purposes of reference, a standard of comparison is required; and this
standard, so long as its nature is not overlooked, may, indeed must be, to
some extent, an arbitrary one. Thus in the phanerogamous plants there is
assumed to exist, in all cases, an axis (stem, branches, roots, thalamus, &c.),
bearing leaves and flowers. These latter consist of four whorls, calyx,
corolla, stamens, and pistils, each whorl consisting of so many separate
pieces in determinate position and numbers, and of regular proportionate
size. A very close approach to such a flower occurs normally in Limnanthes
and Crassula, and, indeed, in a large proportion of all flowers in an early
they are associated with other and often compensatory changes. Hence it is
often necessary, in studying any given malformation, to refer to two or
more subdivisions, and in this way a certain amount of repetition becomes
unavoidable. The details of the several cases of malformation given in these
pages are generally arranged according to their apparent degree of
importance. Thus, in a case of prolification associated with multiplication of
the petals, the former change is a greater deviation from the customary form
than the latter, hence reference should be made, in the first instance, to the
sections treating on prolification, and afterwards to those on multiplication.
To facilitate such research, numerous cross references are supplied.
In the investigation of teratological phenomena constant reference must be
made to the normal condition, and vice versâ, else neither the one nor the
other can be thoroughly understood. It cannot, however, be overlooked that
the form and arrangement called normal are often merely those which are
the most common, while the abnormal or unusual arrangement is often
more in consonance with that considered to be typical than the ordinary
one. Thus, too, it is often found that the structural arrangements, which in
one flower are normal, are in another abnormal, in so far that they are not
usual in that particular instance.
For purposes of reference, a standard of comparison is required; and this
standard, so long as its nature is not overlooked, may, indeed must be, to
some extent, an arbitrary one. Thus in the phanerogamous plants there is
assumed to exist, in all cases, an axis (stem, branches, roots, thalamus, &c.),
bearing leaves and flowers. These latter consist of four whorls, calyx,
corolla, stamens, and pistils, each whorl consisting of so many separate
pieces in determinate position and numbers, and of regular proportionate
size. A very close approach to such a flower occurs normally in Limnanthes
and Crassula, and, indeed, in a large proportion of all flowers in an early
Page 31
stage of development. To a standard type, such as just mentioned, all the
varied forms that are met with, either in normal or abnormal morphology,
may be referred by bearing in mind the different modifications and
adaptations that the organs have to undergo in the course of their
development. Some parts after a time may cease to grow, others may grow
in an inordinate degree, and so on; and thus, great as may be the ultimate
divergences from the assumed standard, they may all readily be explained
by the operation, simply or conjointly, of some of the four principal causes
of malformation before alluded to. The fact that so many and such varied
changes can thus readily be explained is not only a matter of convenience,
but may be taken as evidence that the standard of reference is not wholly
arbitrary and artificial, but that it is a close approximation to the truth.
It has already been said that an arrangement like that here considered as
typical is natural to some flowers in their adult state, and to a vast number
in their immature condition. It would be no extravagant hypothesis to
surmise that this was the primitive structure of the flower in the higher
plants. Variations from it may have arisen in course of time, owing to the
action of an inherent tendency to vary, or from external circumstances and
varied requirements which may have induced corresponding adaptations,
and which may have been transmitted in accordance with the principle of
hereditary transmission. This hypothesis necessarily implies a prior
simplicity of organisation, of which, indeed, there is sufficient proof; many
cases of malformation can thus be considered as so many reversions to the
ancestral form.
Thus, teratology often serves as an aid in the study of morphology in
general, and also in that of special groups of plants, and hence may even be
of assistance in the determination of affinities. In any case the data supplied
by teratology require to be used with caution and in conjunction with those
derived from the study of development and from analogy. It is even possible
varied forms that are met with, either in normal or abnormal morphology,
may be referred by bearing in mind the different modifications and
adaptations that the organs have to undergo in the course of their
development. Some parts after a time may cease to grow, others may grow
in an inordinate degree, and so on; and thus, great as may be the ultimate
divergences from the assumed standard, they may all readily be explained
by the operation, simply or conjointly, of some of the four principal causes
of malformation before alluded to. The fact that so many and such varied
changes can thus readily be explained is not only a matter of convenience,
but may be taken as evidence that the standard of reference is not wholly
arbitrary and artificial, but that it is a close approximation to the truth.
It has already been said that an arrangement like that here considered as
typical is natural to some flowers in their adult state, and to a vast number
in their immature condition. It would be no extravagant hypothesis to
surmise that this was the primitive structure of the flower in the higher
plants. Variations from it may have arisen in course of time, owing to the
action of an inherent tendency to vary, or from external circumstances and
varied requirements which may have induced corresponding adaptations,
and which may have been transmitted in accordance with the principle of
hereditary transmission. This hypothesis necessarily implies a prior
simplicity of organisation, of which, indeed, there is sufficient proof; many
cases of malformation can thus be considered as so many reversions to the
ancestral form.
Thus, teratology often serves as an aid in the study of morphology in
general, and also in that of special groups of plants, and hence may even be
of assistance in the determination of affinities. In any case the data supplied
by teratology require to be used with caution and in conjunction with those
derived from the study of development and from analogy. It is even possible
Page 32
that some malformations, especially when they acquire a permanent nature
and become capable of reproducing themselves by seed, may be the
starting-point of new species, as they assuredly are of new races, and
between a race and a species he would be a bold man who would undertake
to draw a hard and fast line.[7]
Discredit has been cast on teratology because it has been incautiously used.
At one time it was made to prove almost everything; what wonder that by
some, now-a-days, it is held to prove nothing. True the evidence it affords is
sometimes negative, often conflicting, but it is so rather from imperfect
interpretation than from any intrinsic worthlessness. If misused the fault lies
with the disciple, not with Nature.
Teratology as a guide to the solution of morphological problems has been
especially disparaged in contrast with organogeny, but unfairly so. There is
no reason to exalt or to disparage either at the expense of the other. Both
should receive the attention they demand. The study of development shows
the primitive condition and gradual evolution of parts in any given
individual or species; it carries us back some stages further in the history of
particular organisms, but so also does teratology. Many cases of arrest of
development show the mode of growth and evolution more distinctly, and
with much greater ease to the observer, than does the investigation of the
evolution of organs under natural circumstances. Organogeny by no means
necessarily, or always, gives us an insight into the principles regulating the
construction of flowers in general. It gives us no archetype except in those
comparatively rare cases where primordial symmetry and regularity exist.
When an explanation of the irregularity of development in these early
stages of the plant's history is required, recourse must be had to the
inferences and deductions drawn from teratological investigations and from
the comparative study of allied forms precisely as in the case of adult
flowers.
and become capable of reproducing themselves by seed, may be the
starting-point of new species, as they assuredly are of new races, and
between a race and a species he would be a bold man who would undertake
to draw a hard and fast line.[7]
Discredit has been cast on teratology because it has been incautiously used.
At one time it was made to prove almost everything; what wonder that by
some, now-a-days, it is held to prove nothing. True the evidence it affords is
sometimes negative, often conflicting, but it is so rather from imperfect
interpretation than from any intrinsic worthlessness. If misused the fault lies
with the disciple, not with Nature.
Teratology as a guide to the solution of morphological problems has been
especially disparaged in contrast with organogeny, but unfairly so. There is
no reason to exalt or to disparage either at the expense of the other. Both
should receive the attention they demand. The study of development shows
the primitive condition and gradual evolution of parts in any given
individual or species; it carries us back some stages further in the history of
particular organisms, but so also does teratology. Many cases of arrest of
development show the mode of growth and evolution more distinctly, and
with much greater ease to the observer, than does the investigation of the
evolution of organs under natural circumstances. Organogeny by no means
necessarily, or always, gives us an insight into the principles regulating the
construction of flowers in general. It gives us no archetype except in those
comparatively rare cases where primordial symmetry and regularity exist.
When an explanation of the irregularity of development in these early
stages of the plant's history is required, recourse must be had to the
inferences and deductions drawn from teratological investigations and from
the comparative study of allied forms precisely as in the case of adult
flowers.
Page 33
The study of development is of the highest importance in the examination
of plants as individuals, but in regard to comparative anatomy and
morphology, and specially in its relation to the study of vegetable homology
it has no superiority over teratology. Those who hold the contrary opinion
do so, apparently, because they overlook the fact that there is no distinction,
save of degree, to be drawn between the laws regulating normal
organisation, and those by which so-called abnormal formations are
regulated.
It is sometimes said, and not wholly without truth, that teratology, as it
stands at present, is little more than a record of facts, but in proportion as
the laws that regulate normal growth are better understood, so will the
knowledge of those that govern the so-called monstrous formations
increase. Sufficient has been already said to prove that there is no intrinsic
difference between the laws of growth in the two cases. As our knowledge
increases we shall be enabled to ascertain approximately of what extent of
variation a given form is capable, under given conditions, and to refer all
formations now considered anomalous to a few well-defined forms. Already
teratology has done much towards showing the erroneous nature of many
morphological statements that still pass current in our text-books, though
their fallacy has been demonstrated again and again. Thus organs are said to
be fused which were never separate, disjunctions and separations are
assigned to parts that were never joined, adhesions and cohesions are
spoken of in cases where, from the nature of things, neither adhesion nor
cohesion could have existed. Some organs are said to be atrophied which
were never larger and more fully developed than they now are, and so on.
So long as these expressions are used in a merely conventional sense and
for purposes of artificial classification or convenience, well and good, but
let us not delude ourselves that we are thus contributing to the philosophical
study either of the conformation of plants or of the affinities existing
between them. What hope is there that we shall ever gain clear conceptions
of plants as individuals, but in regard to comparative anatomy and
morphology, and specially in its relation to the study of vegetable homology
it has no superiority over teratology. Those who hold the contrary opinion
do so, apparently, because they overlook the fact that there is no distinction,
save of degree, to be drawn between the laws regulating normal
organisation, and those by which so-called abnormal formations are
regulated.
It is sometimes said, and not wholly without truth, that teratology, as it
stands at present, is little more than a record of facts, but in proportion as
the laws that regulate normal growth are better understood, so will the
knowledge of those that govern the so-called monstrous formations
increase. Sufficient has been already said to prove that there is no intrinsic
difference between the laws of growth in the two cases. As our knowledge
increases we shall be enabled to ascertain approximately of what extent of
variation a given form is capable, under given conditions, and to refer all
formations now considered anomalous to a few well-defined forms. Already
teratology has done much towards showing the erroneous nature of many
morphological statements that still pass current in our text-books, though
their fallacy has been demonstrated again and again. Thus organs are said to
be fused which were never separate, disjunctions and separations are
assigned to parts that were never joined, adhesions and cohesions are
spoken of in cases where, from the nature of things, neither adhesion nor
cohesion could have existed. Some organs are said to be atrophied which
were never larger and more fully developed than they now are, and so on.
So long as these expressions are used in a merely conventional sense and
for purposes of artificial classification or convenience, well and good, but
let us not delude ourselves that we are thus contributing to the philosophical
study either of the conformation of plants or of the affinities existing
between them. What hope is there that we shall ever gain clear conceptions
Page 34
as to the former, as long as we tie ourselves down to formulas which are the
expressions of facts as they appear to be, rather than as they really are?
What chance is there of our attaining to comprehensive and accurate views
of the genealogy and affinities of plants as long as we are restricted by false
notions as to the conformation and mutual relation of their parts?[8]
That teratology may serve the purposes of systematic botany to a greater
extent than might at first be supposed becomes obvious from a
consideration of such facts as are mentioned under the head of Peloria,
while the presence of rudimentary organs, or the occasional appearance of
additional parts, or other changes, may, and often do, afford a clue to the
relationship existing between plants—a relationship that might otherwise be
unsuspected. So, too, some of the alterations met with appear susceptible of
no other explanations than that they are reversions to some pre-existing
form, or, at any rate, that they are manifestations of a phase of the plant
affected different from that which is habitual, and due, as it were, to a sort
of allotropism.
The mutations and perversions of form, associated as they commonly are
with corresponding changes of function, show the connection between
teratology and physiology—a connection which is seen to be the more
intimate when viewed in the light afforded by the writings and experiments
of Gærtner, Sprengel, and St. Hilaire, and, in our own times, especially by
the writings and experiments of Mr. Darwin, whose works on the 'Origin of
Species,' and particularly on the 'Variation of Animals and Plants under
Domestication' comprise so large a collection of facts for the use of students
in most departments of biology. It will suffice to allude, in support of these
statements, to the writings of Mr. Darwin on such subjects as rudimentary
organs, the use or disuse of certain parts according to circumstances, the
frequently observed tendency of some flowers to become structurally
unisexual, the liability of other flowers perfectly organised to become
expressions of facts as they appear to be, rather than as they really are?
What chance is there of our attaining to comprehensive and accurate views
of the genealogy and affinities of plants as long as we are restricted by false
notions as to the conformation and mutual relation of their parts?[8]
That teratology may serve the purposes of systematic botany to a greater
extent than might at first be supposed becomes obvious from a
consideration of such facts as are mentioned under the head of Peloria,
while the presence of rudimentary organs, or the occasional appearance of
additional parts, or other changes, may, and often do, afford a clue to the
relationship existing between plants—a relationship that might otherwise be
unsuspected. So, too, some of the alterations met with appear susceptible of
no other explanations than that they are reversions to some pre-existing
form, or, at any rate, that they are manifestations of a phase of the plant
affected different from that which is habitual, and due, as it were, to a sort
of allotropism.
The mutations and perversions of form, associated as they commonly are
with corresponding changes of function, show the connection between
teratology and physiology—a connection which is seen to be the more
intimate when viewed in the light afforded by the writings and experiments
of Gærtner, Sprengel, and St. Hilaire, and, in our own times, especially by
the writings and experiments of Mr. Darwin, whose works on the 'Origin of
Species,' and particularly on the 'Variation of Animals and Plants under
Domestication' comprise so large a collection of facts for the use of students
in most departments of biology. It will suffice to allude, in support of these
statements, to the writings of Mr. Darwin on such subjects as rudimentary
organs, the use or disuse of certain parts according to circumstances, the
frequently observed tendency of some flowers to become structurally
unisexual, the liability of other flowers perfectly organised to become
Page 35
functionally imperfect, at least so far as any reciprocal action of the organs
of the same flower is concerned, reversions, classification, general
morphology, and other subjects handled at once with such comprehensive
breadth and minute accuracy of detail by our great physiologist.
In the following pages alterations of function, unless attended by
corresponding alterations of form, are either only incidentally alluded to, or
are wholly passed over; such, for instance, as alterations in the period of
flowering, in the duration of the several organs, and so forth.[9] Pathological
changes, lesions caused by insect puncture or other causes, also find no
place in this book, unless the changes are of such a character as to admit of
definite comparison with normal conformation. Usually such changes are
entirely heteromorphous, and, as it were, foreign to the natural organisation.
The practical applications of teratology deserve the attention of those
cultivators who are concerned in the embellishment of our gardens and the
supply of our tables. The florist lays down a certain arbitrary standard of
perfection, and attempts to make flowers conform to that model. Whether it
be in good taste or not to value all flowers, in proportion as they accord
with an artificial and comparatively inelastic standard of this kind, we need
not stop to enquire; suffice it to say, that taking the matter in its broadest
sense, the aim of the florist is to produce large, symmetrical flowers,
brightly and purely coloured, or if parti-coloured, the colours must be
distinct, harmonious, or contrasted. When all this is done, the flower, in
most instances, becomes 'monstrous' of the eyes in the botanist, though all
the more interesting to the student of morphology on that account. In like
manner the double flowers, the "breaks," the "sports" which the florist
cultivates so anxiously, are all of them greater or less deviations from the
ordinary form, while the broccolies, the cabbages, and many other products
of our kitchen gardens and fields owe the estimation in which they are held
entirely to those peculiarities which, by an unhappy application of words,
of the same flower is concerned, reversions, classification, general
morphology, and other subjects handled at once with such comprehensive
breadth and minute accuracy of detail by our great physiologist.
In the following pages alterations of function, unless attended by
corresponding alterations of form, are either only incidentally alluded to, or
are wholly passed over; such, for instance, as alterations in the period of
flowering, in the duration of the several organs, and so forth.[9] Pathological
changes, lesions caused by insect puncture or other causes, also find no
place in this book, unless the changes are of such a character as to admit of
definite comparison with normal conformation. Usually such changes are
entirely heteromorphous, and, as it were, foreign to the natural organisation.
The practical applications of teratology deserve the attention of those
cultivators who are concerned in the embellishment of our gardens and the
supply of our tables. The florist lays down a certain arbitrary standard of
perfection, and attempts to make flowers conform to that model. Whether it
be in good taste or not to value all flowers, in proportion as they accord
with an artificial and comparatively inelastic standard of this kind, we need
not stop to enquire; suffice it to say, that taking the matter in its broadest
sense, the aim of the florist is to produce large, symmetrical flowers,
brightly and purely coloured, or if parti-coloured, the colours must be
distinct, harmonious, or contrasted. When all this is done, the flower, in
most instances, becomes 'monstrous' of the eyes in the botanist, though all
the more interesting to the student of morphology on that account. In like
manner the double flowers, the "breaks," the "sports" which the florist
cultivates so anxiously, are all of them greater or less deviations from the
ordinary form, while the broccolies, the cabbages, and many other products
of our kitchen gardens and fields owe the estimation in which they are held
entirely to those peculiarities which, by an unhappy application of words,
Page 36
are called monstrous by botanists. Grafting, layering, the "striking" of
cuttings, the formation of adventitious roots and buds, processes on which
the cultivator so greatly relies for the propagation and extension of his
plants, are also matters with which teratology concerns itself. Again the
difficulty experienced occasionally in getting vines, strawberries, &c., to set
properly, may sometimes be accounted for by that inherent tendency which
some plants possess of exchanging an hermaphrodite for a unisexual
condition.
For reasons then of direct practical utility, no less than on purely scientific
grounds, it is desirable to study these irregularities of growth, their nature,
limits, and inducing causes; and to this end it is hoped the present work
may, in some degree, contribute.
FOOTNOTES:
[1] An excellent summary of the history of Vegetable Teratology is given in Kirschleger's
'Essai historique de la Tératologie Végétale,' Strasburg, 1845.
[2] In some instances diagrams and formulæ are given in explanation of the conformation
of monstrous flowers; in general these require no further explanation than is given in the text,
unless it be to state that the horizontal line—is intended to indicate the cohesion of the parts
over which it is placed, while the vertical line | signifies the adhesion of the organs by whose
side it is placed. The formula
S S S S S
------------------------
| P P P P P
|
| ST ST ST ST ST
shows that the sepals (S) are distinct, the petals (P) coherent, and the stamens (ST) adherent to
the petals.
[3] Wolff was the first to call attention to the great importance of the study of
development. He was followed by Turpin, Mirbel, Schleiden, Payer, and others, and its value
is now fully recognised by botanists.
[4] Agardh, "Theoria Syst. Plant.," p. xxiii.
cuttings, the formation of adventitious roots and buds, processes on which
the cultivator so greatly relies for the propagation and extension of his
plants, are also matters with which teratology concerns itself. Again the
difficulty experienced occasionally in getting vines, strawberries, &c., to set
properly, may sometimes be accounted for by that inherent tendency which
some plants possess of exchanging an hermaphrodite for a unisexual
condition.
For reasons then of direct practical utility, no less than on purely scientific
grounds, it is desirable to study these irregularities of growth, their nature,
limits, and inducing causes; and to this end it is hoped the present work
may, in some degree, contribute.
FOOTNOTES:
[1] An excellent summary of the history of Vegetable Teratology is given in Kirschleger's
'Essai historique de la Tératologie Végétale,' Strasburg, 1845.
[2] In some instances diagrams and formulæ are given in explanation of the conformation
of monstrous flowers; in general these require no further explanation than is given in the text,
unless it be to state that the horizontal line—is intended to indicate the cohesion of the parts
over which it is placed, while the vertical line | signifies the adhesion of the organs by whose
side it is placed. The formula
S S S S S
------------------------
| P P P P P
|
| ST ST ST ST ST
shows that the sepals (S) are distinct, the petals (P) coherent, and the stamens (ST) adherent to
the petals.
[3] Wolff was the first to call attention to the great importance of the study of
development. He was followed by Turpin, Mirbel, Schleiden, Payer, and others, and its value
is now fully recognised by botanists.
[4] Agardh, "Theoria Syst. Plant.," p. xxiii.
Page 37
[5] In the memoirs of Hopkirk, Kirschleger, Cramer, Hallier, and others, malformations are
arranged primarily according to the organs affected, an arrangement which has only
convenience to justify it. It is hoped that the index and the headings to the paragraphs in the
present volume will suit the convenience of the reader as well as if the more artificial plan just
alluded to had been adopted.
[6] Cryptogamous plants are only incidentally alluded to in these pages, owing to their
wide difference in structure from flowering plants. Attention may, also, here be called to a
paper of M. de Seynes in a recent number of the Bulletin of the Botanical Society of France,
vol. xiv, p. 290, tab. 5 et 6, in which numerous cases of malformation among agarics are
recorded. See also same publication, vol. iv, p. 744; vol. v, p. 211; vol. vi, p. 496.
[7] On this subject see a paper of M. Naudin in the 'Comptes Rendus,' 1867, t. 64, pp.
929–933.
[8] It is probable that many terms and expressions calculated to mislead in the way above
mentioned are made use of in the following pages. The inconsistency manifested by their use
may be excused on the ground of ignorance of the true structure, and by the circumstance that
in many cases facts alone are recorded without an explanation of them being offered.
Moreover, it is desirable to act in conformity with the usual practice of botanical writers, and
not to change established terminology, even if suspected to convey false ideas, until the true
condition of affairs be thoroughly well ascertained by organogenetic research or other means.
[9] A curious illustration of the latter class of alterations came under the writer's notice last
summer (1868), and which he has reason to believe has not been previously recorded, viz. the
persistence in an unwithered state of the petals at the base of the ripe fruit, in a strawberry. All
the fruits on the particular plants alluded to were thus provided as it were with a white frill.
Whether this be a constant occurrence in the particular variety is not known.
arranged primarily according to the organs affected, an arrangement which has only
convenience to justify it. It is hoped that the index and the headings to the paragraphs in the
present volume will suit the convenience of the reader as well as if the more artificial plan just
alluded to had been adopted.
[6] Cryptogamous plants are only incidentally alluded to in these pages, owing to their
wide difference in structure from flowering plants. Attention may, also, here be called to a
paper of M. de Seynes in a recent number of the Bulletin of the Botanical Society of France,
vol. xiv, p. 290, tab. 5 et 6, in which numerous cases of malformation among agarics are
recorded. See also same publication, vol. iv, p. 744; vol. v, p. 211; vol. vi, p. 496.
[7] On this subject see a paper of M. Naudin in the 'Comptes Rendus,' 1867, t. 64, pp.
929–933.
[8] It is probable that many terms and expressions calculated to mislead in the way above
mentioned are made use of in the following pages. The inconsistency manifested by their use
may be excused on the ground of ignorance of the true structure, and by the circumstance that
in many cases facts alone are recorded without an explanation of them being offered.
Moreover, it is desirable to act in conformity with the usual practice of botanical writers, and
not to change established terminology, even if suspected to convey false ideas, until the true
condition of affairs be thoroughly well ascertained by organogenetic research or other means.
[9] A curious illustration of the latter class of alterations came under the writer's notice last
summer (1868), and which he has reason to believe has not been previously recorded, viz. the
persistence in an unwithered state of the petals at the base of the ripe fruit, in a strawberry. All
the fruits on the particular plants alluded to were thus provided as it were with a white frill.
Whether this be a constant occurrence in the particular variety is not known.
Page 38
VEGETABLE TERATOLOGY.
Page 39
BOOK I.
DEVIATIONS FROM THE ORDINARY
ARRANGEMENT OF ORGANS.
As full details relating to the disposition or arrangement of the general
organs of flowering plants are given in all the ordinary text-books, it is only
necessary in this place to allude to the main facts at present known, and
which serve as the standard of comparison with which all morphological
changes are compared.
Even in the case of the roots, which appear to be very irregular in their
ramification, it has been found that, in the first instance at least, the rootlets
or fibrils are arranged in regular order one over another, in a certain
determinate number of vertical ranks, generally either in two or in four,
sometimes in three or in five series. This regularity of arrangement
(Rhizotaxy), first carefully studied by M. Clos, is connected with the
disposition of the fibro-vascular bundles in the body of the root. This
primitive regularity is soon lost as the plant grows.
In the case of the leaves there are two principal modes of arrangement,
dependent, as it would seem, on their simultaneous or on their successive
development; thus, if two leaves on opposite sides of the stem are
developed at the same time, we have the arrangement called opposite; if
DEVIATIONS FROM THE ORDINARY
ARRANGEMENT OF ORGANS.
As full details relating to the disposition or arrangement of the general
organs of flowering plants are given in all the ordinary text-books, it is only
necessary in this place to allude to the main facts at present known, and
which serve as the standard of comparison with which all morphological
changes are compared.
Even in the case of the roots, which appear to be very irregular in their
ramification, it has been found that, in the first instance at least, the rootlets
or fibrils are arranged in regular order one over another, in a certain
determinate number of vertical ranks, generally either in two or in four,
sometimes in three or in five series. This regularity of arrangement
(Rhizotaxy), first carefully studied by M. Clos, is connected with the
disposition of the fibro-vascular bundles in the body of the root. This
primitive regularity is soon lost as the plant grows.
In the case of the leaves there are two principal modes of arrangement,
dependent, as it would seem, on their simultaneous or on their successive
development; thus, if two leaves on opposite sides of the stem are
developed at the same time, we have the arrangement called opposite; if
Page 40
there are more than two, the disposition is then called verticillate or
whorled. On the other hand, if the leaves are developed in succession, one
after the other, they are found to emerge from the stem in a spiral direction.
In either case the leaves are arranged in a certain regular manner, according
to what are called the laws of Phyllotaxis, which need not be entered into
fully here; but in order the better to estimate the teratological changes
which take place, it may be well to allude to the following circumstances
relating to the alternation of parts. The effect of this alternation is such, that
no two adjacent leaves stand directly over or in front one of the other, but a
little to one side or a little higher up. Now, in the alternate arrangement the
successive leaves of each spiral cycle alternate one with another till the coil
is completed. For the sake of clearness this may be illustrated thus:—
Suppose the spiral cycle to comprise five leaves, numbered 1, 2, 3, 4, 5,
then 2 would intervene between 1 and 3, and so on, while the sixth leaf
would be the commencement of a new series, and would be placed exactly
over 1. This arrangement may be thus formularised:
6 7 8 9 10
1 2 3 4 5
In the verticillate or simultaneous arrangement of leaves the case is
somewhat different. Let us suppose a whorl of eight leaves, surmounted by
a similar whorl of eight. In such a case it will generally be found that the
whorls alternate one with another, as may be represented by this symbol:
9 10 11 12 13 14 15 16
1 2 3 4 5 6 7 8
The simplest illustration of this arrangement is seen in the case of decussate
leaves, where those organs are placed in pairs, and the pairs cross one
another at right angles. This may be expressed by the following symbol:
whorled. On the other hand, if the leaves are developed in succession, one
after the other, they are found to emerge from the stem in a spiral direction.
In either case the leaves are arranged in a certain regular manner, according
to what are called the laws of Phyllotaxis, which need not be entered into
fully here; but in order the better to estimate the teratological changes
which take place, it may be well to allude to the following circumstances
relating to the alternation of parts. The effect of this alternation is such, that
no two adjacent leaves stand directly over or in front one of the other, but a
little to one side or a little higher up. Now, in the alternate arrangement the
successive leaves of each spiral cycle alternate one with another till the coil
is completed. For the sake of clearness this may be illustrated thus:—
Suppose the spiral cycle to comprise five leaves, numbered 1, 2, 3, 4, 5,
then 2 would intervene between 1 and 3, and so on, while the sixth leaf
would be the commencement of a new series, and would be placed exactly
over 1. This arrangement may be thus formularised:
6 7 8 9 10
1 2 3 4 5
In the verticillate or simultaneous arrangement of leaves the case is
somewhat different. Let us suppose a whorl of eight leaves, surmounted by
a similar whorl of eight. In such a case it will generally be found that the
whorls alternate one with another, as may be represented by this symbol:
9 10 11 12 13 14 15 16
1 2 3 4 5 6 7 8
The simplest illustration of this arrangement is seen in the case of decussate
leaves, where those organs are placed in pairs, and the pairs cross one
another at right angles. This may be expressed by the following symbol:
Page 41
7 8
5 6
3 4
1 2
Thus, while in both the annular and the spiral modes of development the
individual members of each complete series necessarily alternate one with
another, in the former case the series themselves alternate, while in the
successive arrangement they are placed directly one over the other. There
are, of course, exceptions, but the rule is as has been stated, and the effect is
to prevent one leaf from interfering with the development and growth of its
neighbours.
In the case of the whorled or simultaneous arrangement the conditions of
growth must be uniform on all sides, but in the successive or spiral
disposition the conditions influencing growth act with unequal force, on
different sides of the stem, at the same time. In the whorl there is an
illustration of radiating symmetry, while in the spiral arrangement there is a
transition to the bilateral symmetry. There are frequent passages from one to
the other even under normal circumstances; thus, while the one arrangement
obtains in the ordinary leaves, the parts of the flower may be disposed
according to the other method. In the annular disposition it generally
happens that the rings are separated one from the other by the development
of the stem between them, the internodes between the constituent leaves
themselves of course being undeveloped; on the other hand, in the spiral or
successive arrangement there is no such alternate growth and arrest of
growth of the stem between the leaves, or between successive cycles, but
the growth is, under favorable conditions, continuous—leaf is separated
from leaf, and cycle from cycle, by the continually elongating stem. Thus,
the two modes of growth correspond precisely with those observed in the
case of definite and indefinite inflorescence respectively.
5 6
3 4
1 2
Thus, while in both the annular and the spiral modes of development the
individual members of each complete series necessarily alternate one with
another, in the former case the series themselves alternate, while in the
successive arrangement they are placed directly one over the other. There
are, of course, exceptions, but the rule is as has been stated, and the effect is
to prevent one leaf from interfering with the development and growth of its
neighbours.
In the case of the whorled or simultaneous arrangement the conditions of
growth must be uniform on all sides, but in the successive or spiral
disposition the conditions influencing growth act with unequal force, on
different sides of the stem, at the same time. In the whorl there is an
illustration of radiating symmetry, while in the spiral arrangement there is a
transition to the bilateral symmetry. There are frequent passages from one to
the other even under normal circumstances; thus, while the one arrangement
obtains in the ordinary leaves, the parts of the flower may be disposed
according to the other method. In the annular disposition it generally
happens that the rings are separated one from the other by the development
of the stem between them, the internodes between the constituent leaves
themselves of course being undeveloped; on the other hand, in the spiral or
successive arrangement there is no such alternate growth and arrest of
growth of the stem between the leaves, or between successive cycles, but
the growth is, under favorable conditions, continuous—leaf is separated
from leaf, and cycle from cycle, by the continually elongating stem. Thus,
the two modes of growth correspond precisely with those observed in the
case of definite and indefinite inflorescence respectively.
Page 42
Fig. 1.—Diagram showing the arrangement of parts in a complete, regular, pentamerous flower: s,
sepals; p, petals; st, stamens; o, ovaries.
The same arrangements, that are observed in the disposition of the leaves,
apply equally well to the several parts of the flower; thus, in what is for
convenience considered the typical flower, there is a calyx of five or more
distinct sepals, equal in size, and arranged in a whorl, a corolla of a similar
number of petals alternating with the sepals, five stamens placed in the
same position with reference to the petals, and five carpels alternating with
the stamens. Throughout this book this arrangement is taken as the standard
of reference. Nevertheless the spiral order does occur in the floral leaves as
well as in those of the stem; it often happens, especially when the organs
are numerous, that they form spiral series; and the same holds good very
generally, when the parts of the flower are uneven in number, as in the very
common quincuncial arrangement of the sepals, &c.
To these general remarks, intended to show the agreement between the
disposition of the leaves of the stem and those of the flower, it is merely
necessary to add that the arrangement of the placentas, as well as that of the
ovules borne on them, is also definite, and takes place according to methods
explained in all the text-books, and on which, therefore, it is not necessary
to dilate in this place.
The branches of the stem or axis correspond for the most part in disposition
with that of the leaves from the axils of which they originate, subject,
however, to numerous disturbing causes, and to alterations from the usual
sepals; p, petals; st, stamens; o, ovaries.
The same arrangements, that are observed in the disposition of the leaves,
apply equally well to the several parts of the flower; thus, in what is for
convenience considered the typical flower, there is a calyx of five or more
distinct sepals, equal in size, and arranged in a whorl, a corolla of a similar
number of petals alternating with the sepals, five stamens placed in the
same position with reference to the petals, and five carpels alternating with
the stamens. Throughout this book this arrangement is taken as the standard
of reference. Nevertheless the spiral order does occur in the floral leaves as
well as in those of the stem; it often happens, especially when the organs
are numerous, that they form spiral series; and the same holds good very
generally, when the parts of the flower are uneven in number, as in the very
common quincuncial arrangement of the sepals, &c.
To these general remarks, intended to show the agreement between the
disposition of the leaves of the stem and those of the flower, it is merely
necessary to add that the arrangement of the placentas, as well as that of the
ovules borne on them, is also definite, and takes place according to methods
explained in all the text-books, and on which, therefore, it is not necessary
to dilate in this place.
The branches of the stem or axis correspond for the most part in disposition
with that of the leaves from the axils of which they originate, subject,
however, to numerous disturbing causes, and to alterations from the usual
Page 43
or typical order brought about by the development of buds. These latter
organs, as it seems, may be found in almost any situation, though their
ordinary position is in the axil of a leaf or at the end of a stem or branch.
The points just mentioned are of primary importance in structural botany,
and as such are seized on not only by the morphologist, but by the
systematic botanist, who finds in them the characters by which he may
separate one group from another. Thanks to the labours of those observers
who have devoted their attention to that difficult but most important branch
of study, organogeny, or the investigation of the development of the various
organs, and to the researches of the students of comparative anatomy or
morphology, the main principles regulating the arrangement and form of the
organs of flowering plants seem to be fairly well established, though in
matters of detail much remains to be cleared up, even in such important
points as the share which the axis takes in the construction of the flower and
fruit, the nature of the placenta, the construction of the ovules, and other
points.
The facts already known justify the adoption of a standard or typical
arrangement as just mentioned. The intrinsic value of this type is shown by
the facility with which all varieties of form or arrangement may be
explained by reference to certain modifications of it. It must, however, be
considered as an abstraction, and should be looked on in the light rather of a
scaffolding, which enables us to see the building and its several parts, than
of the edifice itself, but which latter, from our imperfect knowledge and
limited powers, we could not see without some such assistance.
The typical form may be, hypothetically at least, considered as the primitive
one transmitted by hereditary descent from generation to generation, and
modified to suit the requirements of the individual, or in accordance with
circumstances. If it be borne in mind that it is but an artificial contrivance,
organs, as it seems, may be found in almost any situation, though their
ordinary position is in the axil of a leaf or at the end of a stem or branch.
The points just mentioned are of primary importance in structural botany,
and as such are seized on not only by the morphologist, but by the
systematic botanist, who finds in them the characters by which he may
separate one group from another. Thanks to the labours of those observers
who have devoted their attention to that difficult but most important branch
of study, organogeny, or the investigation of the development of the various
organs, and to the researches of the students of comparative anatomy or
morphology, the main principles regulating the arrangement and form of the
organs of flowering plants seem to be fairly well established, though in
matters of detail much remains to be cleared up, even in such important
points as the share which the axis takes in the construction of the flower and
fruit, the nature of the placenta, the construction of the ovules, and other
points.
The facts already known justify the adoption of a standard or typical
arrangement as just mentioned. The intrinsic value of this type is shown by
the facility with which all varieties of form or arrangement may be
explained by reference to certain modifications of it. It must, however, be
considered as an abstraction, and should be looked on in the light rather of a
scaffolding, which enables us to see the building and its several parts, than
of the edifice itself, but which latter, from our imperfect knowledge and
limited powers, we could not see without some such assistance.
The typical form may be, hypothetically at least, considered as the primitive
one transmitted by hereditary descent from generation to generation, and
modified to suit the requirements of the individual, or in accordance with
circumstances. If it be borne in mind that it is but an artificial contrivance,
Page 44
more or less true—a means to an end, and not the end itself—no harm will
arise from its employment; and as knowledge increases, or as circumstances
demand, the hypothetical type can be replaced by another more in
accordance with the actual state of science.
Teratological changes in the arrangement of organs depend upon arrest of
growth, as when parts usually spirally arranged remain verticillate, owing to
the non-development of the internodes, or to excessive growth, or
development; but in many instances it is impossible, without studying the
development of the malformed flower, to ascertain whether the altered
arrangement is due to an excessive or to a diminished action. Practically,
however, it is of comparatively little importance to know whether, say, the
isolation of parts, that are usually combined together, is congenital (i.e. the
result of an arrest of growth preventing their union), or whether it be due to
a separation of parts primitively undivided; the effect remains the same,
though the cause may have been very different.
The principal alterations to be mentioned under this head may therefore be
conveniently arranged under the following categories:—Union,
Independence, Displacement, Prolification, Heterotaxy, and Heterogamy.
PART I.
UNION OF ORGANS.
The union of parts, usually separate in their adult condition, is of very
common occurrence as a malformation. The instances of its manifestation
admit of being grouped under the heads of Cohesion, where parts of the
same whorl, or of the same organ, are united together; and of Adhesion,
arise from its employment; and as knowledge increases, or as circumstances
demand, the hypothetical type can be replaced by another more in
accordance with the actual state of science.
Teratological changes in the arrangement of organs depend upon arrest of
growth, as when parts usually spirally arranged remain verticillate, owing to
the non-development of the internodes, or to excessive growth, or
development; but in many instances it is impossible, without studying the
development of the malformed flower, to ascertain whether the altered
arrangement is due to an excessive or to a diminished action. Practically,
however, it is of comparatively little importance to know whether, say, the
isolation of parts, that are usually combined together, is congenital (i.e. the
result of an arrest of growth preventing their union), or whether it be due to
a separation of parts primitively undivided; the effect remains the same,
though the cause may have been very different.
The principal alterations to be mentioned under this head may therefore be
conveniently arranged under the following categories:—Union,
Independence, Displacement, Prolification, Heterotaxy, and Heterogamy.
PART I.
UNION OF ORGANS.
The union of parts, usually separate in their adult condition, is of very
common occurrence as a malformation. The instances of its manifestation
admit of being grouped under the heads of Cohesion, where parts of the
same whorl, or of the same organ, are united together; and of Adhesion,
Page 45
where the union takes place between members of different whorls, or
between two or more ordinarily wholly detached and distinct parts. In either
case, the apparent union may be congenital (that is, the result of a primitive
integrity or a lack of separation), or it may really consist in a coalition of
parts originally distinct and separate. In practice it is not always easy to
distinguish between these two different conditions. Indeed, in most cases it
cannot be done without tracing the development of the flower throughout
all its stages. It is needless to make more than a passing allusion to the
frequency with which both congenital integrity or subsequent coalescence
of organs exist under ordinary circumstances. Considered as a teratological
phenomenon, union admits of being grouped into several subdivisions, such
as Cohesion, Adhesion, Synanthy, Syncarpy, Synophty, &c. Each of these
subdivisions will be separately treated, but it maybe here said that, in all or
any case, the degree of fusion may be very slight, or it may be so perfect
that there may be a complete amalgamation of two or more parts, while to
all outward appearance the organ may be single. The column of Orchids
may be referred to as an illustration under natural circumstances of the
complete union of many usually distinct parts.
In the uncertainty that exists in many cases as to the real nature of the
occurrence, it would be idle to attempt to explain the causes of fusions. It is
clear, however, that an arrest of development will tend towards the
maintenance of primordial integrity (congenital fusion), and that pressure
will induce the coalition of organs primarily distinct.
CHAPTER I.
COHESION.
Following Augustin Pyranius De Candolle, botanists have applied the term
cohesion to the coalescence of parts of the same organ or of members of the
between two or more ordinarily wholly detached and distinct parts. In either
case, the apparent union may be congenital (that is, the result of a primitive
integrity or a lack of separation), or it may really consist in a coalition of
parts originally distinct and separate. In practice it is not always easy to
distinguish between these two different conditions. Indeed, in most cases it
cannot be done without tracing the development of the flower throughout
all its stages. It is needless to make more than a passing allusion to the
frequency with which both congenital integrity or subsequent coalescence
of organs exist under ordinary circumstances. Considered as a teratological
phenomenon, union admits of being grouped into several subdivisions, such
as Cohesion, Adhesion, Synanthy, Syncarpy, Synophty, &c. Each of these
subdivisions will be separately treated, but it maybe here said that, in all or
any case, the degree of fusion may be very slight, or it may be so perfect
that there may be a complete amalgamation of two or more parts, while to
all outward appearance the organ may be single. The column of Orchids
may be referred to as an illustration under natural circumstances of the
complete union of many usually distinct parts.
In the uncertainty that exists in many cases as to the real nature of the
occurrence, it would be idle to attempt to explain the causes of fusions. It is
clear, however, that an arrest of development will tend towards the
maintenance of primordial integrity (congenital fusion), and that pressure
will induce the coalition of organs primarily distinct.
CHAPTER I.
COHESION.
Following Augustin Pyranius De Candolle, botanists have applied the term
cohesion to the coalescence of parts of the same organ or of members of the
Page 46
same whorl; for instance, to the union of the sepals in a gamosepalous
calyx, or of the petals in a gamopetalous corolla. It may arise either from a
union between organs originally distinct, or more frequently from a want of
separation between parts, which under general circumstances become
divided during their development. Nothing is more common as a normal
occurrence, while viewed as a teratological phenomenon it is also very
frequent. For the purposes of convenience it admits of subdivision into
those cases wherein the union takes place between the branches of the same
plant, or between the margins of the same leaf-organ, or between those of
different members of the same whorl.
Cohesion between the axes of the same plant.—This cohesion may occur
in various manners. Firstly. The branches of the main stem may become
united one to the other. Secondly. Two or more stems become joined
together. Thirdly. The branches become united to the stem; or, lastly, the
roots may become fused one with another.
calyx, or of the petals in a gamopetalous corolla. It may arise either from a
union between organs originally distinct, or more frequently from a want of
separation between parts, which under general circumstances become
divided during their development. Nothing is more common as a normal
occurrence, while viewed as a teratological phenomenon it is also very
frequent. For the purposes of convenience it admits of subdivision into
those cases wherein the union takes place between the branches of the same
plant, or between the margins of the same leaf-organ, or between those of
different members of the same whorl.
Cohesion between the axes of the same plant.—This cohesion may occur
in various manners. Firstly. The branches of the main stem may become
united one to the other. Secondly. Two or more stems become joined
together. Thirdly. The branches become united to the stem; or, lastly, the
roots may become fused one with another.
Page 47
Fig. 2—Cohesion of two branches in Dipsacus sylvestris.
The first of these is most commonly met with, doubtless owing to the
number of the branches and the facilities for their union. An illustration of it
is afforded by the figure (fig. 2), showing cohesion affecting the branches of
a teazle (Dipsacus sylvestris). Union of the branches may be the result of an
original cohesion of the buds, while in other cases the fusion does not take
place until after development has proceeded to some extent. Of this latter
kind illustrations are common where the branches are in close
approximation; if the bark be removed by friction the two surfaces are very
likely to become united (natural grafting). Such a union of the branches is
very common in the ivy, the elder, the beech, and other plants. It may take
place in various directions, lengthwise, obliquely, or transversely, according
to circumstances. This mode of union belongs, perhaps, rather to the
domain of pathology than of teratology. Some of the instances that have
been recorded of very large trees, such as the chestnut of Mount Ætna, are
The first of these is most commonly met with, doubtless owing to the
number of the branches and the facilities for their union. An illustration of it
is afforded by the figure (fig. 2), showing cohesion affecting the branches of
a teazle (Dipsacus sylvestris). Union of the branches may be the result of an
original cohesion of the buds, while in other cases the fusion does not take
place until after development has proceeded to some extent. Of this latter
kind illustrations are common where the branches are in close
approximation; if the bark be removed by friction the two surfaces are very
likely to become united (natural grafting). Such a union of the branches is
very common in the ivy, the elder, the beech, and other plants. It may take
place in various directions, lengthwise, obliquely, or transversely, according
to circumstances. This mode of union belongs, perhaps, rather to the
domain of pathology than of teratology. Some of the instances that have
been recorded of very large trees, such as the chestnut of Mount Ætna, are
Page 48
really cases where fusion has taken place between several of the branches,
or suckers, thrown out from the same original stem.[10] The same process of
grafting occurs sometimes in the roots, as in Taxus baccata mentioned by
Moquin, and also in the aerial roots of many of the tropical climbing plants,
such as Clusia rosea, &c.
Fig. 3.—Fasciation in Lettuce.
Fasciation.—In the preceding instances of union between the branches,
&c., the actual number of the fused parts is not increased; but if it happen
that an unusual number of buds be formed in close apposition, so that they
are liable to be compressed during their growth, union is very likely to take
place, the more so from the softness of the young tissues. In this way it is
probable that what is termed fasciation is brought about. This is one of the
most common of all malformations, and seems to affect certain plants more
frequently than others. In its simplest form it consists of a flat, ribbon-like
or suckers, thrown out from the same original stem.[10] The same process of
grafting occurs sometimes in the roots, as in Taxus baccata mentioned by
Moquin, and also in the aerial roots of many of the tropical climbing plants,
such as Clusia rosea, &c.
Fig. 3.—Fasciation in Lettuce.
Fasciation.—In the preceding instances of union between the branches,
&c., the actual number of the fused parts is not increased; but if it happen
that an unusual number of buds be formed in close apposition, so that they
are liable to be compressed during their growth, union is very likely to take
place, the more so from the softness of the young tissues. In this way it is
probable that what is termed fasciation is brought about. This is one of the
most common of all malformations, and seems to affect certain plants more
frequently than others. In its simplest form it consists of a flat, ribbon-like
Page 49
expansion of the stem or branch; cylindrical below, the branches gradually
lose their pristine form, and assume the flattened condition.
Fig. 4.—Fasciation in Asparagus.
lose their pristine form, and assume the flattened condition.
Fig. 4.—Fasciation in Asparagus.
Page 50
Fig. 5.—Fasciated branch of Pinus Pinaster.
Very generally the surface is striated by the prominence of the woody fibres
which, running parallel for a time, converge or diverge at the summit
according to the shape of the branch. If the rate of growth be equal, or
nearly so, on both sides, the stem retains its straight direction, but it more
generally happens that the growth on one side is more rapid and more
vigorous than on the other, and hence arises that curvature of the fasciated
branch so commonly met with, e.g. in the ash (Fraxinus), wherein it has
been likened to a shepherd's crook. It is probable that almost any plant may
present this change. It occurs alike in herbaceous and in woody plants,
originating in the latter case while the branches are still soft. It may be
remarked that, in the case of herbaceous plants, the fasciation always affects
the principal stem, while, on the other hand, in the case of trees and shrubs
the deformity occurs most frequently in the branches; thus, while in the
former it may be said that the whole of the stem is more or less affected, in
Very generally the surface is striated by the prominence of the woody fibres
which, running parallel for a time, converge or diverge at the summit
according to the shape of the branch. If the rate of growth be equal, or
nearly so, on both sides, the stem retains its straight direction, but it more
generally happens that the growth on one side is more rapid and more
vigorous than on the other, and hence arises that curvature of the fasciated
branch so commonly met with, e.g. in the ash (Fraxinus), wherein it has
been likened to a shepherd's crook. It is probable that almost any plant may
present this change. It occurs alike in herbaceous and in woody plants,
originating in the latter case while the branches are still soft. It may be
remarked that, in the case of herbaceous plants, the fasciation always affects
the principal stem, while, on the other hand, in the case of trees and shrubs
the deformity occurs most frequently in the branches; thus, while in the
former it may be said that the whole of the stem is more or less affected, in
Page 51
the latter it is rare to see more than one or two branches of the same tree
thus deformed. It is a common thing for the fasciated branch to divide at the
summit into a number of subdivisions. These latter may be deformed like
the parent branch, or they may resume the ordinary aspect of the twigs.
Fig. 6.—Fasciation and spiral torsion in the stem of Asparagus.
Sometimes the flattened stem is destitute of buds, at other times, these
organs are scattered irregularly over its surface or are crowded together in a
sort of crest along the apex. When, as often happens, the deformity is
accompanied with a twisting of the branch spirally, the buds may be placed
irregularly, or in other cases along the free edge of the spiral curve. In a
specimen of Bupleurum falcatum mentioned by Moquin the spiral
thus deformed. It is a common thing for the fasciated branch to divide at the
summit into a number of subdivisions. These latter may be deformed like
the parent branch, or they may resume the ordinary aspect of the twigs.
Fig. 6.—Fasciation and spiral torsion in the stem of Asparagus.
Sometimes the flattened stem is destitute of buds, at other times, these
organs are scattered irregularly over its surface or are crowded together in a
sort of crest along the apex. When, as often happens, the deformity is
accompanied with a twisting of the branch spirally, the buds may be placed
irregularly, or in other cases along the free edge of the spiral curve. In a
specimen of Bupleurum falcatum mentioned by Moquin the spiral
Page 52
arrangement of the leaves was replaced by a series of perfect whorls, each
consisting of five, six, seven, or eight segments, and there was a flower-
stalk in the axil of each leaf.
When flowers are borne on these fasciated stems they are generally altered
in structure; sometimes the thalamus itself becomes more or less fasciated
or flattened, and the different organs of the flower are arranged on an
elliptical axis. A case of this nature is described by Schlechtendal ('Bot.
Zeit.,' 1857, p. 880), in Cytisus nigricans, and M. Moquin-Tandon describes
an instance in the vine in one flower of which sepals, petals, stamens, and
ovary were abortive, while the receptacle was hypertrophied and fasciated,
and bore on its surface a few adventitious buds.[11] The pedicels of
Streptocarpus Rexii have also been observed in a fasciated state.[12]
It has been occasionally observed that the fasciated condition is hereditary;
thus, Moquin relates that some seeds of a fasciated Cirsium reproduced the
same condition in the seedlings,[13] while a similar tendency is inherited in
the case of the cockscomb (Celosia).
With reference to the nature of the deformity in question there is a
difference of opinion; while most authors consider it to be due to the causes
before mentioned, Moquin was of opinion that fasciation was due to a
flattening of a single stem or branch. Linnæus, on the other hand,
considered such stems to be the result of the formation of an unusual
number of buds, the shoots resulting from which became coherent as
growth proceeded:—"Fasciata dici solet planta cum plures caules
connascuntur, ut unus ex plurimis instar fasciæ evadat et compressus"
(Linn., 'Phil. Bot.,' 274). A similar opinion was held by J. D. Major in a
singular book entitled 'De Plantâ, Monstrosa, Gottorpiensi,' Schleswig,
1665, wherein the stem of a Chrysanthemum is depicted in the fasciated
condition.
consisting of five, six, seven, or eight segments, and there was a flower-
stalk in the axil of each leaf.
When flowers are borne on these fasciated stems they are generally altered
in structure; sometimes the thalamus itself becomes more or less fasciated
or flattened, and the different organs of the flower are arranged on an
elliptical axis. A case of this nature is described by Schlechtendal ('Bot.
Zeit.,' 1857, p. 880), in Cytisus nigricans, and M. Moquin-Tandon describes
an instance in the vine in one flower of which sepals, petals, stamens, and
ovary were abortive, while the receptacle was hypertrophied and fasciated,
and bore on its surface a few adventitious buds.[11] The pedicels of
Streptocarpus Rexii have also been observed in a fasciated state.[12]
It has been occasionally observed that the fasciated condition is hereditary;
thus, Moquin relates that some seeds of a fasciated Cirsium reproduced the
same condition in the seedlings,[13] while a similar tendency is inherited in
the case of the cockscomb (Celosia).
With reference to the nature of the deformity in question there is a
difference of opinion; while most authors consider it to be due to the causes
before mentioned, Moquin was of opinion that fasciation was due to a
flattening of a single stem or branch. Linnæus, on the other hand,
considered such stems to be the result of the formation of an unusual
number of buds, the shoots resulting from which became coherent as
growth proceeded:—"Fasciata dici solet planta cum plures caules
connascuntur, ut unus ex plurimis instar fasciæ evadat et compressus"
(Linn., 'Phil. Bot.,' 274). A similar opinion was held by J. D. Major in a
singular book entitled 'De Plantâ, Monstrosa, Gottorpiensi,' Schleswig,
1665, wherein the stem of a Chrysanthemum is depicted in the fasciated
condition.
Page 53
Fig. 7.—Fasciation in the scape of the Dandelion (Leontodon Taraxacum).
The striæ, which these stems almost invariably present, exhibit the lines of
junction, and the spiral or other curvatures and contraction, which are so
often met with, may be accounted for by the unequal growth of one portion
of the stem as contrasted with that of another. Against this view Moquin
cites the instances of one-stemmed plants, such as Androsace maxima, but,
on the other hand, those herbaceous plants having usually but a single stem
not unfrequently produce several which may remain distinct, but not
uncommonly become united together. Prof. Hincks[14] cites cases of this
kind in Primula vulgaris, Hieracium aureum, and Ranunculus bulbosus. I
have myself met with several cases of the kind in Primula veris, in the
Polyanthus, in the Daisy, and in the Leontodon Taraxacum, in which latter a
The striæ, which these stems almost invariably present, exhibit the lines of
junction, and the spiral or other curvatures and contraction, which are so
often met with, may be accounted for by the unequal growth of one portion
of the stem as contrasted with that of another. Against this view Moquin
cites the instances of one-stemmed plants, such as Androsace maxima, but,
on the other hand, those herbaceous plants having usually but a single stem
not unfrequently produce several which may remain distinct, but not
uncommonly become united together. Prof. Hincks[14] cites cases of this
kind in Primula vulgaris, Hieracium aureum, and Ranunculus bulbosus. I
have myself met with several cases of the kind in Primula veris, in the
Polyanthus, in the Daisy, and in the Leontodon Taraxacum, in which latter a
Page 54
fusion of two or more flower-stems bearing at the top a composite flower,
and made up of two, three, four, or more flowers combined together, and
containing all the organs that would be present in the same flowers if
separate, is very common.
Moquin's second objection is founded upon the fact that, in certain fasciated
stems, the branches are not increased in number or altered in arrangement
from what is usual; but however true this may be in particular cases, it is
quite certain that in the majority of instances a large increase in the number
of leaves and buds is a prominent characteristic of fasciated stems.
Another argument used by the distinguished French botanist to show that
fasciated stems are not due to cohesion of two or more stems, is founded on
the fact that a transverse section of a fasciated stem generally shows an
elliptical outline with but a single central canal. On the other hand, if two
branches become united and a transverse section be made, the form of the
cut surface would be more or less like that of the figure 8[symbol: 8 turned
90°], although in old stems this may give place to an elliptical outline, but
even then traces of two medullary canals may be found. This argument is
very deceptive, for the appearance of the transverse section must depend,
not only on the intimacy of their union, but also on the internal structure of
the stems themselves. When two flowers cohere without much pressure
they exhibit uniting circles somewhat resembling the figure of 8[symbol: 8
turned 90°], but when more completely combined they have an outline of a
very elongated figure, and something similar is to be expected in
herbaceous stems. Even the elongated pith of a transversely cut, woody,
fasciated stem only marks the intimate union of several branches, and Prof.
Hincks, whose views the writer entirely shares, has noticed instances of the
union of two, and of only two, stems where the internal appearance was the
same as in other fasciations.
and made up of two, three, four, or more flowers combined together, and
containing all the organs that would be present in the same flowers if
separate, is very common.
Moquin's second objection is founded upon the fact that, in certain fasciated
stems, the branches are not increased in number or altered in arrangement
from what is usual; but however true this may be in particular cases, it is
quite certain that in the majority of instances a large increase in the number
of leaves and buds is a prominent characteristic of fasciated stems.
Another argument used by the distinguished French botanist to show that
fasciated stems are not due to cohesion of two or more stems, is founded on
the fact that a transverse section of a fasciated stem generally shows an
elliptical outline with but a single central canal. On the other hand, if two
branches become united and a transverse section be made, the form of the
cut surface would be more or less like that of the figure 8[symbol: 8 turned
90°], although in old stems this may give place to an elliptical outline, but
even then traces of two medullary canals may be found. This argument is
very deceptive, for the appearance of the transverse section must depend,
not only on the intimacy of their union, but also on the internal structure of
the stems themselves. When two flowers cohere without much pressure
they exhibit uniting circles somewhat resembling the figure of 8[symbol: 8
turned 90°], but when more completely combined they have an outline of a
very elongated figure, and something similar is to be expected in
herbaceous stems. Even the elongated pith of a transversely cut, woody,
fasciated stem only marks the intimate union of several branches, and Prof.
Hincks, whose views the writer entirely shares, has noticed instances of the
union of two, and of only two, stems where the internal appearance was the
same as in other fasciations.
Page 55
Moquin, moreover, raises the objection that it is unlikely that several
branches should become united lengthwise in one plane only, and, further,
that in the greater number of fasciations all the other branches which should
be present are to be found—not one is wanting, not one has disappeared, as
might have been anticipated had fusion taken place. In raising this
objection, Moquin seems not sufficiently to have considered the
circumstance that the buds in these cases are in one plane from the first, and
are all about equal in point of age and size.
The last objection that Moquin raises to the opinion that fasciation is the
result of a grafting process is, that in such a case, examples should be found
wherein the branches are incompletely fused, and where on a transverse
section traces of the medullary canals belonging to each branch should be
visible. The arrangement of leaves or buds on the surface should also in
such a case indicate a fusion of several spiral cycles or whorls. To this it
may be replied that such cases are met with very frequently indeed. A figure
is given by De Candolle[15] of a stem of Spartium junceum having several
branches only imperfectly fasciated.
Fasciated stems, then, seem to be best explained, as is stated by Prof.
Hincks, "on the principle of adhesion arising in cases where from
superabundant nourishment, especially if accompanied by some check or
injury, numerous buds have been produced in close proximity, and the
supposition that these growths are produced by the dilatation of a single
stem is founded on a false analogy between fasciated stems and certain
other anomalous growths."
It will not, of course, be forgotten that this fasciated condition occurs so
frequently in some plants as almost to constitute their natural state, e.g.
Sedum cristatum, Celosia, &c. This condition may be induced by the art of
the gardener—"Fit idem arte, si plures caules enascentes cogantur
branches should become united lengthwise in one plane only, and, further,
that in the greater number of fasciations all the other branches which should
be present are to be found—not one is wanting, not one has disappeared, as
might have been anticipated had fusion taken place. In raising this
objection, Moquin seems not sufficiently to have considered the
circumstance that the buds in these cases are in one plane from the first, and
are all about equal in point of age and size.
The last objection that Moquin raises to the opinion that fasciation is the
result of a grafting process is, that in such a case, examples should be found
wherein the branches are incompletely fused, and where on a transverse
section traces of the medullary canals belonging to each branch should be
visible. The arrangement of leaves or buds on the surface should also in
such a case indicate a fusion of several spiral cycles or whorls. To this it
may be replied that such cases are met with very frequently indeed. A figure
is given by De Candolle[15] of a stem of Spartium junceum having several
branches only imperfectly fasciated.
Fasciated stems, then, seem to be best explained, as is stated by Prof.
Hincks, "on the principle of adhesion arising in cases where from
superabundant nourishment, especially if accompanied by some check or
injury, numerous buds have been produced in close proximity, and the
supposition that these growths are produced by the dilatation of a single
stem is founded on a false analogy between fasciated stems and certain
other anomalous growths."
It will not, of course, be forgotten that this fasciated condition occurs so
frequently in some plants as almost to constitute their natural state, e.g.
Sedum cristatum, Celosia, &c. This condition may be induced by the art of
the gardener—"Fit idem arte, si plures caules enascentes cogantur
Page 56
penetrare coarctatum spatium et parturiri tanquam ex angusto utero, sic
sæpe in Ranunculo, Beta, Asparago, Hesperide Pinu, Celosiâ,
Tragopogone, Scorzonerâ Cotula fœtida," Linnæus op. cit.
Plot, in his 'History of Oxfordshire,' considers fasciation to arise from the
ascent of too much nourishment for one stalk and not enough for two,
"which accident of plants," says Plot, the German virtuosi ('Misc. Curios.
Med. Physic. Acad. Nat. Cur.,' Ann. i, Observ. 102,) "think only to happen
after hard and late winters, by reason whereof, indeed, the sap, being
restrained somewhat longer than ordinary, upon sudden thaws may
probably be sent up more forcibly, and so produce these fasciated stalks,
whereas the natural and graduated ascent would have produced them but
single." Prof. Hincks' explanation is, however, more near to the truth, and
his opinion is borne out by the frequency with which this change is met
with in certain plants which are frequently forced on during their growth, as
lettuce, asparagus, endive, &c., all of which are very subject to this change.
In the 'Transactions of the Horticultural Society of London,' vol. iv, p. 321,
Mr. Knight gives an account of the cultivation of the cockscomb, so as to
ensure the production of the very large flower-stalks for which this plant is
admired. The principal points in the culture were the application of a large
quantity of stimulating manure and the maintenance of a high temperature.
One of them so grown measured eighteen inches in width.
The list which is appended is intended to show those plants in which
fasciation has been most frequently observed. It makes no pretension to be
complete, but is sufficiently so for the purpose indicated: the * denotes the
especial frequency of the change in question; the ! indicates that the writer
has himself seen the plant, so marked, affected in this way. The remainder
have been copied from various sources.
Exogens.
sæpe in Ranunculo, Beta, Asparago, Hesperide Pinu, Celosiâ,
Tragopogone, Scorzonerâ Cotula fœtida," Linnæus op. cit.
Plot, in his 'History of Oxfordshire,' considers fasciation to arise from the
ascent of too much nourishment for one stalk and not enough for two,
"which accident of plants," says Plot, the German virtuosi ('Misc. Curios.
Med. Physic. Acad. Nat. Cur.,' Ann. i, Observ. 102,) "think only to happen
after hard and late winters, by reason whereof, indeed, the sap, being
restrained somewhat longer than ordinary, upon sudden thaws may
probably be sent up more forcibly, and so produce these fasciated stalks,
whereas the natural and graduated ascent would have produced them but
single." Prof. Hincks' explanation is, however, more near to the truth, and
his opinion is borne out by the frequency with which this change is met
with in certain plants which are frequently forced on during their growth, as
lettuce, asparagus, endive, &c., all of which are very subject to this change.
In the 'Transactions of the Horticultural Society of London,' vol. iv, p. 321,
Mr. Knight gives an account of the cultivation of the cockscomb, so as to
ensure the production of the very large flower-stalks for which this plant is
admired. The principal points in the culture were the application of a large
quantity of stimulating manure and the maintenance of a high temperature.
One of them so grown measured eighteen inches in width.
The list which is appended is intended to show those plants in which
fasciation has been most frequently observed. It makes no pretension to be
complete, but is sufficiently so for the purpose indicated: the * denotes the
especial frequency of the change in question; the ! indicates that the writer
has himself seen the plant, so marked, affected in this way. The remainder
have been copied from various sources.
Exogens.
Page 57
α. Herbaceous.
Page 58
Ranunculus tripartitus.
*bulbosus!
Philonotis.
Delphinium elatum.
*sp.!
Hesperis matronalis.
*Cheiranthus Cheiri!
*Matthiola incana!
*Brassica oleracea! var. pl. inflor.
Linum usitatissimum!
Althæa rosea!
Lavatera trimestris.
Geranii sp.
Tropæolum majus!
Viola odorata inflor.!
Reseda odorata!
Fragaria vesca.
Ervum lens.
Trifolium resupinatum.
repens!
pratense!
Saxifraga mutata.
irrigua.
Bupleurum falcatum.
Bunium flexuosum.
*Sedum reflexum!
cristatum!
Epilobium augustifolium!
Momordica Elaterium!
Gaura biennis.
Cotula fœtida.
Barkhausia taraxacifolia.
Carlina vulgaris!
Apargia autumnalis.
*Leontodon Taraxacum inflor.!
Centaurea Scabiosa.
*Cichorium Intybus!
Hieracium Pilosella.
aureum.
*bulbosus!
Philonotis.
Delphinium elatum.
*sp.!
Hesperis matronalis.
*Cheiranthus Cheiri!
*Matthiola incana!
*Brassica oleracea! var. pl. inflor.
Linum usitatissimum!
Althæa rosea!
Lavatera trimestris.
Geranii sp.
Tropæolum majus!
Viola odorata inflor.!
Reseda odorata!
Fragaria vesca.
Ervum lens.
Trifolium resupinatum.
repens!
pratense!
Saxifraga mutata.
irrigua.
Bupleurum falcatum.
Bunium flexuosum.
*Sedum reflexum!
cristatum!
Epilobium augustifolium!
Momordica Elaterium!
Gaura biennis.
Cotula fœtida.
Barkhausia taraxacifolia.
Carlina vulgaris!
Apargia autumnalis.
*Leontodon Taraxacum inflor.!
Centaurea Scabiosa.
*Cichorium Intybus!
Hieracium Pilosella.
aureum.
Page 59
umbellatum.
*Chrysanthemum Leucanthemum.
indicum!
Anthemis nobilis.
arvensis.
Cirsium lanceolatum.
Conyza squarrosa!
Inula dysenterica!
Tragopogon porrifolium.
Cnicus palustris.
Carduus arvensis!
Helianthus tuberosus!
annuus.
Cineraria palustris.
Helianthus sp.!
Dahlia variabilis.
Bellis perennis inflor.!
Coreopsis sp.!
Crepis virens.
Lactuca sativa!
Zinnia elegans.
*Campanula medium!
rapunculoides.
thyrsoidea.
Dipsacus pilosus.
fullonum.
silvestris.
Knautia arvensis.
Phyteuma orbiculare.
Jasione montana.
*Linaria purpurea!
Antirrhinum majus!
Veronica amethystea.
Veronica maritima.
sp.
Russellia juncea!
Digitalis purpurea!
Ajuga pyramidalis.
Hyssopus officinalis.
*Chrysanthemum Leucanthemum.
indicum!
Anthemis nobilis.
arvensis.
Cirsium lanceolatum.
Conyza squarrosa!
Inula dysenterica!
Tragopogon porrifolium.
Cnicus palustris.
Carduus arvensis!
Helianthus tuberosus!
annuus.
Cineraria palustris.
Helianthus sp.!
Dahlia variabilis.
Bellis perennis inflor.!
Coreopsis sp.!
Crepis virens.
Lactuca sativa!
Zinnia elegans.
*Campanula medium!
rapunculoides.
thyrsoidea.
Dipsacus pilosus.
fullonum.
silvestris.
Knautia arvensis.
Phyteuma orbiculare.
Jasione montana.
*Linaria purpurea!
Antirrhinum majus!
Veronica amethystea.
Veronica maritima.
sp.
Russellia juncea!
Digitalis purpurea!
Ajuga pyramidalis.
Hyssopus officinalis.
Page 60
Dracocephalum moldavicum.
Myosotis scorpioides.
Echium pyrenaicum.
simplex.
Stapeliæ sp.
Lysimachia vulgaris!
Androsace maxima.
Primula veris inflor.!
denticulata inflor.!
Polemonium cœruleum.
Convolvulus sepium!
arvensis!
Plantago media.
*Euphorbia Characias.
exigua.
*Cyparissias.
Suæda maritima.
*Celosia sp.
Beta vulgaris inflor.!
Phytolacca sp.
β. Woody.
Berberis vulgaris.
Hibiscus syriacus!
Acer pseudo-platanus!
Dodonæa viscosa.
Sterculia platanifolia.
Euonymus japonicus!
Vitis vinifera inflor.!
Spartium Scoparium!
Spartium junceum!
Cytisus Laburnum.
nigricans.
Chorozema ilicifolium.
Amorpha sp.
Phaseolus sp.
Prunus sylvestris.
Laurocerasus!
Rosa sp.!
Myosotis scorpioides.
Echium pyrenaicum.
simplex.
Stapeliæ sp.
Lysimachia vulgaris!
Androsace maxima.
Primula veris inflor.!
denticulata inflor.!
Polemonium cœruleum.
Convolvulus sepium!
arvensis!
Plantago media.
*Euphorbia Characias.
exigua.
*Cyparissias.
Suæda maritima.
*Celosia sp.
Beta vulgaris inflor.!
Phytolacca sp.
β. Woody.
Berberis vulgaris.
Hibiscus syriacus!
Acer pseudo-platanus!
Dodonæa viscosa.
Sterculia platanifolia.
Euonymus japonicus!
Vitis vinifera inflor.!
Spartium Scoparium!
Spartium junceum!
Cytisus Laburnum.
nigricans.
Chorozema ilicifolium.
Amorpha sp.
Phaseolus sp.
Prunus sylvestris.
Laurocerasus!
Rosa sp.!
Page 61
Spiræa sp.!
Cotoneaster microphylla!
Ailanthus glandulosus.
*Fraxinus Ornus!
*excelsior!
Melia Azedarach.
Xanthoxylum sp.!
Sambucus nigra.!
Aucuba japonica.
Erica sp. cult.
Jasminum nudiflorum!
officinale!
Olea europœa.
Punica Granatum.
Ilex aquifolium!
Daphne indica.
Daphne odora.
Suæda fruticosa.
Ulmus campestris.
Alnus incana.
Salix vitellina, &c.!
Thuja orientalis.
Pinus pinaster!
sylvestris!
Abies excelsa!
Taxus baccata.
Larix europœa.
Endogens.
Lilium Martagon.
candidum!
*Fritillaria imperialis!
Asparagus officinalis!
Hyacinthus orientalis!
Tamus communis!
Narcissi sp.!
Gladiolus sp.
Zea Mays.
Filices.
Cotoneaster microphylla!
Ailanthus glandulosus.
*Fraxinus Ornus!
*excelsior!
Melia Azedarach.
Xanthoxylum sp.!
Sambucus nigra.!
Aucuba japonica.
Erica sp. cult.
Jasminum nudiflorum!
officinale!
Olea europœa.
Punica Granatum.
Ilex aquifolium!
Daphne indica.
Daphne odora.
Suæda fruticosa.
Ulmus campestris.
Alnus incana.
Salix vitellina, &c.!
Thuja orientalis.
Pinus pinaster!
sylvestris!
Abies excelsa!
Taxus baccata.
Larix europœa.
Endogens.
Lilium Martagon.
candidum!
*Fritillaria imperialis!
Asparagus officinalis!
Hyacinthus orientalis!
Tamus communis!
Narcissi sp.!
Gladiolus sp.
Zea Mays.
Filices.
Page 62
See also—Moquin-Tandon, 'Elem. Ter. Veget.,' p. 146; C. O. Weber, 'Verhandl. Nat. Hist.,'
Vereins, f. d. Preuss., Rheinl. und Westphal., 1860, p. 347, tab. vii; Hallier, 'Phytopathol.,'
p. 128; Boehmer, 'De plantis Fasciatis,' Wittenb., 1752.
Cohesion of foliar organs.—This takes place in several ways, and in very
various degrees; the simplest case is that characterised by the cohesion of
the margins of the same organ, as in the condition called perfoliate in
descriptive works, and which is due either to a cohesion of the margins of
the basal lobes of the leaf, or to the development of the leaf in a sheathing
or tubular manner. As an abnormal occurrence, I have met with this
perfoliation in a leaf of Goodenia ovata. The condition in question is often
loosely confounded with connation, or the union of two leaves by their
bases. In other cases the union takes place between the margins of two or
more leaves.
Cohesion of margins of single organs.—The leaves of Hazels may often
be found with their margins coherent at the base, so as to become peltate,
while in other cases, the disc of the leaf is so depressed that a true pitcher is
formed. This happens also in the Lime Tilia, in which genus pitcher- or
hood-like leaves (folia cucullata) may frequently be met with. There are
trees with leaves of this character in the cemetery of a Cistercian Monastery
at Sedlitz, on which it is said that certain monks were once hung: hence the
legend has arisen, that the peculiar form of the leaf was given in order to
perpetuate the memory of the martyred monks. ('Bayer. Monogr. Tiliæ,'
Berlin, 1861.) It is also stated that this condition is not perpetuated by
grafting.
Vereins, f. d. Preuss., Rheinl. und Westphal., 1860, p. 347, tab. vii; Hallier, 'Phytopathol.,'
p. 128; Boehmer, 'De plantis Fasciatis,' Wittenb., 1752.
Cohesion of foliar organs.—This takes place in several ways, and in very
various degrees; the simplest case is that characterised by the cohesion of
the margins of the same organ, as in the condition called perfoliate in
descriptive works, and which is due either to a cohesion of the margins of
the basal lobes of the leaf, or to the development of the leaf in a sheathing
or tubular manner. As an abnormal occurrence, I have met with this
perfoliation in a leaf of Goodenia ovata. The condition in question is often
loosely confounded with connation, or the union of two leaves by their
bases. In other cases the union takes place between the margins of two or
more leaves.
Cohesion of margins of single organs.—The leaves of Hazels may often
be found with their margins coherent at the base, so as to become peltate,
while in other cases, the disc of the leaf is so depressed that a true pitcher is
formed. This happens also in the Lime Tilia, in which genus pitcher- or
hood-like leaves (folia cucullata) may frequently be met with. There are
trees with leaves of this character in the cemetery of a Cistercian Monastery
at Sedlitz, on which it is said that certain monks were once hung: hence the
legend has arisen, that the peculiar form of the leaf was given in order to
perpetuate the memory of the martyred monks. ('Bayer. Monogr. Tiliæ,'
Berlin, 1861.) It is also stated that this condition is not perpetuated by
grafting.
Page 63
Fig. 8.—Pitcher-shaped leaf of Pelargonium.
I have in my possession a leaf of Antirrhinum majus, and also a specimen of
Pelargonium, wherein the blade of the leaf is funnel-like, and the petiole is
cylindrical, not compressed, and grooved on the upper surface, as is usually
the case. A comparison of the leaves of Pelargonium peltatum with those of
P. cucullatum ('Cav. Diss.,' tab., 106) will show how easy the passage is
from a peltate to a tubular leaf. In these cases the tubular form may rather
be due to dilatation than to cohesion. M. Kickx[16] mentions an instance of
the kind in the leaves of a species of Nicotiana, and also figures the leaf of a
rose in which two opposite leaflets presented themselves in the form of
stalked cups. Schlechtendal[17] notices something of the same kind in the
leaf of Amorpha fruticosa; Treviranus[18] in that of Aristolochia Sipho.
I have in my possession a leaf of Antirrhinum majus, and also a specimen of
Pelargonium, wherein the blade of the leaf is funnel-like, and the petiole is
cylindrical, not compressed, and grooved on the upper surface, as is usually
the case. A comparison of the leaves of Pelargonium peltatum with those of
P. cucullatum ('Cav. Diss.,' tab., 106) will show how easy the passage is
from a peltate to a tubular leaf. In these cases the tubular form may rather
be due to dilatation than to cohesion. M. Kickx[16] mentions an instance of
the kind in the leaves of a species of Nicotiana, and also figures the leaf of a
rose in which two opposite leaflets presented themselves in the form of
stalked cups. Schlechtendal[17] notices something of the same kind in the
leaf of Amorpha fruticosa; Treviranus[18] in that of Aristolochia Sipho.
Page 64
M. Puel[19] describes a leaf of Polygonatum multiflorum, the margins of
which were so completely united together, as only to leave a circular
aperture at the top, through which passed the ends of the leaves. The Rev.
Mr. Hincks, at the meeting of the British Association at Newcastle (1838),
showed a leaf of a Tulip, whose margins were so united that the whole leaf
served as a hood, and was carried upwards by the growing flower like the
calyptra of a Moss.
The margins of the stipules are also occasionally united, so as to form a
little horn-shaped tube. I have met with instances of this kind in the
common white clover, Trifolium repens, where on each side of the base of
the petiole the stipules had the form just indicated. That the bracts also may
assume this condition, may be inferred from the peculiar horn-like
structures of Marcgraavia, which appear to originate from the union of the
margins of the reflected leaf.
Tubular petals occur normally in some flowers, as Helleborus, Epimedium,
Viola, &c., and as an exceptional occurrence I have seen them in
Ranunculus repens, while in Eranthis hyemalis transitions may frequently
be seen between the flat outer segments of the perianth and the tubular
petals. To Dr. Sankey, of Sandywell Park, I am indebted for the flower of a
Pelargonium, in which one of the petals had the form of a cup supported on
a long stalk. This cup-shaped organ was placed at the back of the flower,
and had the dark colour proper to the petals in that situation. I have seen a
petal of Clarkia similarly tubular, while some of the cultivated varieties of
Primula sinensis exhibit tubular petals so perfect in shape as closely to
resemble perfect corollas.
which were so completely united together, as only to leave a circular
aperture at the top, through which passed the ends of the leaves. The Rev.
Mr. Hincks, at the meeting of the British Association at Newcastle (1838),
showed a leaf of a Tulip, whose margins were so united that the whole leaf
served as a hood, and was carried upwards by the growing flower like the
calyptra of a Moss.
The margins of the stipules are also occasionally united, so as to form a
little horn-shaped tube. I have met with instances of this kind in the
common white clover, Trifolium repens, where on each side of the base of
the petiole the stipules had the form just indicated. That the bracts also may
assume this condition, may be inferred from the peculiar horn-like
structures of Marcgraavia, which appear to originate from the union of the
margins of the reflected leaf.
Tubular petals occur normally in some flowers, as Helleborus, Epimedium,
Viola, &c., and as an exceptional occurrence I have seen them in
Ranunculus repens, while in Eranthis hyemalis transitions may frequently
be seen between the flat outer segments of the perianth and the tubular
petals. To Dr. Sankey, of Sandywell Park, I am indebted for the flower of a
Pelargonium, in which one of the petals had the form of a cup supported on
a long stalk. This cup-shaped organ was placed at the back of the flower,
and had the dark colour proper to the petals in that situation. I have seen a
petal of Clarkia similarly tubular, while some of the cultivated varieties of
Primula sinensis exhibit tubular petals so perfect in shape as closely to
resemble perfect corollas.
Page 65
Fig. 9.—Eranthis hyemalis. Transition from flat sepal to tubular petal.
Like the petals, the stamens, and even the styles, assume a hollow tubular
form. This change of form in the case of the stamens is, of course, usually
attended by the petaloid expansion of the filament, or anther, and the more
or less complete obliteration of the pollen sacs, as in Fuchsias, and in some
double-flowered Antirrhinums.[20] So also in some semi-double varieties of
Narcissus poeticus, and in Aquilegia. By the late Professor Charles Morren,
this affection of the stamens and pistils was called Solenaidie,[21] but as a
similar condition exists in other organs, it hardly seems worth while to
adopt a special term for the phenomenon, as it presents itself in one set of
organs.
In many of these cases it is difficult to say whether the cup-like or tubular
form is due to a dilatation or hollowing out of the organ affected, or to a
fusion of its edges. The arrangement of the veins will in some cases supply
the clue, and in others the regularity of form will indicate the nature of the
malformation, for in those instances where the cup is the result of
expansion, its margin is more likely to be regular and even than in those
where the hollow form is the result of fusion.
Cohesion of several organs by their margins:—leaves, &c.—The union
of the margins of two or more different organs is of more common
Like the petals, the stamens, and even the styles, assume a hollow tubular
form. This change of form in the case of the stamens is, of course, usually
attended by the petaloid expansion of the filament, or anther, and the more
or less complete obliteration of the pollen sacs, as in Fuchsias, and in some
double-flowered Antirrhinums.[20] So also in some semi-double varieties of
Narcissus poeticus, and in Aquilegia. By the late Professor Charles Morren,
this affection of the stamens and pistils was called Solenaidie,[21] but as a
similar condition exists in other organs, it hardly seems worth while to
adopt a special term for the phenomenon, as it presents itself in one set of
organs.
In many of these cases it is difficult to say whether the cup-like or tubular
form is due to a dilatation or hollowing out of the organ affected, or to a
fusion of its edges. The arrangement of the veins will in some cases supply
the clue, and in others the regularity of form will indicate the nature of the
malformation, for in those instances where the cup is the result of
expansion, its margin is more likely to be regular and even than in those
where the hollow form is the result of fusion.
Cohesion of several organs by their margins:—leaves, &c.—The union
of the margins of two or more different organs is of more common
Page 66
occurrence than the preceding, the leaves being frequently subjected to this
change. Occasionally, the leaflets of a compound leaf have been observed
united by their margins, as in the strawberry, the white trefoil, and others.
Sometimes the union takes place by means of the stalks only. I have an
instance of this in a Pelargonium, in Tropæolum majus, and Strelitzia
regina; in other cases, the whole extent of the leaf becomes joined to its
neighbour, the leaves thus becoming completely united by their edges, as in
those of Justicia, oxyphylla.[22] M. Clos[23] has observed the same thing in
the leaves of the lentil Ervum lens, conjoined with fasciation of the stem,
and many other examples might be given. Some of the recorded cases are
probably really due to fission of one leaf into two rather than to fusion.
Although usually the lower portions of the leaf are united together, leaving
the upper parts more or less detached, there are some instances in which the
margins of the leaf at their upper portion have been noticed to be coherent,
while their lower portions, with their stalks, were completely free.[24]
Cohesion of the leaves frequently accompanies the union of the branches
and fasciation as might have been anticipated. Moquin cites the fenestrated
leaves of Dracontium pertusum, as well as some cases of a similar kind that
are occasionally met with, as instances of the cohesion of the margins at the
base and apex of the leaf, which thus appears perforated. This appearance,
however, is probably due to some other cause. When the leaves are
verticillate and numerous, and they become coherent by their margins, they
form a foliaceous tube around the stem. When there are but two opposite
leaves, and these become united by their margins, we have a state of things
precisely resembling that to which the term connate is applied.
Fusion of the edges of the cotyledons also occasionally takes place, as in
Ebenus cretica.[25] It has also been observed in Tithonia, and is of constant
occurrence in the seed leaves of some Mesembryanthema. This condition
must be carefully distinguished from the very similar appearance produced
change. Occasionally, the leaflets of a compound leaf have been observed
united by their margins, as in the strawberry, the white trefoil, and others.
Sometimes the union takes place by means of the stalks only. I have an
instance of this in a Pelargonium, in Tropæolum majus, and Strelitzia
regina; in other cases, the whole extent of the leaf becomes joined to its
neighbour, the leaves thus becoming completely united by their edges, as in
those of Justicia, oxyphylla.[22] M. Clos[23] has observed the same thing in
the leaves of the lentil Ervum lens, conjoined with fasciation of the stem,
and many other examples might be given. Some of the recorded cases are
probably really due to fission of one leaf into two rather than to fusion.
Although usually the lower portions of the leaf are united together, leaving
the upper parts more or less detached, there are some instances in which the
margins of the leaf at their upper portion have been noticed to be coherent,
while their lower portions, with their stalks, were completely free.[24]
Cohesion of the leaves frequently accompanies the union of the branches
and fasciation as might have been anticipated. Moquin cites the fenestrated
leaves of Dracontium pertusum, as well as some cases of a similar kind that
are occasionally met with, as instances of the cohesion of the margins at the
base and apex of the leaf, which thus appears perforated. This appearance,
however, is probably due to some other cause. When the leaves are
verticillate and numerous, and they become coherent by their margins, they
form a foliaceous tube around the stem. When there are but two opposite
leaves, and these become united by their margins, we have a state of things
precisely resembling that to which the term connate is applied.
Fusion of the edges of the cotyledons also occasionally takes place, as in
Ebenus cretica.[25] It has also been observed in Tithonia, and is of constant
occurrence in the seed leaves of some Mesembryanthema. This condition
must be carefully distinguished from the very similar appearance produced
Page 67
by quite a different cause, viz., the splitting of one cotyledon into two,
which gives rise to the appearance as if two were partially united together.
Some of the ascidia or pitcher-like formations are due to the cohesion of the
margins of two leaves, as in a specimen of Crassula arborescens, observed
by C. Morren.
Fig. 10.—Two-leaved pitcher of Crassula arborescens, after C. Morren.
The stipules may also be fused together in different ways; their edges
sometimes cohere between the leaf and the stem, and thus form a solitary
intra-axillary stipule. At other times they become united in such a manner as
to produce a single notched stipule opposite to the leaf. Again, in other
cases, they are so united on each side of the stem, that in place of four there
seem only to exist two, common to the two leaves as in the Hop.
which gives rise to the appearance as if two were partially united together.
Some of the ascidia or pitcher-like formations are due to the cohesion of the
margins of two leaves, as in a specimen of Crassula arborescens, observed
by C. Morren.
Fig. 10.—Two-leaved pitcher of Crassula arborescens, after C. Morren.
The stipules may also be fused together in different ways; their edges
sometimes cohere between the leaf and the stem, and thus form a solitary
intra-axillary stipule. At other times they become united in such a manner as
to produce a single notched stipule opposite to the leaf. Again, in other
cases, they are so united on each side of the stem, that in place of four there
seem only to exist two, common to the two leaves as in the Hop.
Page 68
To the Rev. M. J. Berkeley I am indebted for specimens of a curious
pitcher-like formation in the garden Pea. The structure in question consisted
of a stalked foliaceous cup proceeding from the inflorescence. On
examination of the ordinary inflorescence, there will be seen at the base of
the upper of two flowers a small rudimentary bract, having a swollen
circular or ring-like base, from which proceeds a small awl-shaped process,
representing the midrib of an abortive leaf. In some of Mr. Berkeley's
specimens, the stipules were developed as leafy appendages at the base of
the leaf-stalk or midrib, the latter retaining its shortened form, while, in
others, the two stipules had become connate into a cup, and all trace of the
midrib was lost. The cup in question would thus seem to have been formed
from the connation of two stipules which are ordinarily abortive.
Cohesion of the bracts by their edges, so as to form a tubular involucre, or
by their surfaces, so as to form a cupule, is not of uncommon occurrence,
under natural conditions, and may be met with in plants which ordinarily do
not exhibit this appearance.
Cohesion of the sepals in a normally polypetalous calyx renders the latter
gamosepalous, and is not of uncommon occurrence, to a partial extent,
though rarely met with complete. I have observed a junction of the sepals to
be one of the commonest malformations among Orchids, indeed such a
state of things occurs normally in Masdevallia Cypripedium, &c. An
illustration of this occurrence is given by Mr. J. T. Moggridge in Ophrys
insectifera, in 'Seemann's Journal of Botany,' 1866, p. 168, tab. 47. In
Orchids, this cohesion of sepals is very often co-existent with other more
important changes, such as absence of the labellum, dislocation of the parts
of the flower, &c.
pitcher-like formation in the garden Pea. The structure in question consisted
of a stalked foliaceous cup proceeding from the inflorescence. On
examination of the ordinary inflorescence, there will be seen at the base of
the upper of two flowers a small rudimentary bract, having a swollen
circular or ring-like base, from which proceeds a small awl-shaped process,
representing the midrib of an abortive leaf. In some of Mr. Berkeley's
specimens, the stipules were developed as leafy appendages at the base of
the leaf-stalk or midrib, the latter retaining its shortened form, while, in
others, the two stipules had become connate into a cup, and all trace of the
midrib was lost. The cup in question would thus seem to have been formed
from the connation of two stipules which are ordinarily abortive.
Cohesion of the bracts by their edges, so as to form a tubular involucre, or
by their surfaces, so as to form a cupule, is not of uncommon occurrence,
under natural conditions, and may be met with in plants which ordinarily do
not exhibit this appearance.
Cohesion of the sepals in a normally polypetalous calyx renders the latter
gamosepalous, and is not of uncommon occurrence, to a partial extent,
though rarely met with complete. I have observed a junction of the sepals to
be one of the commonest malformations among Orchids, indeed such a
state of things occurs normally in Masdevallia Cypripedium, &c. An
illustration of this occurrence is given by Mr. J. T. Moggridge in Ophrys
insectifera, in 'Seemann's Journal of Botany,' 1866, p. 168, tab. 47. In
Orchids, this cohesion of sepals is very often co-existent with other more
important changes, such as absence of the labellum, dislocation of the parts
of the flower, &c.
Page 69
Fig. 11.—Gamopetalous flower of Papaver bracteatum.
Cohesion of the petals.—Linnæus mentions the occurrence of cohesion of
the petals in Saponaria.[26] Moquin notices a Rose in which the petals were
united into a long tube, their upper portions were free and bent downwards,
forming a sort of irregular limb. An instance of the polypetalous regular
perianth of Clematis viticella being changed into a monopetalous irregular
one, like the corolla of Labiates, is recorded by Jaeger.[27] There is in
cultivation a variety of Papaver bracteatum, in which the petals are united
by their margins so as to form a large cup. Under normal circumstances, the
petals become fused together by their edges along their whole extent, at the
base only, at the apex only, as in the Vine, or at the base and apex, leaving
the central portions detached. Indications of the junction of the petals may
generally be traced by the arrangement of the veins, or by the notches or
lobes left by imperfect union. In Crocuses I have frequently met with
cohesion of the segments of the perianth, by means of their surfaces, but the
Cohesion of the petals.—Linnæus mentions the occurrence of cohesion of
the petals in Saponaria.[26] Moquin notices a Rose in which the petals were
united into a long tube, their upper portions were free and bent downwards,
forming a sort of irregular limb. An instance of the polypetalous regular
perianth of Clematis viticella being changed into a monopetalous irregular
one, like the corolla of Labiates, is recorded by Jaeger.[27] There is in
cultivation a variety of Papaver bracteatum, in which the petals are united
by their margins so as to form a large cup. Under normal circumstances, the
petals become fused together by their edges along their whole extent, at the
base only, at the apex only, as in the Vine, or at the base and apex, leaving
the central portions detached. Indications of the junction of the petals may
generally be traced by the arrangement of the veins, or by the notches or
lobes left by imperfect union. In Crocuses I have frequently met with
cohesion of the segments of the perianth, by means of their surfaces, but the
Page 70
union was confined to the centre of the segment, leaving the rest of the
surfaces free.
Cohesion of the stamens.—Under natural circumstances, cohesion of the
stamens is said to take place either by the union of their filaments, so as to
form one, two, or more parcels (Monadelphia, Diadelphia, Polyadelphia); at
other times, by the cohesion of the anthers (Syngenesia), in which latter
case the union is generally very slight. It must be remembered, however,
that the so-called cohesion of the filaments is in many cases due rather to
the formation of compound stamens, i.e. to the formation from one original
staminal tubercle of numerous secondary ones, so that the process is rather
one of over development than of fusion or of disjunction. These conditions
may be met with as accidental occurrences in plants or in flowers, not
usually showing this arrangement. Thus, for instance, Professor Andersson,
of Stockholm, describes a monstrosity of Salix calyculata, in which the
stamens were so united together as to form a tube open at the top like a
follicle.[28] This is an exaggerated degree of that fusion which exists
normally in Salix monandra, in Cucurbits and other plants.
Cohesion of the pistils is also of very frequent occurrence in plants, under
ordinary circumstances, but is less commonly met with than might have
been expected as a teratological phenomenon.
Further details relating to cohesion of the various parts of the flower are cited in Moquin-
Tandon, 'El. Ter. Veg.,' p. 248; 'Weber. Verhandl. Nat. Hist. Vereins f. d. Preuss. Rheinl. und
Westphal.,' 1860, p. 332, tabs. 6 et 7.
Formation of ascidia or pitchers.—In the preceding paragraphs, the
formation of tubular or horn-like structures, from the union of the margins
of one organ, or from the coalescence, or it may be from the want of
separation of various organs, has been alluded to, so that it seems only
necessary now, by way of summary, to mention the classification of ascidia
surfaces free.
Cohesion of the stamens.—Under natural circumstances, cohesion of the
stamens is said to take place either by the union of their filaments, so as to
form one, two, or more parcels (Monadelphia, Diadelphia, Polyadelphia); at
other times, by the cohesion of the anthers (Syngenesia), in which latter
case the union is generally very slight. It must be remembered, however,
that the so-called cohesion of the filaments is in many cases due rather to
the formation of compound stamens, i.e. to the formation from one original
staminal tubercle of numerous secondary ones, so that the process is rather
one of over development than of fusion or of disjunction. These conditions
may be met with as accidental occurrences in plants or in flowers, not
usually showing this arrangement. Thus, for instance, Professor Andersson,
of Stockholm, describes a monstrosity of Salix calyculata, in which the
stamens were so united together as to form a tube open at the top like a
follicle.[28] This is an exaggerated degree of that fusion which exists
normally in Salix monandra, in Cucurbits and other plants.
Cohesion of the pistils is also of very frequent occurrence in plants, under
ordinary circumstances, but is less commonly met with than might have
been expected as a teratological phenomenon.
Further details relating to cohesion of the various parts of the flower are cited in Moquin-
Tandon, 'El. Ter. Veg.,' p. 248; 'Weber. Verhandl. Nat. Hist. Vereins f. d. Preuss. Rheinl. und
Westphal.,' 1860, p. 332, tabs. 6 et 7.
Formation of ascidia or pitchers.—In the preceding paragraphs, the
formation of tubular or horn-like structures, from the union of the margins
of one organ, or from the coalescence, or it may be from the want of
separation of various organs, has been alluded to, so that it seems only
necessary now, by way of summary, to mention the classification of ascidia
Page 71
proposed by Professor Charles Morren[29], who divides the structures in
question into two heads, according as they are formed from one or more
leaves. The following list is arranged according to the views of the Belgian
savant, and comprises a few additional illustrations. Those to which the ! is
affixed have been seen by the writer himself; the * indicates the more
frequent occurrence of the phenomenon in some than in other plants. Those
plants, such as Nepenthes, &c., which occur normally and constantly, are
not here included. Possibly some of the cases would be more properly
classed under dilatation or excavation.
Ascidia.
A. Monophyllous.
1. Sarracenia-like pitchers, formed by a single leaf, the edges of which are united for the
greater portion of their length, but are disunited near the top, so as to leave an oblique
aperture.
*Brassica oleracea (several of the cultivated varieties)!
*Tilia europæa!
Pelargonium inquinans!
Staphylea pinnata.
Amorpha fruticosa.
Pisum sativum!
Lathyrus tuberosus.
Vicia sp.
Gleditschia sp.
Ceratonia siliqua.
Trifolium repens!
Cassia marylandica.
Mimosa Lophantha.
Rosa centifolia.
gallica.
Begonia sp.
Bellis perennis!
Nicotiana sp.
Goodenia ovata!
Antirrhinum majus!
question into two heads, according as they are formed from one or more
leaves. The following list is arranged according to the views of the Belgian
savant, and comprises a few additional illustrations. Those to which the ! is
affixed have been seen by the writer himself; the * indicates the more
frequent occurrence of the phenomenon in some than in other plants. Those
plants, such as Nepenthes, &c., which occur normally and constantly, are
not here included. Possibly some of the cases would be more properly
classed under dilatation or excavation.
Ascidia.
A. Monophyllous.
1. Sarracenia-like pitchers, formed by a single leaf, the edges of which are united for the
greater portion of their length, but are disunited near the top, so as to leave an oblique
aperture.
*Brassica oleracea (several of the cultivated varieties)!
*Tilia europæa!
Pelargonium inquinans!
Staphylea pinnata.
Amorpha fruticosa.
Pisum sativum!
Lathyrus tuberosus.
Vicia sp.
Gleditschia sp.
Ceratonia siliqua.
Trifolium repens!
Cassia marylandica.
Mimosa Lophantha.
Rosa centifolia.
gallica.
Begonia sp.
Bellis perennis!
Nicotiana sp.
Goodenia ovata!
Antirrhinum majus!
Page 72
Vinca rosea.
Polygonum orientale.
Aristolochia sipho?
Codiæum variegatum var.!
Spinacia oleracea.
Corylus avellana!
Polygonatum multiflorum.
Xanthosoma appendiculatum!
2. Calyptriform or hood-like pitchers, formed by the complete union of the margins, and
falling off by a transverse fissure (as in the calyx of Escholtzia).
Tulipa Gesneriana.
B. Polyphyllous.
1. Diphyllous, formed by the union of two leaves into a single cup, tube, or funnel, &c.
Pisum sativum (stipules)!
Crassula arborescens.
Polygonatum multiflorum.
2. Triphyllous, formed by the union of three leaves.
Paris quadrifolia var.
Besides the above varieties of ascidia formed from the union of one or more leaves, there
are others which seem to be the result of a peculiar excrescence or hypertrophy of the leaf.
Such are some of the curious pitcher-like structures met with occasionally in the leaves of
cabbages, lettuces, Aristolochia, &c. See Hypertrophy, cup-like deformities, &c.
In addition to other publications previously mentioned, reference may be made to the
following treatises on the subject of ascidia:—Bonnet, 'Rech. Us. Feuilles,' p. 216, tab.
xxvi, f. 1, Brassica; De Candolle, 'Trans. Hort. Soc.,' t. v, pl. 1, Brassica; Id., 'Org. Veget.,'
I, 316; 'Bull. Soc. Bot. Fr.,' I, p. 62, Polygonatum; 'Bull. Acad. Belg.,' 1851, p. 591, Rosa;
Hoffmann, 'Tijdschrift v. Natuur. Geschied.,' vol. viii, p. 318, tab. 9, Ceratonia; C. Mulder,
'Tijdschrift, &c.,' vol. vi, p. 106, tab. 5, 6, Trifolium, Mimosa, Staphylea;' Molkenboer,' p.
115, t. 4, Brassica.
FOOTNOTES:
[10] See a curious instance of this kind in the branches of Pinus. 'Regel. Garten Flora,' vol.
8, tab. 268.
[11] 'Bull. Soc. Bot. France,' 1860, p. 881.
Polygonum orientale.
Aristolochia sipho?
Codiæum variegatum var.!
Spinacia oleracea.
Corylus avellana!
Polygonatum multiflorum.
Xanthosoma appendiculatum!
2. Calyptriform or hood-like pitchers, formed by the complete union of the margins, and
falling off by a transverse fissure (as in the calyx of Escholtzia).
Tulipa Gesneriana.
B. Polyphyllous.
1. Diphyllous, formed by the union of two leaves into a single cup, tube, or funnel, &c.
Pisum sativum (stipules)!
Crassula arborescens.
Polygonatum multiflorum.
2. Triphyllous, formed by the union of three leaves.
Paris quadrifolia var.
Besides the above varieties of ascidia formed from the union of one or more leaves, there
are others which seem to be the result of a peculiar excrescence or hypertrophy of the leaf.
Such are some of the curious pitcher-like structures met with occasionally in the leaves of
cabbages, lettuces, Aristolochia, &c. See Hypertrophy, cup-like deformities, &c.
In addition to other publications previously mentioned, reference may be made to the
following treatises on the subject of ascidia:—Bonnet, 'Rech. Us. Feuilles,' p. 216, tab.
xxvi, f. 1, Brassica; De Candolle, 'Trans. Hort. Soc.,' t. v, pl. 1, Brassica; Id., 'Org. Veget.,'
I, 316; 'Bull. Soc. Bot. Fr.,' I, p. 62, Polygonatum; 'Bull. Acad. Belg.,' 1851, p. 591, Rosa;
Hoffmann, 'Tijdschrift v. Natuur. Geschied.,' vol. viii, p. 318, tab. 9, Ceratonia; C. Mulder,
'Tijdschrift, &c.,' vol. vi, p. 106, tab. 5, 6, Trifolium, Mimosa, Staphylea;' Molkenboer,' p.
115, t. 4, Brassica.
FOOTNOTES:
[10] See a curious instance of this kind in the branches of Pinus. 'Regel. Garten Flora,' vol.
8, tab. 268.
[11] 'Bull. Soc. Bot. France,' 1860, p. 881.
Page 73
[12] Ibid., 1861, p. 708.
[13] Ibid., 1860, p. 923.
[14] 'Proc. Linn. Soc.,' April 5, 1853.
[15] 'Organ. Végét.,' pl. iii, fig. 1.
[16] 'Bull. Acad. Roy. Bruxelles,' t. xviii, p. i and p. 591.
[17] 'Linnæa,' tom. 13, p. 383.
[18] 'Verhandl. Nat. Hist. Vereins,' 1859, Bonn, tom. xvi, tab. 3.
[19] 'Bull. Soc. Bot. Fr.,' vol. i, p. 62.
[20] 'Report of Internat. Bot. Congress,' London, 1866, p. 131, tab. vii, figs. 10–13.
[21] 'Bull. Acad. Roy. Belg.,' t. xviii, 2nd part, p. 179.
[22] D. C., 'Organ. Végét.,' pl. xvii, fig. 3, and pl. xlviii, fig. 2.
[23] 'Mém. Acad. Toulouse,' 1862.
[24] Bonnet, 'Recherches Us. feuill.,' pl. xxi, fig. 2.
[25] De Candolle, 'Mém. Lég.,' pl. v, fig. 14.
[26] 'Phil. Bot.,' § 125.
[27] 'Nov. Act. Acad. Nat. Cur.,' 14, p. 642, t. xxxvii.
[28] 'Journal of the Linn. Soc. Bot.,' vol. iv, p. 55.
[29] 'Bull. Acad. Roy. Bruxelles,' 1838, t. v, p. 582. 'Bull. Acad. Roy. Belg.,' 1852, t. xix,
part iii, p. 437.
CHAPTER II.
ADHESION.
[13] Ibid., 1860, p. 923.
[14] 'Proc. Linn. Soc.,' April 5, 1853.
[15] 'Organ. Végét.,' pl. iii, fig. 1.
[16] 'Bull. Acad. Roy. Bruxelles,' t. xviii, p. i and p. 591.
[17] 'Linnæa,' tom. 13, p. 383.
[18] 'Verhandl. Nat. Hist. Vereins,' 1859, Bonn, tom. xvi, tab. 3.
[19] 'Bull. Soc. Bot. Fr.,' vol. i, p. 62.
[20] 'Report of Internat. Bot. Congress,' London, 1866, p. 131, tab. vii, figs. 10–13.
[21] 'Bull. Acad. Roy. Belg.,' t. xviii, 2nd part, p. 179.
[22] D. C., 'Organ. Végét.,' pl. xvii, fig. 3, and pl. xlviii, fig. 2.
[23] 'Mém. Acad. Toulouse,' 1862.
[24] Bonnet, 'Recherches Us. feuill.,' pl. xxi, fig. 2.
[25] De Candolle, 'Mém. Lég.,' pl. v, fig. 14.
[26] 'Phil. Bot.,' § 125.
[27] 'Nov. Act. Acad. Nat. Cur.,' 14, p. 642, t. xxxvii.
[28] 'Journal of the Linn. Soc. Bot.,' vol. iv, p. 55.
[29] 'Bull. Acad. Roy. Bruxelles,' 1838, t. v, p. 582. 'Bull. Acad. Roy. Belg.,' 1852, t. xix,
part iii, p. 437.
CHAPTER II.
ADHESION.
Page 74
Adhesion, so called, occurs either from actual union of originally distinct
members of different whorls or from the non-occurrence of that separation
which usually takes place between them. It is thus in some degree a graver
deviation than cohesion, and is generally a consequence of, or at least is
coexistent with, more serious changes; thus if two leaves of the same whorl
are coherent the change is not very great, but if two leaves belonging to
different whorls, or two leaves in the same spiral cycle are adherent, a
deformation in the axis or a certain amount of dislocation must almost
necessarily exist. Adhesion as a normal occurrence is usually the result of a
lack of separation rather than of union of parts primitively separate.
Instances of adhesion between different organs is seen under ordinary
circumstances in the bract of the Lime tree, which adheres to the peduncle,
also in Neuropeltis, while in Erythrochiton hypophyllanthus the cymose
peduncles are adherent to the under surface of the leaf.
Adhesion between the axes of the same plant is sufficiently treated of under
the head of Cohesion, from which it is in this instance impossible to make a
distinction. Adhesion of the inflorescence is necessarily a frequent
accompaniment of fasciation and cohesion of the branches.
Adhesion of foliar organs may occur either between the margins or
between the surfaces of the affected parts; in the former case there is almost
necessarily more or less displacement and change of direction, such as a
twisting of the stem and a vertical rather than a horizontal attachment of the
foliar organ to it; hence it generally forms but a part of other and more
important deviations.
Adhesion of leaves by their surfaces.—The union of leaves by their
surfaces is not of very frequent occurrence, many of the instances cited
being truly referable to other conditions. Bonnet describes the union of two
lettuce leaves, and Turpin that of two leaves of Agave americana, in which
members of different whorls or from the non-occurrence of that separation
which usually takes place between them. It is thus in some degree a graver
deviation than cohesion, and is generally a consequence of, or at least is
coexistent with, more serious changes; thus if two leaves of the same whorl
are coherent the change is not very great, but if two leaves belonging to
different whorls, or two leaves in the same spiral cycle are adherent, a
deformation in the axis or a certain amount of dislocation must almost
necessarily exist. Adhesion as a normal occurrence is usually the result of a
lack of separation rather than of union of parts primitively separate.
Instances of adhesion between different organs is seen under ordinary
circumstances in the bract of the Lime tree, which adheres to the peduncle,
also in Neuropeltis, while in Erythrochiton hypophyllanthus the cymose
peduncles are adherent to the under surface of the leaf.
Adhesion between the axes of the same plant is sufficiently treated of under
the head of Cohesion, from which it is in this instance impossible to make a
distinction. Adhesion of the inflorescence is necessarily a frequent
accompaniment of fasciation and cohesion of the branches.
Adhesion of foliar organs may occur either between the margins or
between the surfaces of the affected parts; in the former case there is almost
necessarily more or less displacement and change of direction, such as a
twisting of the stem and a vertical rather than a horizontal attachment of the
foliar organ to it; hence it generally forms but a part of other and more
important deviations.
Adhesion of leaves by their surfaces.—The union of leaves by their
surfaces is not of very frequent occurrence, many of the instances cited
being truly referable to other conditions. Bonnet describes the union of two
lettuce leaves, and Turpin that of two leaves of Agave americana, in which
Page 75
latter the upper surface of one leaf was adherent to the lower surface of the
leaf next above it, and I have myself met with similar instances in the
wallflower and in lettuce and cabbage leaves; other instances have been
mentioned in Saxifraga, Gesnera, &c.[30]
In these cases, owing to the non-development of the internodes, the nascent
leaves are closely packed, and the conditions for adhesion are favorable, but
in most of the so-called cases of adhesion of leaf to leaf by the surface, a
preferable explanation is afforded either by an exuberant development
(hypertrophy) or by chorisis (see sections on those subjects). Thus, when a
leaf of this kind is apparently so united, that the lower surface of one is
adherent to the corresponding surface of another, the phenomenon is
probably due rather to extra development or to fission. There is an
exception to this, however, in the case of two vertically-erect leaves on
opposite sides of the stem; here the two upper or inner surfaces may
become adherent, as in an orange, where two leaves were thus united, the
terminal bud between them being suppressed or abortive.
Adhesion between the membranous bract of Narcissus poeticus and the
upper surface of the leaf is described by Moquin.[31] The same author
mentions having seen a remarkable example of adhesion in the involucels
of Caucalis leptophylla, the bracts of which were soldered to the outer
surface of the flowers. M. Bureau[32] mentions an instance wherein the
spathe of Narcissus biflorus was partially twisted in such a manner that the
lower surface of its median nerve was adherent to the corresponding surface
of one of the sepals, mid-rib to mid-rib, thus apparently confirming a law of
G. de Hilaire, that when two parts of the same individual unite, they
generally do so by the corresponding surfaces or edges, but the rule is
probably not so general in its application as has been supposed.
leaf next above it, and I have myself met with similar instances in the
wallflower and in lettuce and cabbage leaves; other instances have been
mentioned in Saxifraga, Gesnera, &c.[30]
In these cases, owing to the non-development of the internodes, the nascent
leaves are closely packed, and the conditions for adhesion are favorable, but
in most of the so-called cases of adhesion of leaf to leaf by the surface, a
preferable explanation is afforded either by an exuberant development
(hypertrophy) or by chorisis (see sections on those subjects). Thus, when a
leaf of this kind is apparently so united, that the lower surface of one is
adherent to the corresponding surface of another, the phenomenon is
probably due rather to extra development or to fission. There is an
exception to this, however, in the case of two vertically-erect leaves on
opposite sides of the stem; here the two upper or inner surfaces may
become adherent, as in an orange, where two leaves were thus united, the
terminal bud between them being suppressed or abortive.
Adhesion between the membranous bract of Narcissus poeticus and the
upper surface of the leaf is described by Moquin.[31] The same author
mentions having seen a remarkable example of adhesion in the involucels
of Caucalis leptophylla, the bracts of which were soldered to the outer
surface of the flowers. M. Bureau[32] mentions an instance wherein the
spathe of Narcissus biflorus was partially twisted in such a manner that the
lower surface of its median nerve was adherent to the corresponding surface
of one of the sepals, mid-rib to mid-rib, thus apparently confirming a law of
G. de Hilaire, that when two parts of the same individual unite, they
generally do so by the corresponding surfaces or edges, but the rule is
probably not so general in its application as has been supposed.
Page 76
Adhesion of foliar to axile organs.—The appendicular organs may
likewise be found united to the axile ones. This union takes place in many
ways; sometimes the leaves do not become detached from the stem for a
considerable distance, as in the so-called decurrent leaves, at other times the
leaves are prolonged at their base into lobes, which are directed along the
stem, and are united with it. Turpin records a tendril of a vine which was
fused with the stem for some distance, and bore leaves and other tendrils.
Union of the leaf or bract with the flower-stalk is not uncommon. It occurs
normally in the Lime and other plants.
Adhesion of the sepals to the petals is spoken of by Morren as
calyphyomy, καλυξ φυομαι.[33] Moquin cites an instance in Geranium
nodosum, in which one petal was united by its lower surface to one of the
segments of the calyx. A similar circumstance has been observed in Petunia
violacea by Morren. Duchartre describes an instance wherein one of the
outer sepals of Cattleya Forbesii was adherent to the labellum.[34]
Adhesion of the stamens to the petals is of common occurrence under
natural circumstances. Cassini has described a malformation of Centaurea
collina, in which two of the five stamens were completely grafted with the
corolla, the three others remaining perfectly free. Adhesion of the petals to
the column is not of infrequent occurrence among Orchids. I have observed
cases of the adhesion of the segments of the perianth to the stamen in
Ophrys aranifera, Odontoglossum, sp. &c. It is the ordinary condition in
Gongora and some other genera. I have seen it also in Lilium lancifolium.
Some forms of Crocus, occasionally met with, present a very singular
appearance, owing to the adhesion of the stamens to the outer segments of
the perianth, the former, moreover, being partially petaloid in aspect. M. de
la Vaud[35] speaks of a similar union in Tigridia pavonia. Morren[36]
describes a malformation of Fuchsia wherein the petals were so completely
adherent to the stamens, that the former were dragged out of their ordinary
likewise be found united to the axile ones. This union takes place in many
ways; sometimes the leaves do not become detached from the stem for a
considerable distance, as in the so-called decurrent leaves, at other times the
leaves are prolonged at their base into lobes, which are directed along the
stem, and are united with it. Turpin records a tendril of a vine which was
fused with the stem for some distance, and bore leaves and other tendrils.
Union of the leaf or bract with the flower-stalk is not uncommon. It occurs
normally in the Lime and other plants.
Adhesion of the sepals to the petals is spoken of by Morren as
calyphyomy, καλυξ φυομαι.[33] Moquin cites an instance in Geranium
nodosum, in which one petal was united by its lower surface to one of the
segments of the calyx. A similar circumstance has been observed in Petunia
violacea by Morren. Duchartre describes an instance wherein one of the
outer sepals of Cattleya Forbesii was adherent to the labellum.[34]
Adhesion of the stamens to the petals is of common occurrence under
natural circumstances. Cassini has described a malformation of Centaurea
collina, in which two of the five stamens were completely grafted with the
corolla, the three others remaining perfectly free. Adhesion of the petals to
the column is not of infrequent occurrence among Orchids. I have observed
cases of the adhesion of the segments of the perianth to the stamen in
Ophrys aranifera, Odontoglossum, sp. &c. It is the ordinary condition in
Gongora and some other genera. I have seen it also in Lilium lancifolium.
Some forms of Crocus, occasionally met with, present a very singular
appearance, owing to the adhesion of the stamens to the outer segments of
the perianth, the former, moreover, being partially petaloid in aspect. M. de
la Vaud[35] speaks of a similar union in Tigridia pavonia. Morren[36]
describes a malformation of Fuchsia wherein the petals were so completely
adherent to the stamens, that the former were dragged out of their ordinary
Page 77
position, so as to become opposite to the sepals; the fusion was here so
complete that, no trace of it could be seen externally. It should be remarked
that it was the outer series of stamens that were thus fused.[37]
Fig. 12.—Crocus. Adhesion of petaloid stamens to perianth.
Adhesion of stamens to pistils.—The stamens also may be united to the
pistils, as in gynandrous plants. Moquin speaks of such a case in a
Scabious; M. Clos in Verbascum australe.[38] I have seen cases of the same
kind in the Wallflower, Cowslip (Primula veris), Tulip, Orange, in the
garden Azalea and other plants.
Miscellaneous adhesions.—Sometimes organs, comparatively speaking,
widely separated one from the other, become united together. Miquel has
recorded the union of a stigma with the middle lobe of the lower lip of the
corolla of Salvia pratensis.[39] In the accompanying figure [fig. 13], taken
complete that, no trace of it could be seen externally. It should be remarked
that it was the outer series of stamens that were thus fused.[37]
Fig. 12.—Crocus. Adhesion of petaloid stamens to perianth.
Adhesion of stamens to pistils.—The stamens also may be united to the
pistils, as in gynandrous plants. Moquin speaks of such a case in a
Scabious; M. Clos in Verbascum australe.[38] I have seen cases of the same
kind in the Wallflower, Cowslip (Primula veris), Tulip, Orange, in the
garden Azalea and other plants.
Miscellaneous adhesions.—Sometimes organs, comparatively speaking,
widely separated one from the other, become united together. Miquel has
recorded the union of a stigma with the middle lobe of the lower lip of the
corolla of Salvia pratensis.[39] In the accompanying figure [fig. 13], taken
Page 78
from a double wallflower, there is shown an adhesion between a petal and
an open carpel on the one side, and a stamen on the other.
Moquin speaks of some pears, which were united, at an early stage, with
one or two small leaves borne by the peduncle and grafted to the fruit by the
whole of their upper surface. As the pear increased in size the leaves
became detached from it, leaving on the surface of the fruit an impression
of the same form as the leaf, and differing in colour from the rest of the
surface of the fruit. Traces of the principal nerves were seen on the pear.
Fig. 13.—Cheiranthus cheiri. Adhesion of petal to stamen and open carpel.
It is curious to notice how very rare it is for the calyx to adhere to the ovary
in flowers where that organ is normally superior. The "calyx inferus" seems
scarcely ever to become "calyx superus," while, on the other hand, the
"calyx normaliter superus" frequently becomes inferior from detachment
from, or from want of union with the surface of the ovary.
an open carpel on the one side, and a stamen on the other.
Moquin speaks of some pears, which were united, at an early stage, with
one or two small leaves borne by the peduncle and grafted to the fruit by the
whole of their upper surface. As the pear increased in size the leaves
became detached from it, leaving on the surface of the fruit an impression
of the same form as the leaf, and differing in colour from the rest of the
surface of the fruit. Traces of the principal nerves were seen on the pear.
Fig. 13.—Cheiranthus cheiri. Adhesion of petal to stamen and open carpel.
It is curious to notice how very rare it is for the calyx to adhere to the ovary
in flowers where that organ is normally superior. The "calyx inferus" seems
scarcely ever to become "calyx superus," while, on the other hand, the
"calyx normaliter superus" frequently becomes inferior from detachment
from, or from want of union with the surface of the ovary.
Page 79
Adhesion of fruit to branch.—Of this Mr. Berkeley[40] cites an instance in
a vegetable marrow (Cucumis), where a female flower had become
confluent with the branch, at whose base it was placed, and also with two or
more flowers at the upper part of the same branch, so as to make an oblique
scar running down from the apex of the fruit to the branch.
Synanthy.—Adhesion of two or more flowers takes place in various ways;
sometimes merely the stalks are united together, so that we have a single
peduncle, bearing at its extremity two flowers placed in approximation very
slightly adherent one to the other. In this manner I have seen three flowers
of the vegetable marrow on a common stalk, the flowers themselves being
only united at the extreme base. Occasionally cases may be met with
wherein the pedicels of a stalked flower become adherent to the side of a
sessile flower. I have noticed this commonly in Umbelliferæ. Union of this
kind occurs frequently in the common cornel (Cornus), wherein one of the
lower flowers becomes adherent to one of the upper ones. In De Candolle's
'Organographie Végétale,' Plates 14 and 15, are figured cases of fusion of
the flower stems of the Hyacinth and of a Centaurea. In other cases the
union involves not only the stalk but the flowers themselves; thus fusion of
the flowers is a common accompaniment of fasciation, as was the case in
the Campanula figured in the cut (fig. 14).
a vegetable marrow (Cucumis), where a female flower had become
confluent with the branch, at whose base it was placed, and also with two or
more flowers at the upper part of the same branch, so as to make an oblique
scar running down from the apex of the fruit to the branch.
Synanthy.—Adhesion of two or more flowers takes place in various ways;
sometimes merely the stalks are united together, so that we have a single
peduncle, bearing at its extremity two flowers placed in approximation very
slightly adherent one to the other. In this manner I have seen three flowers
of the vegetable marrow on a common stalk, the flowers themselves being
only united at the extreme base. Occasionally cases may be met with
wherein the pedicels of a stalked flower become adherent to the side of a
sessile flower. I have noticed this commonly in Umbelliferæ. Union of this
kind occurs frequently in the common cornel (Cornus), wherein one of the
lower flowers becomes adherent to one of the upper ones. In De Candolle's
'Organographie Végétale,' Plates 14 and 15, are figured cases of fusion of
the flower stems of the Hyacinth and of a Centaurea. In other cases the
union involves not only the stalk but the flowers themselves; thus fusion of
the flowers is a common accompaniment of fasciation, as was the case in
the Campanula figured in the cut (fig. 14).
Page 80
Fig. 14.—Synanthic flowers of Campanula medium.
Synanthy may take place without much derangement of the structure of
either flower, or the union may be attended with abortion or suppression of
some of the parts of one or both flowers. Occasionally this union is carried
to such an extent that a bloom appears to be single, when it is, in reality,
composed of two or more, the parts of which have become not only fused,
but, as it were, thrust into and completely incorporated one with another,
and in such a manner as to occupy the place of some parts of the flower
which have been suppressed. It must not be overlooked that this adhesion of
one flower to another is a very common occurrence under natural
circumstances, as in Lonicera, in the common tomato, in Pomax,
Opercularia, Symphyomyrtus, &c., while the large size of some of the
cultivated sunflowers is in like manner due to the union of two or more
flower-heads.
One of the simplest instances of synanthy is that mentioned by M.
Duchartre,[41] in which two flowers of a hyacinth were united together
simply by means of two segments of the perianth one from each flower. A
Synanthy may take place without much derangement of the structure of
either flower, or the union may be attended with abortion or suppression of
some of the parts of one or both flowers. Occasionally this union is carried
to such an extent that a bloom appears to be single, when it is, in reality,
composed of two or more, the parts of which have become not only fused,
but, as it were, thrust into and completely incorporated one with another,
and in such a manner as to occupy the place of some parts of the flower
which have been suppressed. It must not be overlooked that this adhesion of
one flower to another is a very common occurrence under natural
circumstances, as in Lonicera, in the common tomato, in Pomax,
Opercularia, Symphyomyrtus, &c., while the large size of some of the
cultivated sunflowers is in like manner due to the union of two or more
flower-heads.
One of the simplest instances of synanthy is that mentioned by M.
Duchartre,[41] in which two flowers of a hyacinth were united together
simply by means of two segments of the perianth one from each flower. A
Page 81
similar occurrence has been cited by M. Gay in Narcissus chrysanthus. In
like manner the blossoms of Fuchsias or Loniceras occasionally become
adherent merely by their surface, without involving any other change in the
conformation of the flowers. M. Maugin alludes to a case of this kind in
Aristolochia Clematitis.[42]
But it is more usual for some of the organs to be suppressed, so that the
number of existing parts is less than would be the case in two or more
uncombined flowers. A few illustrations will exemplify this. In two flowers
of Matthiola incana, that I observed to be joined together, there were eight
sepals, eight petals, and ten perfect stamens, eight long and two short,
instead of twelve. Closer examination showed that the point of union
between the two flowers occurred just where, under ordinary circumstances,
the two short stamens would be. In this instance but little suppression had
occurred. In similar flowers of Narcissus incomparabilis I remarked a ten-
parted perianth, ten stamens within a single cup, two styles, and a five-
celled ovary. Here, then, it would appear that two segments of the perianth,
two stamens, and one carpel were suppressed. In a Polyanthus there were
nine sepals, nine petals, nine stamens, and a double ovary.
like manner the blossoms of Fuchsias or Loniceras occasionally become
adherent merely by their surface, without involving any other change in the
conformation of the flowers. M. Maugin alludes to a case of this kind in
Aristolochia Clematitis.[42]
But it is more usual for some of the organs to be suppressed, so that the
number of existing parts is less than would be the case in two or more
uncombined flowers. A few illustrations will exemplify this. In two flowers
of Matthiola incana, that I observed to be joined together, there were eight
sepals, eight petals, and ten perfect stamens, eight long and two short,
instead of twelve. Closer examination showed that the point of union
between the two flowers occurred just where, under ordinary circumstances,
the two short stamens would be. In this instance but little suppression had
occurred. In similar flowers of Narcissus incomparabilis I remarked a ten-
parted perianth, ten stamens within a single cup, two styles, and a five-
celled ovary. Here, then, it would appear that two segments of the perianth,
two stamens, and one carpel were suppressed. In a Polyanthus there were
nine sepals, nine petals, nine stamens, and a double ovary.
Page 82
Fig. 15.—Union of three flowers of Calanthe vestita.
Page 83
Fig. 16.—Shows the abortion of the central spur in synanthic flowers of Calanthe vestita.
As an illustration of a more complicated nature reference may be made to
three flowers of Aconitum Napellus, figured by A. de Chamisso, 'Linnæa,'
vol. vii, 1832, p. 205, tab. vii, figs. 1, 2. In this specimen the two outer
blossoms had each four sepals present, namely, the upper hooded one, one
of the lateral sepals, and both of the inferior ones; the central flower had
only the upper sepal and one other, probably one of the lower sepals; thus
there were but ten sepals instead of fifteen. The nectary-like petals, the
stamens, and pistils were all present in the lateral flowers, but were
completely suppressed in the middle one. A less degree of suppression was
exemplified in a triple flower of Calanthe vestita sent me by Dr. Moore, of
Glasnevin, in which all the parts usually existing in three separate flowers
were to be found, with the exception of the spur belonging to the labellum
of the middle flower (figs. 15, 16).
One of the most common malformations in the Foxglove (Digitalis) results
from the fusion of several of the terminal flowers into one. In these cases
the number of parts is very variable in different instances; the sepals are
more or less blended together, and the corollas as well as the stamens are
usually free and distinct, the latter often of equal length, so that the
blossom, although truly complex, is, as to its external form, less irregular
than under natural circumstances. The centre of these flowers is occupied
by a two to five-celled pistil, between the carpels of which, not
unfrequently, the stem of the plant projects, bearing on its sides bracts and
rudimentary flowers. (See Prolification.) An instance of this nature is
figured in the 'Gardeners' Chronicle,' 1850, p. 435, from which the cut (fig.
17) is borrowed.
As an illustration of a more complicated nature reference may be made to
three flowers of Aconitum Napellus, figured by A. de Chamisso, 'Linnæa,'
vol. vii, 1832, p. 205, tab. vii, figs. 1, 2. In this specimen the two outer
blossoms had each four sepals present, namely, the upper hooded one, one
of the lateral sepals, and both of the inferior ones; the central flower had
only the upper sepal and one other, probably one of the lower sepals; thus
there were but ten sepals instead of fifteen. The nectary-like petals, the
stamens, and pistils were all present in the lateral flowers, but were
completely suppressed in the middle one. A less degree of suppression was
exemplified in a triple flower of Calanthe vestita sent me by Dr. Moore, of
Glasnevin, in which all the parts usually existing in three separate flowers
were to be found, with the exception of the spur belonging to the labellum
of the middle flower (figs. 15, 16).
One of the most common malformations in the Foxglove (Digitalis) results
from the fusion of several of the terminal flowers into one. In these cases
the number of parts is very variable in different instances; the sepals are
more or less blended together, and the corollas as well as the stamens are
usually free and distinct, the latter often of equal length, so that the
blossom, although truly complex, is, as to its external form, less irregular
than under natural circumstances. The centre of these flowers is occupied
by a two to five-celled pistil, between the carpels of which, not
unfrequently, the stem of the plant projects, bearing on its sides bracts and
rudimentary flowers. (See Prolification.) An instance of this nature is
figured in the 'Gardeners' Chronicle,' 1850, p. 435, from which the cut (fig.
17) is borrowed.
Page 84
Fig. 17.—Synanthy and other changes in a Foxglove.
One of the most singular recorded instances of changes connected with
fusion of the flowers is that cited by Reinsch,[43] where two female flowers
of Salix cinerea were so united with a male one as to produce an
hermaphrodite blossom.
It follows, from what has been said, that the number of parts that are met
with in these fused flowers varies according to the number of blossoms and
of the organs which have been suppressed. Comparatively rarely do we find
all the organs present; but when two flowers are united together we find
every possible variety between the number of parts naturally belonging to
the two flowers and that belonging to a single one. Sometimes instances are
met with wherein the calyx does not present the normal number of parts,
while the other parts of the flower are in excess. I have seen in a
Calceolaria a single calyx, with the ordinary number of sepals, enclosing
One of the most singular recorded instances of changes connected with
fusion of the flowers is that cited by Reinsch,[43] where two female flowers
of Salix cinerea were so united with a male one as to produce an
hermaphrodite blossom.
It follows, from what has been said, that the number of parts that are met
with in these fused flowers varies according to the number of blossoms and
of the organs which have been suppressed. Comparatively rarely do we find
all the organs present; but when two flowers are united together we find
every possible variety between the number of parts naturally belonging to
the two flowers and that belonging to a single one. Sometimes instances are
met with wherein the calyx does not present the normal number of parts,
while the other parts of the flower are in excess. I have seen in a
Calceolaria a single calyx, with the ordinary number of sepals, enclosing
Page 85
two corollas, adherent simply by their upper lips, and containing stamens
and pistils in the usual way. In this instance, then, the sepals of one flower
must have been suppressed, while no such suppression took place in the
other parts of the flower.
Professor Charles Morren paid special attention to the various methods in
which the flowers of Calceolarias may become fused, and to the
complications that ensue from the suppression of some parts, the complete
amalgamation of others, &c. Referring the reader to the Belgian savant's
papers for the full details of the changes observed, it is only necessary to
allude to a few of the most salient features.
Fig. 18.—Synanthic flowers of Calceolaria in which, with two upper lips, there was but a single
lower one.
Sometimes the upper lips of two flowers are fused into one, the two lower
remaining distinct. In other cases, the upper lip disappears altogether, while
and pistils in the usual way. In this instance, then, the sepals of one flower
must have been suppressed, while no such suppression took place in the
other parts of the flower.
Professor Charles Morren paid special attention to the various methods in
which the flowers of Calceolarias may become fused, and to the
complications that ensue from the suppression of some parts, the complete
amalgamation of others, &c. Referring the reader to the Belgian savant's
papers for the full details of the changes observed, it is only necessary to
allude to a few of the most salient features.
Fig. 18.—Synanthic flowers of Calceolaria in which, with two upper lips, there was but a single
lower one.
Sometimes the upper lips of two flowers are fused into one, the two lower
remaining distinct. In other cases, the upper lip disappears altogether, while
Page 86
there are two lower lips placed opposite one another;, of the stamens,
sometimes the outermost, at other times the innermost disappear.[44]
Occasionally there appears to be, as it were, a transference of the parts of
one flower to another. One of the simplest and most intelligible cases of this
kind is recorded by Wigand in the 'Flora' for 1856, in a compound flower of
Polygonatum anceps, in which within a twelve-parted perianth there were
twelve stamens and two pistils, one four-celled, the other two-celled; hence
it would appear as if a carpel belonging to one flower had become united to
those constituting the pistil of the adjacent one. Among Orchids this fusion
of some of the elements of different flowers, together with the suppression
of others, is carried to such an extent as to render the real structure difficult
to decipher. Sometimes flowers of Ophrys aranifera, at first sight seeming
normal as to the number, and almost so as regards the arrangement of their
parts, have yet, on examination, proved to be the result of a confluence of
two flowers. Mr. Moggridge has observed similar phenomena in the same
species at Mentone.
Sometimes the fusion affects flowers belonging to different branches of the
same inflorescence, as in Centranthus ruber, described by Buchenau,
'Flora,' 1857, p. 293, and even a blossom of one generation of axes may be
united with a flower belonging to another generation. Thus M. Michalet[45]
speaks of a case wherein the terminal flower of Betonica alopecuros was
affected with Peloria, and fused with an adjacent one belonging to a
secondary axis of inflorescence, and not yet expanded. This latter flower
had no calyx, but in its place were three bracts, surrounding the corolla; this
again was united to the calyx of the terminal bloom in a most singular
manner, the limb of the corolla and that of the calyx being so joined one to
the other as to form but a single tube. It is not uncommon, as has been
before stated, to find two corollas enclosed within one calyx, but this is
sometimes the outermost, at other times the innermost disappear.[44]
Occasionally there appears to be, as it were, a transference of the parts of
one flower to another. One of the simplest and most intelligible cases of this
kind is recorded by Wigand in the 'Flora' for 1856, in a compound flower of
Polygonatum anceps, in which within a twelve-parted perianth there were
twelve stamens and two pistils, one four-celled, the other two-celled; hence
it would appear as if a carpel belonging to one flower had become united to
those constituting the pistil of the adjacent one. Among Orchids this fusion
of some of the elements of different flowers, together with the suppression
of others, is carried to such an extent as to render the real structure difficult
to decipher. Sometimes flowers of Ophrys aranifera, at first sight seeming
normal as to the number, and almost so as regards the arrangement of their
parts, have yet, on examination, proved to be the result of a confluence of
two flowers. Mr. Moggridge has observed similar phenomena in the same
species at Mentone.
Sometimes the fusion affects flowers belonging to different branches of the
same inflorescence, as in Centranthus ruber, described by Buchenau,
'Flora,' 1857, p. 293, and even a blossom of one generation of axes may be
united with a flower belonging to another generation. Thus M. Michalet[45]
speaks of a case wherein the terminal flower of Betonica alopecuros was
affected with Peloria, and fused with an adjacent one belonging to a
secondary axis of inflorescence, and not yet expanded. This latter flower
had no calyx, but in its place were three bracts, surrounding the corolla; this
again was united to the calyx of the terminal bloom in a most singular
manner, the limb of the corolla and that of the calyx being so joined one to
the other as to form but a single tube. It is not uncommon, as has been
before stated, to find two corollas enclosed within one calyx, but this is
Page 87
probably the only recorded instance of the fusion of the calyx and corolla of
two different flowers belonging to two different axes.
From the preceding details, as well as from others which it is not necessary
to give in this place, it would appear that synanthy is more liable to occur
where the flowers are naturally crowded together[46] than where they are
remote; so too, the upper or younger portions of the inflorescence are those
most subject to this change. In like manner the derangements consequent on
the coalescence of flowers are often more grave in the central organs, which
are most exposed to pressure, and have the least opportunities of resisting
the effects of that agency, than they are in the outer portions of the flowers
where growth is less restricted.
Morren in his papers on synanthic Calceolarias, before referred to,
considers that the direction in which fusion acts is centripetal, e.g. from the
circumference towards the centre of the flower, thus reversing the natural
order of things. He considers that there is a radical antagonism between the
normal organizing forces and the teratological disorganizing forces, and
explains in this way the frequent sterility of monsters from an imperfect
formation of stamens, or pistils, or both.
The greater tendency in synanthic flowers of parts of one whorl to adhere to
the corresponding organs in another flower has often been remarked,
though the dislocation of parts may be so great as to prevent this from being
carried out in all cases. It appears also that synanthy is more frequently met
with among flowers which have an inferior ovary than in those in which the
relative position of the organ in question is reversed. This remark applies
particularly to individual cases; the proportion as regards the genera may
not be so large. The explanation of this must of course depend on the
circumstances of each particular case; and it would be wrong to attempt to
lay down a general rule, when organogenists have not yet fully decided in
two different flowers belonging to two different axes.
From the preceding details, as well as from others which it is not necessary
to give in this place, it would appear that synanthy is more liable to occur
where the flowers are naturally crowded together[46] than where they are
remote; so too, the upper or younger portions of the inflorescence are those
most subject to this change. In like manner the derangements consequent on
the coalescence of flowers are often more grave in the central organs, which
are most exposed to pressure, and have the least opportunities of resisting
the effects of that agency, than they are in the outer portions of the flowers
where growth is less restricted.
Morren in his papers on synanthic Calceolarias, before referred to,
considers that the direction in which fusion acts is centripetal, e.g. from the
circumference towards the centre of the flower, thus reversing the natural
order of things. He considers that there is a radical antagonism between the
normal organizing forces and the teratological disorganizing forces, and
explains in this way the frequent sterility of monsters from an imperfect
formation of stamens, or pistils, or both.
The greater tendency in synanthic flowers of parts of one whorl to adhere to
the corresponding organs in another flower has often been remarked,
though the dislocation of parts may be so great as to prevent this from being
carried out in all cases. It appears also that synanthy is more frequently met
with among flowers which have an inferior ovary than in those in which the
relative position of the organ in question is reversed. This remark applies
particularly to individual cases; the proportion as regards the genera may
not be so large. The explanation of this must of course depend on the
circumstances of each particular case; and it would be wrong to attempt to
lay down a general rule, when organogenists have not yet fully decided in
Page 88
what plants the inferior ovary is an axial structure, and in what others the
appearance is due to the adhesion of the base of the calyx to the carpels.
The list which follows is not intended as a complete one, but it may serve to
show what plants are more particularly subject to this anomaly; the *
indicates unusual frequency of occurrence, the ! signifies that the writer has
himself seen instances in the plants named. Many of the recorded cases of
Synanthy are really cases of adhesion of the inflorescence rather than of the
flowers.
Ranunculus Lingua.
bulbosus!
Aconitum Napellus.
Delphinium sp.!
Matthiola incana!
Arabis sagittata.
Silene sp.
Reseda odorata!
Vitis vinifera.
Citrus aurantium.
*Fuchsia var. hort.!
Œnothera sp.
Saxifraga sp.
Podalyria myrtillifolia.
Prunus Armeniaca.
spinosa.
Pyrus Malus.
Persica vulgaris.
Cratægus monogyna.
Robinia pseudacacia.
Gleditschia triacanthos.
Syringa persica.
Cornus sanguinea.
Viburnum sp.
*Lonicera sp. plur!
Centranthus ruber!
Valantia cruciata.
appearance is due to the adhesion of the base of the calyx to the carpels.
The list which follows is not intended as a complete one, but it may serve to
show what plants are more particularly subject to this anomaly; the *
indicates unusual frequency of occurrence, the ! signifies that the writer has
himself seen instances in the plants named. Many of the recorded cases of
Synanthy are really cases of adhesion of the inflorescence rather than of the
flowers.
Ranunculus Lingua.
bulbosus!
Aconitum Napellus.
Delphinium sp.!
Matthiola incana!
Arabis sagittata.
Silene sp.
Reseda odorata!
Vitis vinifera.
Citrus aurantium.
*Fuchsia var. hort.!
Œnothera sp.
Saxifraga sp.
Podalyria myrtillifolia.
Prunus Armeniaca.
spinosa.
Pyrus Malus.
Persica vulgaris.
Cratægus monogyna.
Robinia pseudacacia.
Gleditschia triacanthos.
Syringa persica.
Cornus sanguinea.
Viburnum sp.
*Lonicera sp. plur!
Centranthus ruber!
Valantia cruciata.
Page 89
Centaurea moschata.
Jacea.
Zinnia elegans.
Zinnia revoluta.
Helianthus sp.!
Spilanthes oleracea.
Dahlia.
*Leontodon Taraxacum!
Senecio Doria.
Cichorium Intybus.
Lactuca sativa.
Anthemis retusa.
*Campanula medium!
persicifolia.
Azalea indica!
Vinca minor.
Atropa Belladonna.
*Solanum Lycopersicum!
*Petunia violacea!
Galeopsis ochroleuca.
Betonica alopecuros.
*Digitalis purpurea!
*Antirrhinum majus!
*Linaria purpurea!
*Pedicularis sylvatica!
*Calceolaria var. hort.!
Scrophularia nodosa.
Salpiglossis straminea.
Streptocarpus Rexii.
*Gesnera var. hort.!
Æschynanthus sp.!
Thyrsacanthus rutilans!
Anagallis collina.
*Primula veris!
Auricula.
*Primula acaulis, var. umbellata!
elatior?
*sinensis!
Aristolochia Clematitis.
Jacea.
Zinnia elegans.
Zinnia revoluta.
Helianthus sp.!
Spilanthes oleracea.
Dahlia.
*Leontodon Taraxacum!
Senecio Doria.
Cichorium Intybus.
Lactuca sativa.
Anthemis retusa.
*Campanula medium!
persicifolia.
Azalea indica!
Vinca minor.
Atropa Belladonna.
*Solanum Lycopersicum!
*Petunia violacea!
Galeopsis ochroleuca.
Betonica alopecuros.
*Digitalis purpurea!
*Antirrhinum majus!
*Linaria purpurea!
*Pedicularis sylvatica!
*Calceolaria var. hort.!
Scrophularia nodosa.
Salpiglossis straminea.
Streptocarpus Rexii.
*Gesnera var. hort.!
Æschynanthus sp.!
Thyrsacanthus rutilans!
Anagallis collina.
*Primula veris!
Auricula.
*Primula acaulis, var. umbellata!
elatior?
*sinensis!
Aristolochia Clematitis.
Page 90
Blitum sp.
Chenopodium sp.
Rumex sp.
Salix cinerea.
*Hyacinthus orientalis!
Lilium bulbiferum!
croceum, et sp. alix, pl.
Tulipa, sp.
Polygonatum anceps.
Fritillaria imperalis!
Agave americana.
Iris versicolor.
sambucina.
Crocus, sp.
Colchicum autumnale.
Narcissus incomparabilis!
Tazetta.
biflorus.
chrysanthus.
*Ophrys aranifera!
Calanthe vestita!
Oncidium bicolor.
ornithorhyncum.
&c. &c.
In addition to the works before cited, additional information on this subject may be gained
from the following:—Jaeger, 'Missbilld.,' p. 92. v. Schlechtend, 'Bot. Zeit.,' 1856, Robinia.
Weber, 'Verhandl. Nat. Hist. Vereins. Preuss. Rheinl.,' 1849, p. 290, Primula. Hincks, 'Rep.
Brit. Assoc. Newcastle,' 1838, Salpiglossis. Clos, 'Mém. Acad. Toulouse,' vol. vi, 1862,
Anagallis. Wigand, 'Flora,' 1856, tab. 8, Pedicularis. Henfrey, 'Botan. Gazette,' i, p. 280,
Reseda. P. Reinsch, 'Flora,' 1860, tab. 7, Petasites. Weber, Verhandl. Nat. Hist. Vereins. f.d.
Preuss. Rheinl. u. Westphal.,' 1860, p. 332, tabs. 6 et 7, Prunus, Persica, Campanula,
Taraxacum, Saxifraga, Silene, Hyacinthus, &c. Miquel, 'Linnæa,' xi, p. 423, Colchicum.
Michel, 'Traité du Citronnier,' tab. 6, Citrus.
Syncarpy.—In the preceding section it has been shown that the carpels, like
other parts of the flower, are subject to be united together. This union may
either take place between the carpels of a single flower or between the
pistils of different flowers. In the latter case the other floral whorls are
Chenopodium sp.
Rumex sp.
Salix cinerea.
*Hyacinthus orientalis!
Lilium bulbiferum!
croceum, et sp. alix, pl.
Tulipa, sp.
Polygonatum anceps.
Fritillaria imperalis!
Agave americana.
Iris versicolor.
sambucina.
Crocus, sp.
Colchicum autumnale.
Narcissus incomparabilis!
Tazetta.
biflorus.
chrysanthus.
*Ophrys aranifera!
Calanthe vestita!
Oncidium bicolor.
ornithorhyncum.
&c. &c.
In addition to the works before cited, additional information on this subject may be gained
from the following:—Jaeger, 'Missbilld.,' p. 92. v. Schlechtend, 'Bot. Zeit.,' 1856, Robinia.
Weber, 'Verhandl. Nat. Hist. Vereins. Preuss. Rheinl.,' 1849, p. 290, Primula. Hincks, 'Rep.
Brit. Assoc. Newcastle,' 1838, Salpiglossis. Clos, 'Mém. Acad. Toulouse,' vol. vi, 1862,
Anagallis. Wigand, 'Flora,' 1856, tab. 8, Pedicularis. Henfrey, 'Botan. Gazette,' i, p. 280,
Reseda. P. Reinsch, 'Flora,' 1860, tab. 7, Petasites. Weber, Verhandl. Nat. Hist. Vereins. f.d.
Preuss. Rheinl. u. Westphal.,' 1860, p. 332, tabs. 6 et 7, Prunus, Persica, Campanula,
Taraxacum, Saxifraga, Silene, Hyacinthus, &c. Miquel, 'Linnæa,' xi, p. 423, Colchicum.
Michel, 'Traité du Citronnier,' tab. 6, Citrus.
Syncarpy.—In the preceding section it has been shown that the carpels, like
other parts of the flower, are subject to be united together. This union may
either take place between the carpels of a single flower or between the
pistils of different flowers. In the latter case the other floral whorls are
Page 91
generally more or less altered. Where, however, the ovary is, as it is called,
inferior, it may happen that the pistils of different flowers may coalesce
more or less without much alteration in the other parts of the flower, as
happens normally in many Caprifoliaceæ, Rubiaceæ, &c. &c. In some of
these cases it must be remembered that the real structure of the apparent
fruit is not made out beyond dispute, the main points of controversy being
as to what, if any, share the dilated fruit-stalk or axis takes in the formation
of such organs. Again, it will be borne in mind that in some cases the so-
called fruit is made up of a number of flowers all fused together, as in the
Mulberry or the Pineapple, in which plants what is, in ordinary language,
called the fruit really consists of the whole mass of flowers constituting the
inflorescence fused together. Union of the fruits may also in some cases
take place between the carpels after the fall of the other floral whorls,
particularly when the outer layers of the pericarp assume a succulent
condition, so that under the general head of syncarpy really different
conditions are almost necessarily grouped together, and, in seeking to
investigate the causes of the phenomenon, the particular circumstances of
each individual case must be taken into account. Syncarpy takes place in
various degrees; sometimes only the stalks are joined; at other times the
whole extent of the fruit, as in cherries, &c. This peculiarity did not escape
the observant mind of Shakespeare—
"A double cherry seeming parted.
But yet a union in partition,
Two lovely berries moulded on one stem."
'Midsummer Night's Dream,' act iii, sc. 2.
A similar union has been observed in peaches, gooseberries, gourds,
melons, and a great many other fruits. In the Barbarossa grape I have
frequently seen a fusion of two, three, four or more berries quite at the end
of the bunch, so that the clusters were terminated by a compound grape.
Seringe has remarked sometimes two, sometimes three, fruits of
inferior, it may happen that the pistils of different flowers may coalesce
more or less without much alteration in the other parts of the flower, as
happens normally in many Caprifoliaceæ, Rubiaceæ, &c. &c. In some of
these cases it must be remembered that the real structure of the apparent
fruit is not made out beyond dispute, the main points of controversy being
as to what, if any, share the dilated fruit-stalk or axis takes in the formation
of such organs. Again, it will be borne in mind that in some cases the so-
called fruit is made up of a number of flowers all fused together, as in the
Mulberry or the Pineapple, in which plants what is, in ordinary language,
called the fruit really consists of the whole mass of flowers constituting the
inflorescence fused together. Union of the fruits may also in some cases
take place between the carpels after the fall of the other floral whorls,
particularly when the outer layers of the pericarp assume a succulent
condition, so that under the general head of syncarpy really different
conditions are almost necessarily grouped together, and, in seeking to
investigate the causes of the phenomenon, the particular circumstances of
each individual case must be taken into account. Syncarpy takes place in
various degrees; sometimes only the stalks are joined; at other times the
whole extent of the fruit, as in cherries, &c. This peculiarity did not escape
the observant mind of Shakespeare—
"A double cherry seeming parted.
But yet a union in partition,
Two lovely berries moulded on one stem."
'Midsummer Night's Dream,' act iii, sc. 2.
A similar union has been observed in peaches, gooseberries, gourds,
melons, and a great many other fruits. In the Barbarossa grape I have
frequently seen a fusion of two, three, four or more berries quite at the end
of the bunch, so that the clusters were terminated by a compound grape.
Seringe has remarked sometimes two, sometimes three, fruits of
Page 92
Ranunculus tripartitus soldered together. He has also seen three melons
similarly joined.[47] Turpin mentions having seen a complete union between
the three smooth and leathery pericarps which are naturally separate and
enclosed within the spiny cupule of the chestnut.[48] Poiteau and Turpin
have figured and described in their treatise on fruit trees, under the name of
Néfle de Correa, four or five medlars, joined together and surmounted by
all the persistent leaflets of the calyces.[49]
A very remarkable example of Syncarpy has been recorded by E. Kœnig in
which nine strawberries were borne on one stem (Fragaria botryformis),[50]
and a similar malformation has been observed in the Pineapple.
When two fruits are united together they may be of about equal size, while
in other cases one of the two is much smaller than the other. This was the
case in two cucumbers given to me by Mr. James Salter. These were united
together along their whole length excepting at the very tips; the upper one
of the two was much larger than the lower, and contained three cells, the
lower fruit was one-celled by suppression. Both fruits were curved, the
curvature being evidently due to the more rapid growth of the upper as
compared with the lower one.
similarly joined.[47] Turpin mentions having seen a complete union between
the three smooth and leathery pericarps which are naturally separate and
enclosed within the spiny cupule of the chestnut.[48] Poiteau and Turpin
have figured and described in their treatise on fruit trees, under the name of
Néfle de Correa, four or five medlars, joined together and surmounted by
all the persistent leaflets of the calyces.[49]
A very remarkable example of Syncarpy has been recorded by E. Kœnig in
which nine strawberries were borne on one stem (Fragaria botryformis),[50]
and a similar malformation has been observed in the Pineapple.
When two fruits are united together they may be of about equal size, while
in other cases one of the two is much smaller than the other. This was the
case in two cucumbers given to me by Mr. James Salter. These were united
together along their whole length excepting at the very tips; the upper one
of the two was much larger than the lower, and contained three cells, the
lower fruit was one-celled by suppression. Both fruits were curved, the
curvature being evidently due to the more rapid growth of the upper as
compared with the lower one.
Page 93
Fig. 19.—Adhesion of two apples.
Fig. 20.—Section of united apples.
In many of these cases, where the fruits are united by their bases, the
summits become separated one from the other, so as to resemble the letter
V. Such divergence is of frequent occurrence where fruits are united by their
stalks, because, as growth goes on, the tendency must necessarily be
towards separation and divergence of the tips of the fruit.
Fig. 20.—Section of united apples.
In many of these cases, where the fruits are united by their bases, the
summits become separated one from the other, so as to resemble the letter
V. Such divergence is of frequent occurrence where fruits are united by their
stalks, because, as growth goes on, the tendency must necessarily be
towards separation and divergence of the tips of the fruit.
Page 94
In some cases of Syncarpy the fusion and interpenetration of the carpels is
carried to such an extent that it is very difficult to trace on the outer surface
the lines of union. The fruit in these cases resembles a single one of much
larger size than usual. Moquin mentions a double apple in which the
connection was so close that the fruit was not very different in form from
what is customary, and a similar thing happens with the tomato. In the case
of stone-fruits it sometimes happens, not only that the outer portions are
adherent, but that the stones are so likewise.
M. Rœper has observed two apples grafted together, one of which had its
stalk broken, and seemed evidently borne and nourished by the other apple;
[51] and a similar occurrence happens not infrequently in the cucumber.
Moquin has seen three united cherries having only a single stalk jointed to
the central fruit, the lateral cherries having each a slight depression or
cicatrix marking the situation of the suppressed stalks. Schlotterbec has
figured three apples presenting precisely similar appearances.[52]
Fusion of two or more nuts (Corylus) is not uncommon; I have seen as
many as five so united.[53] In these cases the fruits may be united together
in a ring or in linear series.
In some Leguminosæ, contrary to the general rule in the order, more than
one carpel is found; thus peas, French beans, and other similar plants, are
occasionally met with having two or more pods within the same calyx, and
in Gleditschia triacanthos and Cæsalpinia digyna this is so commonly the
case as to be considered almost the normal state. (De Cand. 'Mem. Leg.,' pl.
2, fig. 6; pl. 3, fig. 2.) At times these carpels become fused together, and it
becomes difficult, when the traces of the flower have disappeared, to
ascertain whether these carpels were formed in one flower, or whether they
were the result of the fusion of several blossoms. I have seen an instance of
this kind in a plum in which there were two carpels in the same flower, the
carried to such an extent that it is very difficult to trace on the outer surface
the lines of union. The fruit in these cases resembles a single one of much
larger size than usual. Moquin mentions a double apple in which the
connection was so close that the fruit was not very different in form from
what is customary, and a similar thing happens with the tomato. In the case
of stone-fruits it sometimes happens, not only that the outer portions are
adherent, but that the stones are so likewise.
M. Rœper has observed two apples grafted together, one of which had its
stalk broken, and seemed evidently borne and nourished by the other apple;
[51] and a similar occurrence happens not infrequently in the cucumber.
Moquin has seen three united cherries having only a single stalk jointed to
the central fruit, the lateral cherries having each a slight depression or
cicatrix marking the situation of the suppressed stalks. Schlotterbec has
figured three apples presenting precisely similar appearances.[52]
Fusion of two or more nuts (Corylus) is not uncommon; I have seen as
many as five so united.[53] In these cases the fruits may be united together
in a ring or in linear series.
In some Leguminosæ, contrary to the general rule in the order, more than
one carpel is found; thus peas, French beans, and other similar plants, are
occasionally met with having two or more pods within the same calyx, and
in Gleditschia triacanthos and Cæsalpinia digyna this is so commonly the
case as to be considered almost the normal state. (De Cand. 'Mem. Leg.,' pl.
2, fig. 6; pl. 3, fig. 2.) At times these carpels become fused together, and it
becomes difficult, when the traces of the flower have disappeared, to
ascertain whether these carpels were formed in one flower, or whether they
were the result of the fusion of several blossoms. I have seen an instance of
this kind in a plum in which there were two carpels in the same flower, the
Page 95
one being partially fused to the other. The nature of such cases may usually
be determined by an inspection of the peduncle which shows no traces of
fusion. (See chapter on Multiplication.)
When, however, the fruits are sessile, and they become grafted together, the
kind of syncarpy is difficult to distinguish. It, may, nevertheless, be said as
a general rule that the union brought about by the approximation of two
fruits, after the fall of the floral whorls, is never so complete or so intimate
as that determined by synanthy; and also that in those cases where there are
supernumerary carpels in the flower, and those carpels become united
together, they are rarely so completely fused that their individuality is lost.
An analogous phenomenon takes place not uncommonly in mosses, the
spore capsules of which become united together in various ways and
degrees. Schimper[54] cites the following species as subject to this anomaly:
—Buxbaumia indusiata, Leskea sericea, Hypnum lutescens, Anomodon
alternatus, Clinacium dendroides, Bryum cæspititium, Brachythecium
plumosum, Mnium serratum, Splachnum vasculosum. It has also been
observed in Trichostomum rigidulum and Hypnum triquetrum.
In addition to the authorities already mentioned, the reader may consult Moquin-Tandon,
'El. Ter. Veg.,' p. 270. Turpin. 'Mém. greffe. Ann. Sc. Nat.,' ser. i, t. xxiv, p. 334. De
Candolle, 'Organ. Veget.,' t. i. Duhamel, 'Phys. des Arbres,' t. i, p. 304, tab. xiii, xiv. Weber.
'Verhandl. Nat. Hist. Vereina f. d. Preuss. Rheinl. u. Westphal.,' 1860, p. 332, tab. vi. et vii.
Synspermy, or Union of the Seeds.—Seeds may be united together in
various degrees, either by their integuments,[55] or by their inner parts. Such
union of the seeds, however, is of rare occurrence. It takes place normally,
to a slight extent, in certain cultivated forms of cotton, wherein the seeds
are aggregated together into a reniform mass, whence the term kidney
cotton. Union of the parts of the embryo is treated under another head (see
Synophty).
be determined by an inspection of the peduncle which shows no traces of
fusion. (See chapter on Multiplication.)
When, however, the fruits are sessile, and they become grafted together, the
kind of syncarpy is difficult to distinguish. It, may, nevertheless, be said as
a general rule that the union brought about by the approximation of two
fruits, after the fall of the floral whorls, is never so complete or so intimate
as that determined by synanthy; and also that in those cases where there are
supernumerary carpels in the flower, and those carpels become united
together, they are rarely so completely fused that their individuality is lost.
An analogous phenomenon takes place not uncommonly in mosses, the
spore capsules of which become united together in various ways and
degrees. Schimper[54] cites the following species as subject to this anomaly:
—Buxbaumia indusiata, Leskea sericea, Hypnum lutescens, Anomodon
alternatus, Clinacium dendroides, Bryum cæspititium, Brachythecium
plumosum, Mnium serratum, Splachnum vasculosum. It has also been
observed in Trichostomum rigidulum and Hypnum triquetrum.
In addition to the authorities already mentioned, the reader may consult Moquin-Tandon,
'El. Ter. Veg.,' p. 270. Turpin. 'Mém. greffe. Ann. Sc. Nat.,' ser. i, t. xxiv, p. 334. De
Candolle, 'Organ. Veget.,' t. i. Duhamel, 'Phys. des Arbres,' t. i, p. 304, tab. xiii, xiv. Weber.
'Verhandl. Nat. Hist. Vereina f. d. Preuss. Rheinl. u. Westphal.,' 1860, p. 332, tab. vi. et vii.
Synspermy, or Union of the Seeds.—Seeds may be united together in
various degrees, either by their integuments,[55] or by their inner parts. Such
union of the seeds, however, is of rare occurrence. It takes place normally,
to a slight extent, in certain cultivated forms of cotton, wherein the seeds
are aggregated together into a reniform mass, whence the term kidney
cotton. Union of the parts of the embryo is treated under another head (see
Synophty).
Page 96
Adhesion between the axes of different plants.—Under this head may be
classed the union that takes place between the stems, branches, or roots of
different plants of the same species, and that which occurs between
individuals of different species; the first is not very different in its nature
from cohesion of the branches of the same plant (figs. 21, 22). It finds its
parallel, under natural circumstances, among the lower cryptogams, in
which it often happens that several individual plants, originally distinct,
become inseparably blended together into one mass. In the gardening
operations of inarching, and to some extent in budding, this adhesion of
axis to axis occurs, the union taking place the more readily in proportion as
the contact between the younger growing portions of the two axes
respectively is close. The huge size of some trees has been, in some cases,
attributed to the adnation of different stems. This is said to be the case with
the famous plane trees of Bujukdere, near Constantinople, and in which
nine trunks are more or less united together.[56]
classed the union that takes place between the stems, branches, or roots of
different plants of the same species, and that which occurs between
individuals of different species; the first is not very different in its nature
from cohesion of the branches of the same plant (figs. 21, 22). It finds its
parallel, under natural circumstances, among the lower cryptogams, in
which it often happens that several individual plants, originally distinct,
become inseparably blended together into one mass. In the gardening
operations of inarching, and to some extent in budding, this adhesion of
axis to axis occurs, the union taking place the more readily in proportion as
the contact between the younger growing portions of the two axes
respectively is close. The huge size of some trees has been, in some cases,
attributed to the adnation of different stems. This is said to be the case with
the famous plane trees of Bujukdere, near Constantinople, and in which
nine trunks are more or less united together.[56]
Page 97
Fig. 21.—Adhesion of two distinct stems of oak, or possibly cohesion of branches of the same tree.
'Gard. Chron.,' 1846, p. 252.
A similar anastomosis may take place in the roots. Lindley cites a case
wherein two carrots, of the white Belgian and the red Surrey varieties
respectively, had grown so close to each other that each twisted half round
the other, so that they ultimately became soldered together; the most
singular thing with reference to this union was, that the red carrot (fig. 23,
b), with its small overgrown part above the junction, took the colour and
large dimensions of the white Belgian (d), which, in like manner, with its
larger head above the joining (a), took the colour and small dimensions of
the red one at and below the union (e d). The respective qualities of the two
roots were thus transposed, while the upper portions or crowns were
unaffected: the root of one, naturally weak, became distended and enlarged
by the abundant matter poured into it by its new crown; and in like manner
the root of the other, naturally vigorous, was starved by insufficient food
derived from the new crown, and became diminutive and shrunken (see
Synophty).
'Gard. Chron.,' 1846, p. 252.
A similar anastomosis may take place in the roots. Lindley cites a case
wherein two carrots, of the white Belgian and the red Surrey varieties
respectively, had grown so close to each other that each twisted half round
the other, so that they ultimately became soldered together; the most
singular thing with reference to this union was, that the red carrot (fig. 23,
b), with its small overgrown part above the junction, took the colour and
large dimensions of the white Belgian (d), which, in like manner, with its
larger head above the joining (a), took the colour and small dimensions of
the red one at and below the union (e d). The respective qualities of the two
roots were thus transposed, while the upper portions or crowns were
unaffected: the root of one, naturally weak, became distended and enlarged
by the abundant matter poured into it by its new crown; and in like manner
the root of the other, naturally vigorous, was starved by insufficient food
derived from the new crown, and became diminutive and shrunken (see
Synophty).
Page 98
Fig. 22.—Adhesion of the branches of two elms. 'Gard. Chron.,' 1849, p. 421.
The explanation of the fact that the stumps of felled fir trees occasionally
continue to grow, and to deposit fresh zones of wood over the stump,
depends on similar facts. In Abies pectinata, says Goeppert,[57] the roots of
different individuals frequently unite; hence if one be cut down, its stump
may continue to live, being supplied with nourishment from the adjacent
trees to which it is adherent by means of its roots.
The explanation of the fact that the stumps of felled fir trees occasionally
continue to grow, and to deposit fresh zones of wood over the stump,
depends on similar facts. In Abies pectinata, says Goeppert,[57] the roots of
different individuals frequently unite; hence if one be cut down, its stump
may continue to live, being supplied with nourishment from the adjacent
trees to which it is adherent by means of its roots.
Page 99
Fig. 23.—Adhesion of two roots of carrot. 'Gard. Chron.,' 1851, p. 67.
A not uncommon malformation in mushrooms arises from the confluence of
their stalks (fig. 24), and when the union takes place by means of the pilei,
it sometimes happens, during growth, that the one fungus is detached from
its attachment to the ground, and is borne up with the other, sometimes,
even, being found in an inverted position on the top of its fellow.[58]
The garden operations of budding, grafting and inarching have already been
alluded to as furnishing illustrations of adhesion, but it may be well to refer
briefly to certain other interesting examples of adhesion induced artificially;
thus, the employment of the root as a stock, "root-grafting," is now largely
practised with some plants, as affording a quicker means of propagation
than by cuttings; and a still more curious illustration may be cited in the fact
that it has also been found possible to graft a scion on the leaf in the orange.
[59]
A not uncommon malformation in mushrooms arises from the confluence of
their stalks (fig. 24), and when the union takes place by means of the pilei,
it sometimes happens, during growth, that the one fungus is detached from
its attachment to the ground, and is borne up with the other, sometimes,
even, being found in an inverted position on the top of its fellow.[58]
The garden operations of budding, grafting and inarching have already been
alluded to as furnishing illustrations of adhesion, but it may be well to refer
briefly to certain other interesting examples of adhesion induced artificially;
thus, the employment of the root as a stock, "root-grafting," is now largely
practised with some plants, as affording a quicker means of propagation
than by cuttings; and a still more curious illustration may be cited in the fact
that it has also been found possible to graft a scion on the leaf in the orange.
[59]
Page 100
Fig. 24.—Section through two adherent mushrooms, the upper one inverted.
Mr. Darwin, in his work on the 'Variation of Animals and Plants,' vol. i, p.
395, alludes to the two following remarkable cases of fusion:—"The author
of 'Des Jacinthes' (Amsterdam, 1768, p. 124) says that bulbs of blue and red
hyacinths may be cut in two, and that they will grow together, and throw up
a united stem (and this Mr. Darwin has himself seen), with flowers of the
two colours on the opposite sides. But the remarkable point is, that flowers
are sometimes produced with the two colours blended together." In the
second case related by Mr. Trail, about sixty blue and white potatoes were
cut in halves through the eyes or buds, and the halves were then joined, the
other buds being destroyed. Union took place, and some of the united tubers
produced white, others blue, while some produced tubers partly white and
partly blue.
Adhesion of the axes of plants belonging to different species is a more
singular occurrence than the former, and is of some interest as connected
Mr. Darwin, in his work on the 'Variation of Animals and Plants,' vol. i, p.
395, alludes to the two following remarkable cases of fusion:—"The author
of 'Des Jacinthes' (Amsterdam, 1768, p. 124) says that bulbs of blue and red
hyacinths may be cut in two, and that they will grow together, and throw up
a united stem (and this Mr. Darwin has himself seen), with flowers of the
two colours on the opposite sides. But the remarkable point is, that flowers
are sometimes produced with the two colours blended together." In the
second case related by Mr. Trail, about sixty blue and white potatoes were
cut in halves through the eyes or buds, and the halves were then joined, the
other buds being destroyed. Union took place, and some of the united tubers
produced white, others blue, while some produced tubers partly white and
partly blue.
Adhesion of the axes of plants belonging to different species is a more
singular occurrence than the former, and is of some interest as connected
Page 101
with the operation of grafting. As a general rule horticulturists are of
opinion, and their opinion is borne out by facts, that the operation of
grafting, to be successful, must be practised on plants of close botanical
affinity. On the other hand, it is equally true that some plants very closely
allied cannot be propagated in this manner. Contact between the younger
growing tissues is essential to successful grafting as practised by the
gardener, and is probably quite as necessary in those cases where the
process takes place naturally. Although there is little doubt but that some of
the recorded instances of natural or artificial grafting of plants of distant
botanical affinities are untrustworthy, yet the instances of adhesion between
widely different plants are too numerous and too well attested to allow of
doubt. Moreover, when parasitical plants are considered, such as the
Orobanches, the Cuscutas, and specially the mistleto (Viscum), which may
be found growing on plants of very varied botanical relationship, the
occurrence of occasional adhesion between plants of distant affinity is not
so much to be wondered at. Union between the haulms of wheat and rye,
and other grasses, has been recorded[60]. Moquin-Tandon[61] relates a case
wherein, by accident, a branch of a species of Sophora passed through the
fork, made by two diverging branches of an elder (Sambucus), growing in
the Jardin des Plantes of Toulouse. The branch of the Sophora contracted a
firm adhesion to the elder, and what is remarkable is that, although the latter
has much softer wood than the former, yet the branch of the harder wooded
tree was flattened, as if subjected to great pressure[62]. It is possible that
some of the cases similar to those spoken of by Columella, Virgil[63], and
other classical writers, may have originated in the accidental admission of
seeds into the crevices of trees; in time the seeds grew, and as they did so,
the young plants contracted an adhesion to the supporting tree. Some of the
instances recorded by classical writers may be attributed to intentional or
accidental fallacy, as in the so-called "greffe des charlatans" of more
modern days.
opinion, and their opinion is borne out by facts, that the operation of
grafting, to be successful, must be practised on plants of close botanical
affinity. On the other hand, it is equally true that some plants very closely
allied cannot be propagated in this manner. Contact between the younger
growing tissues is essential to successful grafting as practised by the
gardener, and is probably quite as necessary in those cases where the
process takes place naturally. Although there is little doubt but that some of
the recorded instances of natural or artificial grafting of plants of distant
botanical affinities are untrustworthy, yet the instances of adhesion between
widely different plants are too numerous and too well attested to allow of
doubt. Moreover, when parasitical plants are considered, such as the
Orobanches, the Cuscutas, and specially the mistleto (Viscum), which may
be found growing on plants of very varied botanical relationship, the
occurrence of occasional adhesion between plants of distant affinity is not
so much to be wondered at. Union between the haulms of wheat and rye,
and other grasses, has been recorded[60]. Moquin-Tandon[61] relates a case
wherein, by accident, a branch of a species of Sophora passed through the
fork, made by two diverging branches of an elder (Sambucus), growing in
the Jardin des Plantes of Toulouse. The branch of the Sophora contracted a
firm adhesion to the elder, and what is remarkable is that, although the latter
has much softer wood than the former, yet the branch of the harder wooded
tree was flattened, as if subjected to great pressure[62]. It is possible that
some of the cases similar to those spoken of by Columella, Virgil[63], and
other classical writers, may have originated in the accidental admission of
seeds into the crevices of trees; in time the seeds grew, and as they did so,
the young plants contracted an adhesion to the supporting tree. Some of the
instances recorded by classical writers may be attributed to intentional or
accidental fallacy, as in the so-called "greffe des charlatans" of more
modern days.
Page 102
Adhesion of the roots of different species has been effected artificially, as
between the carrot and the beet root, while Dr. Maclean succeeded in
engrafting, on a red beet, a scion of the white Silesian variety of the same
species. In all these cases, even in the most successful grafts, the amount of
adhesion is very slight; the union in no degree warrants the term fusion, it is
little but simple contact of similar tissues, while new growing matter is
formed all round the cut surfaces, so that the latter become gradually
imbedded in the newly formed matter.
Synophty or adhesion of the embryo.—This often occurs partially in the
embryo plants of the common mistleto (Viscum), but is not of common
occurrence in other plants, even in such cases as the orange (Citrus), the
Cycadeæ, Coniferæ, &c., where there is frequently more than one embryo
in the seed. Alphonse De Candolle has described and figured an instance of
the kind in Euphorbia helioscopia, wherein two embryo plants were
completely grafted together throughout the whole length of their axes,
leaving merely the four cotyledons separate. A similar adnation has been
observed by the same botanist in Lepidium sativum and Sinapis ramosa, as
well as in other plants.[64] I have met with corresponding instances in
Antirrhinum majus and in Cratægus oxyacantha, in the latter case
complicated with the partial atrophy of one of the four cotyledons. It is
necessary to distinguish between such cases and the fallacious appearances
arising from a division of the cotyledons. M. Morren has figured and
described the union of two roots of carrot (Daucus), which were also
spirally twisted. He attributes this union to the blending of two radicles, and
applies the term "rhizocollesy" to this union of the roots.[65] Mr. Thwaites
cites a case wherein two embryos were contained in one seed in a Fuchsia,
and had become adherent. What is still more remarkable, the two embryos
were different, a circumstance attributable to their hybrid origin, the seed
containing them being the result of the fertilisation of Fuchsia coccinea
(quere F. magellanica?) by the pollen of F. fulgens.
between the carrot and the beet root, while Dr. Maclean succeeded in
engrafting, on a red beet, a scion of the white Silesian variety of the same
species. In all these cases, even in the most successful grafts, the amount of
adhesion is very slight; the union in no degree warrants the term fusion, it is
little but simple contact of similar tissues, while new growing matter is
formed all round the cut surfaces, so that the latter become gradually
imbedded in the newly formed matter.
Synophty or adhesion of the embryo.—This often occurs partially in the
embryo plants of the common mistleto (Viscum), but is not of common
occurrence in other plants, even in such cases as the orange (Citrus), the
Cycadeæ, Coniferæ, &c., where there is frequently more than one embryo
in the seed. Alphonse De Candolle has described and figured an instance of
the kind in Euphorbia helioscopia, wherein two embryo plants were
completely grafted together throughout the whole length of their axes,
leaving merely the four cotyledons separate. A similar adnation has been
observed by the same botanist in Lepidium sativum and Sinapis ramosa, as
well as in other plants.[64] I have met with corresponding instances in
Antirrhinum majus and in Cratægus oxyacantha, in the latter case
complicated with the partial atrophy of one of the four cotyledons. It is
necessary to distinguish between such cases and the fallacious appearances
arising from a division of the cotyledons. M. Morren has figured and
described the union of two roots of carrot (Daucus), which were also
spirally twisted. He attributes this union to the blending of two radicles, and
applies the term "rhizocollesy" to this union of the roots.[65] Mr. Thwaites
cites a case wherein two embryos were contained in one seed in a Fuchsia,
and had become adherent. What is still more remarkable, the two embryos
were different, a circumstance attributable to their hybrid origin, the seed
containing them being the result of the fertilisation of Fuchsia coccinea
(quere F. magellanica?) by the pollen of F. fulgens.
Page 103
FOOTNOTES:
[30] Wydler, 'Flora,' 1852, p. 737, tab. ix.
[31] 'El. Ter. Veg.,' p. 254.
[32] 'Bull. Soc. Bot. Fr.,' 1857, p. 451.
[33] 'Bull. Acad. Belg.,' vol. xix, part ii, p. 335.
[34] 'Bull. Soc. Bot. Fr.,' 1860, p. 25.
[35] 'Bull. Soc. Bot. Fr.,' 1861, p. 147.
[36] 'Bull. Acad. Belg.,' vol. xviii, part ii, p. 498.
[37] See also Prillieux, 'Bull. Soc. Bot. Fr.,' 1861, p. 195.
[38] 'Mém. Acad. Toulouse,' 5th Series, vol. iii.
[39] Linnæa, vol. ii. p. 607.
[40] 'Journal Roy. Hort. Soc.,' new ser., vol. i. 1866, p. 200.
[41] 'Bull. Soc. Bot. Fr.,' 1861, p. 159.
[42] Ibid., 1859, p. 467.
[43] 'Flora,' 1858, p. 65, tab. ii.
[44] C. Morren. 'Bull. Acad. Belg.,' vol. xv (Fuchsia, p. 89); vol. xviii, p. 591. (Lobelia, p.
142); vol. xix, p. 352; vol. xx, p. 4.
[45] 'Bull. Soc. Bot. Fr.,' vol. vii, p. 625.
[46] Cramer, 'Bildungsabweichungen,' p. 56, tab. vii, fig. 10, figures a case wherein the two
central flowers of the capitulum of Centaurea Jacea were united together.
[47] 'Bull. Bot.' tab. iii, figs. 4–6.
[48] 'Mém. greffe Ann. Science Nat.,' ser. i, t. xxiv, p. 334.
[49] "Mespilus portentosa." Poit. et Turp., 'Pomol. Franc.,' liv, xxxi, p. 202, pl. 202.
[50] Duchesne, 'Hist. Nat. Frais.,' p. 79.
[51] De Cand., 'Phys. Végét.,' tom. ii, p. 781.
[30] Wydler, 'Flora,' 1852, p. 737, tab. ix.
[31] 'El. Ter. Veg.,' p. 254.
[32] 'Bull. Soc. Bot. Fr.,' 1857, p. 451.
[33] 'Bull. Acad. Belg.,' vol. xix, part ii, p. 335.
[34] 'Bull. Soc. Bot. Fr.,' 1860, p. 25.
[35] 'Bull. Soc. Bot. Fr.,' 1861, p. 147.
[36] 'Bull. Acad. Belg.,' vol. xviii, part ii, p. 498.
[37] See also Prillieux, 'Bull. Soc. Bot. Fr.,' 1861, p. 195.
[38] 'Mém. Acad. Toulouse,' 5th Series, vol. iii.
[39] Linnæa, vol. ii. p. 607.
[40] 'Journal Roy. Hort. Soc.,' new ser., vol. i. 1866, p. 200.
[41] 'Bull. Soc. Bot. Fr.,' 1861, p. 159.
[42] Ibid., 1859, p. 467.
[43] 'Flora,' 1858, p. 65, tab. ii.
[44] C. Morren. 'Bull. Acad. Belg.,' vol. xv (Fuchsia, p. 89); vol. xviii, p. 591. (Lobelia, p.
142); vol. xix, p. 352; vol. xx, p. 4.
[45] 'Bull. Soc. Bot. Fr.,' vol. vii, p. 625.
[46] Cramer, 'Bildungsabweichungen,' p. 56, tab. vii, fig. 10, figures a case wherein the two
central flowers of the capitulum of Centaurea Jacea were united together.
[47] 'Bull. Bot.' tab. iii, figs. 4–6.
[48] 'Mém. greffe Ann. Science Nat.,' ser. i, t. xxiv, p. 334.
[49] "Mespilus portentosa." Poit. et Turp., 'Pomol. Franc.,' liv, xxxi, p. 202, pl. 202.
[50] Duchesne, 'Hist. Nat. Frais.,' p. 79.
[51] De Cand., 'Phys. Végét.,' tom. ii, p. 781.
Page 104
[52] Sched. de monstr. plant. 'Act. Helv.,' tab. i, fig. 8.
[53] 'Mém. greffe,' loc. cit., tab. xxiv, p. 334.
[54] 'Bull. Soc. Bot. Franc.,' 8, pp. 73 and 351, tab. ii; and Röse. 'Bot. Zeit.,' x, p. 410.
[55] Nymphæa lutea, Æsculus Hippocastanum, &c. See Moquin, 'El. Ter. Veg.,' p. 277.
[56] C. Martins, 'Promenade Botanique,' p. 8.
[57] 'Ann. Sc. Nat.,' t. xix, 1843, p. 141, tab. iv.
[58] 'Ann. Nat. Hist.,' ser. 2, vol. ix, tab. xvi. 'Phytologist,' 1857. p. 352, &c.
[59] Quoted from the 'Revue Hortic.' in 'Gard. Chron.,' 1866, p. 386.
[60] Senebier, 'Phys Végét.,' t. iv, p. 426. The same author also cites Romer as having found
two plants of Ranunculus, from the stem of which emerged a daisy. As it is not an uncommon
practice to stick a daisy on a buttercup, it is to be hoped no hoax was played off on M. Romer.
[61] 'El. Ter. Veg.,' p. 289.
[62] An instance of this kind is cited in Dr. Robson's memoir of the late Charles Waterton,
from which it appears that two trees, a spruce fir and an elm, were originally planted side by
side, and had been annually twisted round each other, so that they had in places grown one into
the other, with the result of stunting the growth of both trees, thus illustrating, according to the
opinion of the eccentric naturalist above cited, the incongruous union of Church and State!
[63] See Daubeny, 'Lectures on Roman Husbandry,' p. 156.
[64] A. P. De Candolle, 'Organ Végét.,' t. ii, p. 72, tab. liv, fig. 1.
[65] 'Bull. Acad. Belg.,' t. xx, part i, 1852, p. 43.
PART II.
INDEPENDENCE OR SEPARATION OF ORGANS.
[53] 'Mém. greffe,' loc. cit., tab. xxiv, p. 334.
[54] 'Bull. Soc. Bot. Franc.,' 8, pp. 73 and 351, tab. ii; and Röse. 'Bot. Zeit.,' x, p. 410.
[55] Nymphæa lutea, Æsculus Hippocastanum, &c. See Moquin, 'El. Ter. Veg.,' p. 277.
[56] C. Martins, 'Promenade Botanique,' p. 8.
[57] 'Ann. Sc. Nat.,' t. xix, 1843, p. 141, tab. iv.
[58] 'Ann. Nat. Hist.,' ser. 2, vol. ix, tab. xvi. 'Phytologist,' 1857. p. 352, &c.
[59] Quoted from the 'Revue Hortic.' in 'Gard. Chron.,' 1866, p. 386.
[60] Senebier, 'Phys Végét.,' t. iv, p. 426. The same author also cites Romer as having found
two plants of Ranunculus, from the stem of which emerged a daisy. As it is not an uncommon
practice to stick a daisy on a buttercup, it is to be hoped no hoax was played off on M. Romer.
[61] 'El. Ter. Veg.,' p. 289.
[62] An instance of this kind is cited in Dr. Robson's memoir of the late Charles Waterton,
from which it appears that two trees, a spruce fir and an elm, were originally planted side by
side, and had been annually twisted round each other, so that they had in places grown one into
the other, with the result of stunting the growth of both trees, thus illustrating, according to the
opinion of the eccentric naturalist above cited, the incongruous union of Church and State!
[63] See Daubeny, 'Lectures on Roman Husbandry,' p. 156.
[64] A. P. De Candolle, 'Organ Végét.,' t. ii, p. 72, tab. liv, fig. 1.
[65] 'Bull. Acad. Belg.,' t. xx, part i, 1852, p. 43.
PART II.
INDEPENDENCE OR SEPARATION OF ORGANS.
Page 105
Under this head are included all those instances wherein organs usually
entire, or more or less united, are, or appear to be, split or disunited. It thus
includes such cases as the division of an ordinarily entire leaf into a lobed
or partite one, as well as those characterised by the separation of organs
usually joined together. Union, as has been stated in a previous chapter, is
the result either of persistent integrity or of a junction of originally separate
organs, after their formation; so in like manner, the separation or disjunction
of parts may arise from the absence of that process of union which is
habitual in some cases, or from an actual bonâ fide separation of parts
originally united together. In the former case, the isolation of parts arises
from arrest of development, while in the latter it is due rather to luxuriant
growth. A knowledge, as well of the ordinary as of the unusual course, of
development in any particular flower is thus required in order to ascertain
with accuracy the true nature of the separation of parts. The late Professor
Morren[66] proposed the general term Monosy (μονωσις) for all these cases
of abnormal isolation, subdividing the group into two, as follows—1,
Adesmy (α-δεσμος), including those cases where the separation is
congenital; and 2, Dialysis (διαλυω), comprising those instances where the
isolation is truly a result of the separation of parts previously joined
together. Adesmy, moreover, was by the Belgian savant said to be
homologous when it occurred between members of the same whorl, e.g.
between the sepals of an ordinary monosepalous calyx, or heterologous
when the separation took place between members of different whorls, as
when the calyx is detached from the ovary, &c. The former case would thus
be the converse of cohesion, the latter of adhesion.
To the adoption of these words there is this great objection, that we can but
rarely, in the present state of our knowledge, tell in which group any
particular illustration should be placed.
entire, or more or less united, are, or appear to be, split or disunited. It thus
includes such cases as the division of an ordinarily entire leaf into a lobed
or partite one, as well as those characterised by the separation of organs
usually joined together. Union, as has been stated in a previous chapter, is
the result either of persistent integrity or of a junction of originally separate
organs, after their formation; so in like manner, the separation or disjunction
of parts may arise from the absence of that process of union which is
habitual in some cases, or from an actual bonâ fide separation of parts
originally united together. In the former case, the isolation of parts arises
from arrest of development, while in the latter it is due rather to luxuriant
growth. A knowledge, as well of the ordinary as of the unusual course, of
development in any particular flower is thus required in order to ascertain
with accuracy the true nature of the separation of parts. The late Professor
Morren[66] proposed the general term Monosy (μονωσις) for all these cases
of abnormal isolation, subdividing the group into two, as follows—1,
Adesmy (α-δεσμος), including those cases where the separation is
congenital; and 2, Dialysis (διαλυω), comprising those instances where the
isolation is truly a result of the separation of parts previously joined
together. Adesmy, moreover, was by the Belgian savant said to be
homologous when it occurred between members of the same whorl, e.g.
between the sepals of an ordinary monosepalous calyx, or heterologous
when the separation took place between members of different whorls, as
when the calyx is detached from the ovary, &c. The former case would thus
be the converse of cohesion, the latter of adhesion.
To the adoption of these words there is this great objection, that we can but
rarely, in the present state of our knowledge, tell in which group any
particular illustration should be placed.
Page 106
The terms adopted in the present work are, for the most part, not necessarily
intended to convey any idea as to the organogenetic history of the parts
affected. Where a single organ, that is usually entire, becomes divided the
term Fission is used; in cases where parts of the same whorl become
isolated, the word Dialysis is employed, and in the same sense in which it is
generally used by descriptive botanists, and where the various whorls
become detached one from the other, the occurrence is distinguished by the
application of the term Solution.
FOOTNOTES:
[66] 'Bull. Acad. Belg.,' t. xix, part iii, 1852, p. 315.
CHAPTER I.
FISSION.
When an organ becomes divided it receives at the hands of descriptive
botanists the appellations cleft, partite, or sect, according to the depth of the
division; hence in considering the teratological instances of this nature, the
term fission has suggested itself as an appropriate one to be applied to the
subdivision of an habitually entire or undivided organ. It thus corresponds
pretty nearly in its application with the term Chorisis or "dédoublement," or
with the "disjonctions qui divisent les organes" of Moquin-Tandon.[67] It is
usually, but not always, a concomitant with hypertrophy, and dependent on
luxuriance of growth.
It must be understood therefore that the term, as generally applied, does not
so much indicate the cleavage of a persistent organ, as it does the formation
and development of two or more growing points instead of one, whence
results a branching or forking (di-tri-chotomy) of the affected organ. In
intended to convey any idea as to the organogenetic history of the parts
affected. Where a single organ, that is usually entire, becomes divided the
term Fission is used; in cases where parts of the same whorl become
isolated, the word Dialysis is employed, and in the same sense in which it is
generally used by descriptive botanists, and where the various whorls
become detached one from the other, the occurrence is distinguished by the
application of the term Solution.
FOOTNOTES:
[66] 'Bull. Acad. Belg.,' t. xix, part iii, 1852, p. 315.
CHAPTER I.
FISSION.
When an organ becomes divided it receives at the hands of descriptive
botanists the appellations cleft, partite, or sect, according to the depth of the
division; hence in considering the teratological instances of this nature, the
term fission has suggested itself as an appropriate one to be applied to the
subdivision of an habitually entire or undivided organ. It thus corresponds
pretty nearly in its application with the term Chorisis or "dédoublement," or
with the "disjonctions qui divisent les organes" of Moquin-Tandon.[67] It is
usually, but not always, a concomitant with hypertrophy, and dependent on
luxuriance of growth.
It must be understood therefore that the term, as generally applied, does not
so much indicate the cleavage of a persistent organ, as it does the formation
and development of two or more growing points instead of one, whence
results a branching or forking (di-tri-chotomy) of the affected organ. In
Page 107
some instances it seems rather to be due to the relative deficiency of
cellular, as contrasted with fibro-vascular tissue.
Fission of axile organs.—This condition is scarcely to be distinguished
from multiplication of the axile organs (which see). A little attention,
however, will generally show whether the unusual number of branches is a
consequence of the development of a large number of distinct shoots, as
happens, for instance, when a tree is pollarded, or of a division of one. M.
Fournier[68] gives as an illustration the case of a specimen of Ruscus
aculeatus in which there occurred a division of the foliaceous branches into
two segments, reaching as far as the insertion of the flower, but no further.
He also mentions lateral cleavage effected by a notching of the margin, the
notch being anterior to the flowers and always directed towards their
insertion. In the allied genus Danaë, Webb, 'Phyt. Canar.,' p. 320, describes
the fascicles of flowers as in "crenulis brevibus ad marginem ramulorum
dispositis." Sometimes, on the other hand, Danaë has a fascicle of flowers
inserted on the middle of the upper surface, as in Ruscus. Wigand mentions
an instance in Digitalis lutea, where the upper part of the stem was divided
into six or seven racemes; possibly this was a case of fasciation, but such a
division of the inflorescence is by no means uncommon in the spicate
species of Veronica. I have also seen it in Plantago lanceolata, Reseda
luteola, Campanula medium, Epacris impressa, and a bifurcation of the axis
of the spikelet within the outer glumes in Lolium perenne[69] and
Anthoxanthum odoratum. In the Kew Museum is preserved a cone of Abies
excelsa,[70] dividing into two divisions, each bearing bracts and scales. A
similar thing frequently occurs in the male catkins of Cedrus Libani (fig.
25).
cellular, as contrasted with fibro-vascular tissue.
Fission of axile organs.—This condition is scarcely to be distinguished
from multiplication of the axile organs (which see). A little attention,
however, will generally show whether the unusual number of branches is a
consequence of the development of a large number of distinct shoots, as
happens, for instance, when a tree is pollarded, or of a division of one. M.
Fournier[68] gives as an illustration the case of a specimen of Ruscus
aculeatus in which there occurred a division of the foliaceous branches into
two segments, reaching as far as the insertion of the flower, but no further.
He also mentions lateral cleavage effected by a notching of the margin, the
notch being anterior to the flowers and always directed towards their
insertion. In the allied genus Danaë, Webb, 'Phyt. Canar.,' p. 320, describes
the fascicles of flowers as in "crenulis brevibus ad marginem ramulorum
dispositis." Sometimes, on the other hand, Danaë has a fascicle of flowers
inserted on the middle of the upper surface, as in Ruscus. Wigand mentions
an instance in Digitalis lutea, where the upper part of the stem was divided
into six or seven racemes; possibly this was a case of fasciation, but such a
division of the inflorescence is by no means uncommon in the spicate
species of Veronica. I have also seen it in Plantago lanceolata, Reseda
luteola, Campanula medium, Epacris impressa, and a bifurcation of the axis
of the spikelet within the outer glumes in Lolium perenne[69] and
Anthoxanthum odoratum. In the Kew Museum is preserved a cone of Abies
excelsa,[70] dividing into two divisions, each bearing bracts and scales. A
similar thing frequently occurs in the male catkins of Cedrus Libani (fig.
25).
Page 108
Fig. 25.—Bifurcated male inflorescence, Cedrus Libani.
This subdivision of axial organs is not unfrequently the result of some
injury or mutilation, thus Duval Jouve alludes to the frequency with which
branched stems are produced in the various species of Equisetum, as a
consequence of injuries to the main stem, but this is rather to be considered
as a multiplication of parts than as a subdivision of one.
This subdivision of axial organs is not unfrequently the result of some
injury or mutilation, thus Duval Jouve alludes to the frequency with which
branched stems are produced in the various species of Equisetum, as a
consequence of injuries to the main stem, but this is rather to be considered
as a multiplication of parts than as a subdivision of one.
Page 109
Fig. 26.—Bifurcated leaf of Lamium album, &c.
Fission of foliar organs.—Many leaves exhibit constantly the process of
fission, such as the Salisburia adiantifolia, and which is due perhaps as
much to the absence or relatively small proportion of cellular as compared
with vascular tissue, as to absolute fission. In the same way we have
laciniated leaves of the Persian lilac, Syringa persica, and Moquin mentions
instances in a species of Mercurialis in which the leaves were deeply
slashed. In Chenopodium Quinoa the leaves were so numerous and the
clefts so deep, that the species was hardly recognisable, while on a branch
of Rhus Cotinus observed by De Candolle the lobes were so narrow and so
fine as to give the plant the aspect of an Umbellifer. Wigand ('Flora,' 1856,
p. 706) speaks of the leaves of Dipsacus fullonum with bi-partite leaves;
Moquin mentions the occurrence of a leaf of an oleander bi-lobed at the
summit, so as to give the appearance of a fusion of two leaves. Steinheil has
recorded an instance in Scabiosa atropurpurea in which one of the stem
Fission of foliar organs.—Many leaves exhibit constantly the process of
fission, such as the Salisburia adiantifolia, and which is due perhaps as
much to the absence or relatively small proportion of cellular as compared
with vascular tissue, as to absolute fission. In the same way we have
laciniated leaves of the Persian lilac, Syringa persica, and Moquin mentions
instances in a species of Mercurialis in which the leaves were deeply
slashed. In Chenopodium Quinoa the leaves were so numerous and the
clefts so deep, that the species was hardly recognisable, while on a branch
of Rhus Cotinus observed by De Candolle the lobes were so narrow and so
fine as to give the plant the aspect of an Umbellifer. Wigand ('Flora,' 1856,
p. 706) speaks of the leaves of Dipsacus fullonum with bi-partite leaves;
Moquin mentions the occurrence of a leaf of an oleander bi-lobed at the
summit, so as to give the appearance of a fusion of two leaves. Steinheil has
recorded an instance in Scabiosa atropurpurea in which one of the stem
Page 110
leaves presented the following peculiarities. It was simple below, but
divided above into two equal lobes, provided each with a median nerve.[71]
Steinheil has also recorded a Cerastium in which one of the leaves was
provided with two midribs; above this leaf was a group of ternate leaves. I
have seen similar instances in the common Elm, Ulmus campestris, and also
in the common nettle, Urtica dioica, the leaves of which latter thus
resembled those of Urtica biloba, which are habitually bilobed at the
summit. M. Clos[72] mentions an instance where the terminal leaf and first
bract of Orchis sambucina were divided into two segments. The same
author also mentions the leaves of Anemiopsis californica, which were
divided in their upper halves each into two lobes—also leaves of a lentil
springing from a fasciated stem and completely divided into two segments,
but with only a single bud in the axil. The axillary branches in like manner
showed traces of cleavage. Fig. 26 represents a case of this kind in Lamium
album, conjoined with suppression of the flowers on one side of the stem. I
have also in my herbarium a leaf of Arum maculatum, with a stalk single at
the base, but dividing into two separate stalks, each bearing a hastate
lamina, the form of which is so perfect that were it not from the venation of
the sheath it would be considered that there was here a union of two leaves
rather than a bifurcation of one. A garden Pelargonium presented the same
appearance.
divided above into two equal lobes, provided each with a median nerve.[71]
Steinheil has also recorded a Cerastium in which one of the leaves was
provided with two midribs; above this leaf was a group of ternate leaves. I
have seen similar instances in the common Elm, Ulmus campestris, and also
in the common nettle, Urtica dioica, the leaves of which latter thus
resembled those of Urtica biloba, which are habitually bilobed at the
summit. M. Clos[72] mentions an instance where the terminal leaf and first
bract of Orchis sambucina were divided into two segments. The same
author also mentions the leaves of Anemiopsis californica, which were
divided in their upper halves each into two lobes—also leaves of a lentil
springing from a fasciated stem and completely divided into two segments,
but with only a single bud in the axil. The axillary branches in like manner
showed traces of cleavage. Fig. 26 represents a case of this kind in Lamium
album, conjoined with suppression of the flowers on one side of the stem. I
have also in my herbarium a leaf of Arum maculatum, with a stalk single at
the base, but dividing into two separate stalks, each bearing a hastate
lamina, the form of which is so perfect that were it not from the venation of
the sheath it would be considered that there was here a union of two leaves
rather than a bifurcation of one. A garden Pelargonium presented the same
appearance.
Page 111
Fig. 27.—Bifurcated leaf of Pelargonium.
Fern fronds are particularly liable to this kind of subdivision, and they
exhibit it in almost every degree, from a simple bifurcation of the frond to
the formation of large tufts of small lobes all formed on the same plan by
the repeated forking of the pinnules. These may be considered as cases of
hypertrophy.
Moquin-Tandon, at a meeting of the Botanical Society of France (April 3rd,
1858) exhibited a leaf of Cerasus Lauro-Cerasus divided in such a manner
as to resemble a leaf of Citrus or of Phyllarthron. In this case, therefore, the
disunion must have taken place laterally, and not from apex towards base,
as is most common. The leaves of the common horse-radish, Cochlearia
Armoracia, are very subject to this pinnated subdivision of the margin, and
numerous other illustrations might be given.
Fern fronds are particularly liable to this kind of subdivision, and they
exhibit it in almost every degree, from a simple bifurcation of the frond to
the formation of large tufts of small lobes all formed on the same plan by
the repeated forking of the pinnules. These may be considered as cases of
hypertrophy.
Moquin-Tandon, at a meeting of the Botanical Society of France (April 3rd,
1858) exhibited a leaf of Cerasus Lauro-Cerasus divided in such a manner
as to resemble a leaf of Citrus or of Phyllarthron. In this case, therefore, the
disunion must have taken place laterally, and not from apex towards base,
as is most common. The leaves of the common horse-radish, Cochlearia
Armoracia, are very subject to this pinnated subdivision of the margin, and
numerous other illustrations might be given.
Page 112
Fig. 28.—Bifurcated frond, Scolopendrium vulgare.
A. Braun describes a singular case in a leaf of Irina glabra wherein the
blade of the leaf on one side was deeply and irregularly laciniated, the other
side remaining entire. (Verhandl., d. 35, Naturforscherversammlung, tab. 3.)
Laciniate varieties of plants are of frequent occurrence in gardens where
they are often cultivated for their beauty or singularity; thus, there are
laciniated alders, fern-leaved beeches and limes, oak-leaved laburnums, &c.
A list of several of these is subjoined. A similar fission takes place
constantly in the cotyledons of some plants, sometimes, as in Coniferæ, to
such an extent as to give an appearance as if there were several cotyledons.
[73]
It is not always easy to recognise, at a first glance, whether the division be
the result of disunion or of an incomplete union of two leaves, but we may
be guided by the number of leaves in the cycle or the whorl. The number is
A. Braun describes a singular case in a leaf of Irina glabra wherein the
blade of the leaf on one side was deeply and irregularly laciniated, the other
side remaining entire. (Verhandl., d. 35, Naturforscherversammlung, tab. 3.)
Laciniate varieties of plants are of frequent occurrence in gardens where
they are often cultivated for their beauty or singularity; thus, there are
laciniated alders, fern-leaved beeches and limes, oak-leaved laburnums, &c.
A list of several of these is subjoined. A similar fission takes place
constantly in the cotyledons of some plants, sometimes, as in Coniferæ, to
such an extent as to give an appearance as if there were several cotyledons.
[73]
It is not always easy to recognise, at a first glance, whether the division be
the result of disunion or of an incomplete union of two leaves, but we may
be guided by the number of leaves in the cycle or the whorl. The number is
Page 113
complete in cases of partial disjunction, while in cases of fusion it is
incomplete. Again, in instances of disjunction, there is only one point of
origin, but, when two leaves are grafted together, two such points may
generally be detected at the base of the leaf, or a transverse section of the
leaf-stalk will show indications of fusion. The number and position of the
midribs will also serve as a guide, as in cases of fusion there are generally
two or more midribs, according to the number of fused leaves; but as
Moquin well remarks, this latter character cannot be always depended upon,
for the median nerve may divide without any corresponding separation of
the cellular portions of the leaf. The author just quoted cites examples of
this kind in Cardamine pratensis, Hedera Helix, Plantago major, Geranium
nodosum.
The following list of plants commonly producing leaves that are cleft or
divided, to a greater extent than is usual in the species, is mainly taken from
one given by Schlechtendal, 'Bot. Zeit.,' 1844, p. 441, with additions from
other sources. The ! indicates that the author has himself met with the
deviation in question. Many are cultivated as garden varieties under the
names here given.
Trollius europæus dissectus.
Chelidonium majus laciniatum!
Glaucium luteum.
Brassica oleracea!
Tilia parvifolia laciniata.
asplenifolia!
Acer platanoides laciniatum.
crispum.
Æsculus Hippocastanum incisum!
asplenifolium.
Vitis vinifera apiifolia!
laciniosa.
Ilex Aquifolium!
Rhus Toxicodendron quercifolium.
incomplete. Again, in instances of disjunction, there is only one point of
origin, but, when two leaves are grafted together, two such points may
generally be detected at the base of the leaf, or a transverse section of the
leaf-stalk will show indications of fusion. The number and position of the
midribs will also serve as a guide, as in cases of fusion there are generally
two or more midribs, according to the number of fused leaves; but as
Moquin well remarks, this latter character cannot be always depended upon,
for the median nerve may divide without any corresponding separation of
the cellular portions of the leaf. The author just quoted cites examples of
this kind in Cardamine pratensis, Hedera Helix, Plantago major, Geranium
nodosum.
The following list of plants commonly producing leaves that are cleft or
divided, to a greater extent than is usual in the species, is mainly taken from
one given by Schlechtendal, 'Bot. Zeit.,' 1844, p. 441, with additions from
other sources. The ! indicates that the author has himself met with the
deviation in question. Many are cultivated as garden varieties under the
names here given.
Trollius europæus dissectus.
Chelidonium majus laciniatum!
Glaucium luteum.
Brassica oleracea!
Tilia parvifolia laciniata.
asplenifolia!
Acer platanoides laciniatum.
crispum.
Æsculus Hippocastanum incisum!
asplenifolium.
Vitis vinifera apiifolia!
laciniosa.
Ilex Aquifolium!
Rhus Toxicodendron quercifolium.
Page 114
Cotinus.
Ervum Lens.
Cytisus Laburnum quercifolium!
incisum.
Rubus fraticosus laciniatus!
Pyrcis communis.
Cerasus Lauro-cerasus.
Apium graveolens!
Pimpinella magna.
Saxifraga.
Cratægus Oxyacantha laciniata.
quercifolia!
Ribes nigrum.
Sambucus nigra laciniata!
racemosa laciniata.
Dipsacus fullonum.
Scabiosa atropurpurea!
Symphoricarpus racemosus.
Helianthus sp.!
Lonicera Periclymenum quercifolia!
Syringa persica laciniata!
Syringa vulgaris!
Nerium Oleander!
Lamium purpureum.
album!
Salvia officinalis.
Solanum Dulcamara!
Fraxinus excelsior crispa.
Veronica austriaca.
Polemonium cæruleum.
Juglans regia laciniata!
heterophylla.
filicifolia.
Anemiopsis californica.
Chenopodium Quinoa.
Ulmus americana incisa.
Fagus sylvatica heterophylla!
laciniata!
aspleniifolia!
Ervum Lens.
Cytisus Laburnum quercifolium!
incisum.
Rubus fraticosus laciniatus!
Pyrcis communis.
Cerasus Lauro-cerasus.
Apium graveolens!
Pimpinella magna.
Saxifraga.
Cratægus Oxyacantha laciniata.
quercifolia!
Ribes nigrum.
Sambucus nigra laciniata!
racemosa laciniata.
Dipsacus fullonum.
Scabiosa atropurpurea!
Symphoricarpus racemosus.
Helianthus sp.!
Lonicera Periclymenum quercifolia!
Syringa persica laciniata!
Syringa vulgaris!
Nerium Oleander!
Lamium purpureum.
album!
Salvia officinalis.
Solanum Dulcamara!
Fraxinus excelsior crispa.
Veronica austriaca.
Polemonium cæruleum.
Juglans regia laciniata!
heterophylla.
filicifolia.
Anemiopsis californica.
Chenopodium Quinoa.
Ulmus americana incisa.
Fagus sylvatica heterophylla!
laciniata!
aspleniifolia!
Page 115
incisa.
salicifolia!
Mercurialis perennis.
Urtica dioica.
Quercus Cerris laciniata!
pubescens filicina.
Betula populifolia laciniata.
alba dalecarlica.
Alnus incana laciniata!
glutinosa laciniata!
quercifolia.
oxyacanthifolia.
Corylus Avellana heterophylla!
laciniata!
urticifolia.
Carpinus Betulus incisa!
quercifolia.
heterophylla.
Castanea vesca heterophylla.
quercifolia.
incisa.
Populus alba acerifolia.
palmata.
quercifolia.
balsamifera.
Orchis sambucina.
Arum maculatum.
Filices sp. pl.
See also Schlechtendal, 'Bot. Zeit.,' tom. xiii, p. 823. A. Braun, loc. supra citat. For Ferns
too numerous for insertion, see Moore, 'Nature-Printed Ferns,' 8vo ed., 2 vols. Clos, 'Mém.
Acad. Toulouse,' 1862, p. 51.
Fission of the petals, &c.—The floral leaves are subject to a similar
process of cleavage to that which has just been mentioned as taking place in
the leaves. This, indeed, occurs very often as a normal occurrence as in the
petals of mignonette (Reseda), or those of Alsine media and many other
plants. Here, however, we have only to allude to those instances in which
salicifolia!
Mercurialis perennis.
Urtica dioica.
Quercus Cerris laciniata!
pubescens filicina.
Betula populifolia laciniata.
alba dalecarlica.
Alnus incana laciniata!
glutinosa laciniata!
quercifolia.
oxyacanthifolia.
Corylus Avellana heterophylla!
laciniata!
urticifolia.
Carpinus Betulus incisa!
quercifolia.
heterophylla.
Castanea vesca heterophylla.
quercifolia.
incisa.
Populus alba acerifolia.
palmata.
quercifolia.
balsamifera.
Orchis sambucina.
Arum maculatum.
Filices sp. pl.
See also Schlechtendal, 'Bot. Zeit.,' tom. xiii, p. 823. A. Braun, loc. supra citat. For Ferns
too numerous for insertion, see Moore, 'Nature-Printed Ferns,' 8vo ed., 2 vols. Clos, 'Mém.
Acad. Toulouse,' 1862, p. 51.
Fission of the petals, &c.—The floral leaves are subject to a similar
process of cleavage to that which has just been mentioned as taking place in
the leaves. This, indeed, occurs very often as a normal occurrence as in the
petals of mignonette (Reseda), or those of Alsine media and many other
plants. Here, however, we have only to allude to those instances in which
Page 116
the cleavage occurs in flowers whose sepals or petals are usually entire.
Under this category Moquin mentions a petal of Brassica oleracea
completely split into two. Linné in his 'Flora Lapponica' (pp. 145 and 164)
mentions quadrifid petals of Lychnis dioica, and much divided petals of
Rubus arcticus. Among other plants subject to this division of sepals or
petals may be mentioned as having come within the writer's personal
observation, Ranunculus Lingua, R. acris, Papaver somniferum, and others
of this genus, Saponaria sp., Dianthus, Narcissus, &c.
In some of the garden varieties of Cyclamen the corolla looks at first sight
as if double, and the plan of the flower is oblong or elliptical, instead of
circular. In these flowers each lobe of the corolla is divided almost to the
base into two lobes, so that there appear to be ten lobes to the corolla
instead of five, as usual. The stamens are normal in form and number in
these flowers.
In the paroquet tulips of gardeners the segments of the perianth are deeply
and irregularly gashed, the segments occasionally becoming rolled up and
their margins coherent so as to form little tubular spurs. I have also noticed
the segments of the perianth in Crocus and Colchicum deeply cleft, so much
so sometimes, as to equal in this particular the stigmas. In the flowers of a
species of Oncidium, communicated to me by Mr. Currey, the lip was
divided into three segments perfectly distinct one from the other, but
confluent with the column; the two side pieces had callosities at the upper
edge close to the base, the central piece had a similar wartlike process in its
centre. In these flowers the ovary, the stigma, and the anther were all in a
rudimentary condition. Some verbenas raised by Mr. Wills offer a curious
illustration of this condition. It will be remembered that some of the lobes
or petals of a verbena are normally divided at the base to a slight degree,
but in the flowers in question this is carried to such an extent that the
enlarged lobes are pushed into the centre of the flower and simulate, at a
Under this category Moquin mentions a petal of Brassica oleracea
completely split into two. Linné in his 'Flora Lapponica' (pp. 145 and 164)
mentions quadrifid petals of Lychnis dioica, and much divided petals of
Rubus arcticus. Among other plants subject to this division of sepals or
petals may be mentioned as having come within the writer's personal
observation, Ranunculus Lingua, R. acris, Papaver somniferum, and others
of this genus, Saponaria sp., Dianthus, Narcissus, &c.
In some of the garden varieties of Cyclamen the corolla looks at first sight
as if double, and the plan of the flower is oblong or elliptical, instead of
circular. In these flowers each lobe of the corolla is divided almost to the
base into two lobes, so that there appear to be ten lobes to the corolla
instead of five, as usual. The stamens are normal in form and number in
these flowers.
In the paroquet tulips of gardeners the segments of the perianth are deeply
and irregularly gashed, the segments occasionally becoming rolled up and
their margins coherent so as to form little tubular spurs. I have also noticed
the segments of the perianth in Crocus and Colchicum deeply cleft, so much
so sometimes, as to equal in this particular the stigmas. In the flowers of a
species of Oncidium, communicated to me by Mr. Currey, the lip was
divided into three segments perfectly distinct one from the other, but
confluent with the column; the two side pieces had callosities at the upper
edge close to the base, the central piece had a similar wartlike process in its
centre. In these flowers the ovary, the stigma, and the anther were all in a
rudimentary condition. Some verbenas raised by Mr. Wills offer a curious
illustration of this condition. It will be remembered that some of the lobes
or petals of a verbena are normally divided at the base to a slight degree,
but in the flowers in question this is carried to such an extent that the
enlarged lobes are pushed into the centre of the flower and simulate, at a
Page 117
first glance, a distinct and separate organ, though in reality it is but an
enlargement of what occurs normally.[74]
Fig. 29.—Flower of Oncidium sp. seen from the back. The lip is divided into three unequal segments.
Moquin mentions having seen the stamens of Matthiola incana and Silene
conica completely divided, each section bearing half an anther, exactly as
happens in Polygalaceæ. In tulips and lilies the same author mentions
division of the anther only, the filament remaining entire, as happens
naturally in many species of Vaccinium.
A division of the individual carpels occurs very frequently when those
organs become more or less leafy, as in Trifolium repens, and other plants to
be hereafter mentioned.
The instances given in this chapter have all been cases wherein the division
or the accessory growth has taken place in one plane only and that plane the
enlargement of what occurs normally.[74]
Fig. 29.—Flower of Oncidium sp. seen from the back. The lip is divided into three unequal segments.
Moquin mentions having seen the stamens of Matthiola incana and Silene
conica completely divided, each section bearing half an anther, exactly as
happens in Polygalaceæ. In tulips and lilies the same author mentions
division of the anther only, the filament remaining entire, as happens
naturally in many species of Vaccinium.
A division of the individual carpels occurs very frequently when those
organs become more or less leafy, as in Trifolium repens, and other plants to
be hereafter mentioned.
The instances given in this chapter have all been cases wherein the division
or the accessory growth has taken place in one plane only and that plane the
Page 118
same as that of the affected organ, but there are other examples, probably
equally due to fissiparous division, where the new growth is either parallel
to, or even at angle with the primary organ. Of such nature are some of
those instances wherein two leaves appear to be placed back to back. These
partake of the nature of excrescences or of exaggerated developments, and
hence will be more fully treated of under the head of hypertrophy. It must
be remembered that in some of these cases the fission may be a resumption
of characters proper to the species under natural conditions, but lost by
cultivation or otherwise. Thus, Mr. Buckman accounts for "finger-and-toe"
in root-crops on the principle of reversion to the wild form.
FOOTNOTES:
[67] Loc. cit., p. 295.
[68] 'Bull. Soc. Bot. France,' 1857, p. 758.
[69] Masters, 'Jourl. Linn. Soc.,' vol. vii, p. 121.
[70] Cramer, 'Bildungsabweichungen,' p. 4, tab. vi, fig. 4, figures a case of the same kind in
Pinus Cembra.
[71] 'Ann. des Science Nat.,' 2nd series, t. iv, p. 147, tab. v, figs. 3 and 4.
[72] 'Mém. Acad. Scien. Toulouse,' 5th series, vol. iii.
[73] Duchartre, 'Ann. Sc. Nat.,' 3rd series, 1848, vol. x, p. 207.
[74] Masters, 'Rep. Bot. Congress,' London, 1866, p. 136, tab. 7, f. 15, 16.
CHAPTER II.
DIALYSIS.
This term is here made use of in the same sense as in descriptive botany, to
indicate the isolation of parts of the same whorl; it is thus the opposite of
equally due to fissiparous division, where the new growth is either parallel
to, or even at angle with the primary organ. Of such nature are some of
those instances wherein two leaves appear to be placed back to back. These
partake of the nature of excrescences or of exaggerated developments, and
hence will be more fully treated of under the head of hypertrophy. It must
be remembered that in some of these cases the fission may be a resumption
of characters proper to the species under natural conditions, but lost by
cultivation or otherwise. Thus, Mr. Buckman accounts for "finger-and-toe"
in root-crops on the principle of reversion to the wild form.
FOOTNOTES:
[67] Loc. cit., p. 295.
[68] 'Bull. Soc. Bot. France,' 1857, p. 758.
[69] Masters, 'Jourl. Linn. Soc.,' vol. vii, p. 121.
[70] Cramer, 'Bildungsabweichungen,' p. 4, tab. vi, fig. 4, figures a case of the same kind in
Pinus Cembra.
[71] 'Ann. des Science Nat.,' 2nd series, t. iv, p. 147, tab. v, figs. 3 and 4.
[72] 'Mém. Acad. Scien. Toulouse,' 5th series, vol. iii.
[73] Duchartre, 'Ann. Sc. Nat.,' 3rd series, 1848, vol. x, p. 207.
[74] Masters, 'Rep. Bot. Congress,' London, 1866, p. 136, tab. 7, f. 15, 16.
CHAPTER II.
DIALYSIS.
This term is here made use of in the same sense as in descriptive botany, to
indicate the isolation of parts of the same whorl; it is thus the opposite of
Page 119
cohesion. Morren, as has been previously stated, employed the word in a
different sense, while Moquin-Tandon[75] included cases of this description
under the category of "Disjonctions qui isolent les organes."
Dialysis, as here understood, may be the result of an arrest of development,
in consequence of which parts that under ordinary circumstances would
become fused, do not do so; or, on the other hand, it may be the result of an
actual separation between parts primitively undivided. As it is not possible
in every case to distinguish between the effects of these two diverse causes,
no attempt is here made to do so.
Dialysis of the margins of individual foliar organs.—In cases where the
leaf or leaf-like organ is ordinarily tubular or horn-like in form, owing to
the cohesion of its edges, it may happen either from lack of union or from
actual separation of the previously united edges, that the tubular shape is
replaced by the ordinary flattened expansion. Thus, in Eranthis hyemalis,
wherein the petals (nectaries) are tubular and the sepals flat, I have met with
numerous instances of transition from the one form to the other, as shown in
fig. 9, p. 24.
It is, however, in the carpels that this separation occurs most frequently.
When these organs appear under the guise of leaves, as they often do, their
margins are disunited, so that the carpel becomes flat or open. This happens
in the strawberry (Fragaria), the columbine (Aquilegia), in Trifolium
repens, Ranunculus Ficaria, &c.[76]
Dialysis of the parts of the same whorl:—calyx.—The separation of an
ordinarily coherent series into its constituent parts is necessarily of more
common occurrence than the foregoing. As here understood, it is the precise
converse of cohesion, and it may be represented diagrammatically by a
dotted line above the letters denoting the sepals, petals, &c. When this
different sense, while Moquin-Tandon[75] included cases of this description
under the category of "Disjonctions qui isolent les organes."
Dialysis, as here understood, may be the result of an arrest of development,
in consequence of which parts that under ordinary circumstances would
become fused, do not do so; or, on the other hand, it may be the result of an
actual separation between parts primitively undivided. As it is not possible
in every case to distinguish between the effects of these two diverse causes,
no attempt is here made to do so.
Dialysis of the margins of individual foliar organs.—In cases where the
leaf or leaf-like organ is ordinarily tubular or horn-like in form, owing to
the cohesion of its edges, it may happen either from lack of union or from
actual separation of the previously united edges, that the tubular shape is
replaced by the ordinary flattened expansion. Thus, in Eranthis hyemalis,
wherein the petals (nectaries) are tubular and the sepals flat, I have met with
numerous instances of transition from the one form to the other, as shown in
fig. 9, p. 24.
It is, however, in the carpels that this separation occurs most frequently.
When these organs appear under the guise of leaves, as they often do, their
margins are disunited, so that the carpel becomes flat or open. This happens
in the strawberry (Fragaria), the columbine (Aquilegia), in Trifolium
repens, Ranunculus Ficaria, &c.[76]
Dialysis of the parts of the same whorl:—calyx.—The separation of an
ordinarily coherent series into its constituent parts is necessarily of more
common occurrence than the foregoing. As here understood, it is the precise
converse of cohesion, and it may be represented diagrammatically by a
dotted line above the letters denoting the sepals, petals, &c. When this
Page 120
change happens in the calyx we have the gamosepalous condition replaced
by the polysepalous one, as thus represented:
.............
S S S S S
instead of
_____________
S S S S S
as in a calyx of five coherent sepals.
Detachment of this kind occurs not unfrequently, as in Primula vulgaris,
Trifolium repens, &c. In Rosaceæ and Pomaceæ this separation of the calyx
is of the more moment, as it has reference to the structure of the inferior
ovary, as will be more fully mentioned hereafter. Here, however, a case
recorded by M. J. E. Planchon may be alluded to[77] wherein a quince fruit
(Cydonia) was surmounted by five leaves, the surface of the pome being
marked by as many prominences, which apparently corresponded to the five
stalks of the calycine leaves. In this specimen, then, the inferior position of
the ovary appeared to be not so much due to an expansion of the fruit stalk,
as to the fusion of the hypertrophied stalks of the sepals. Some of the
malformations among Cucurbits point to a similar structure. It is probable
that in many of these cases the so-called inferior ovary is partly axial partly
foliar, i.e., sepaline, and partly carpellary in its nature.
Dialysis of the sepals in calyces that are usually gamosepalous has been
most frequently observed in Rosaceæ, Pomaceæ, Umbelliferæ, less
commonly in Leguminosæ, also in the following genera:—Primula,
Symphytum, Gentiana, Campanula, &c.
by the polysepalous one, as thus represented:
.............
S S S S S
instead of
_____________
S S S S S
as in a calyx of five coherent sepals.
Detachment of this kind occurs not unfrequently, as in Primula vulgaris,
Trifolium repens, &c. In Rosaceæ and Pomaceæ this separation of the calyx
is of the more moment, as it has reference to the structure of the inferior
ovary, as will be more fully mentioned hereafter. Here, however, a case
recorded by M. J. E. Planchon may be alluded to[77] wherein a quince fruit
(Cydonia) was surmounted by five leaves, the surface of the pome being
marked by as many prominences, which apparently corresponded to the five
stalks of the calycine leaves. In this specimen, then, the inferior position of
the ovary appeared to be not so much due to an expansion of the fruit stalk,
as to the fusion of the hypertrophied stalks of the sepals. Some of the
malformations among Cucurbits point to a similar structure. It is probable
that in many of these cases the so-called inferior ovary is partly axial partly
foliar, i.e., sepaline, and partly carpellary in its nature.
Dialysis of the sepals in calyces that are usually gamosepalous has been
most frequently observed in Rosaceæ, Pomaceæ, Umbelliferæ, less
commonly in Leguminosæ, also in the following genera:—Primula,
Symphytum, Gentiana, Campanula, &c.
Page 121
Fig. 30.—Dialysis of the sepals and petals in Correa.
Dialysis of the corolla is likewise of frequent occurrence, either partially or
to such an extent as to render the corolla truly polypetalous. Among
Labiatæ the upper lip of the corolla may be often met with partially cleft, as
it is constantly in Phlomis biloba, or more markedly among the Lobeliaceæ.
In the Compositæ, a similar separation of the petals is not infrequent, thus
showing frequent transitional stages between the labiatifloral and
tubulifloral divisions respectively. The ligulate corollas also may often be
found in Chrysanthemums, Dahlias, &c., more or less deeply divided into
their component parts.
A more complete separation occurs not unfrequently in Campanula,
Rhododendron, Phlox, &c. Figs. 30 and 31 illustrate dialysis of the corolla;
the first in Correa, the second in Campanula.
Dialysis of the corolla is likewise of frequent occurrence, either partially or
to such an extent as to render the corolla truly polypetalous. Among
Labiatæ the upper lip of the corolla may be often met with partially cleft, as
it is constantly in Phlomis biloba, or more markedly among the Lobeliaceæ.
In the Compositæ, a similar separation of the petals is not infrequent, thus
showing frequent transitional stages between the labiatifloral and
tubulifloral divisions respectively. The ligulate corollas also may often be
found in Chrysanthemums, Dahlias, &c., more or less deeply divided into
their component parts.
A more complete separation occurs not unfrequently in Campanula,
Rhododendron, Phlox, &c. Figs. 30 and 31 illustrate dialysis of the corolla;
the first in Correa, the second in Campanula.
Page 122
Fig. 31.—Dialysis of the corolla in Campanula sp., after De Candolle.
In the last-named genus, C. rotundifolia has been found with polypetalous
flowers in a wild state in the mountains of Canton Neufchatel, Switzerland,
and gave rise to the creation of a new genus. This form is now introduced
into gardens.
It must be remembered that in some genera, where this separation of the
petals has been met with, there are species in which a similar isolation
occurs normally, as in Rhododendron. R. linearilobum, a Japanese species,
offers a good illustration of this.
The following list contains the names of the genera in which this separation
of the petals of an ordinarily gamopetalous flower takes place most
frequently.
Correa.
Campanula! sp. pl.
In the last-named genus, C. rotundifolia has been found with polypetalous
flowers in a wild state in the mountains of Canton Neufchatel, Switzerland,
and gave rise to the creation of a new genus. This form is now introduced
into gardens.
It must be remembered that in some genera, where this separation of the
petals has been met with, there are species in which a similar isolation
occurs normally, as in Rhododendron. R. linearilobum, a Japanese species,
offers a good illustration of this.
The following list contains the names of the genera in which this separation
of the petals of an ordinarily gamopetalous flower takes place most
frequently.
Correa.
Campanula! sp. pl.
Page 123
Polemonium.
Phlox!
Cobœa!
Rhododendron!
Erica!
Rhodora.
Azalea!
Compositæ! sp. pl.
Lonicera!
Convolvulus!
Pharbitis.
Antirrhinum!
Verbascum!
Mimulus.
Digitalis!
Orobanche.
Solanum.
Nicotiana.
Gentiana!
Anagallis.
Primula!
Lamium!
Convallaria!
Lilium!
Colchicum!
&c. &c.
This list does not include those very numerous cases in which this change is
associated with more or less complete frondescence or leafy condition of
the petals.
Dialysis of the stamens.—A similar isolation of the stamens occurs
occasionally; for instance, when Mallows (Malvaceæ) become double, one
of the first stages of the process is often the disjunction of the stamens, and
a similar dissociation occurs in Leguminosæ and Compositæ, as in
Tragopogon, as related by Kirschleger, in Hypochæris by Wigand, and in
Coreopsis by Schlechtendal.
Phlox!
Cobœa!
Rhododendron!
Erica!
Rhodora.
Azalea!
Compositæ! sp. pl.
Lonicera!
Convolvulus!
Pharbitis.
Antirrhinum!
Verbascum!
Mimulus.
Digitalis!
Orobanche.
Solanum.
Nicotiana.
Gentiana!
Anagallis.
Primula!
Lamium!
Convallaria!
Lilium!
Colchicum!
&c. &c.
This list does not include those very numerous cases in which this change is
associated with more or less complete frondescence or leafy condition of
the petals.
Dialysis of the stamens.—A similar isolation of the stamens occurs
occasionally; for instance, when Mallows (Malvaceæ) become double, one
of the first stages of the process is often the disjunction of the stamens, and
a similar dissociation occurs in Leguminosæ and Compositæ, as in
Tragopogon, as related by Kirschleger, in Hypochæris by Wigand, and in
Coreopsis by Schlechtendal.
Page 124
Dialysis of the carpels.—In the case of the carpels this disunion is more
frequent than in the stamens. M. Seringe[78] figures carpels of Diplotaxis
tenuifolia more or less completely separated one from the other; indeed, this
separation is very common amongst Cruciferæ and Umbelliferæ.
Generally speaking, the disunion is complicated with frondescence—but
not always so. I have, in my herbarium, specimens of Convallaria majalis,
Commelyna sp., and of Lilium auratum, in all of which the three carpels are
completely disjoined, and present three styles, three stigmas, &c., without
any other change. Engelmann[79] speaks of three classes of this
malformation. 1st, that in which the carpels separate one from the other
without opening, as in the lily just alluded to; 2nd, that in which the ovary
remains closed, but loses its internal partitions, as in a case mentioned by
Moquin in Stachys sylvatica, in which, owing to imperfect disjunction, the
two bi-lobed carpels were changed into a nearly one-celled capsule;[80] and
3rd, those cases in which the carpels are open and foliaceous.
Fig. 32.—Anomalous form of orange.
frequent than in the stamens. M. Seringe[78] figures carpels of Diplotaxis
tenuifolia more or less completely separated one from the other; indeed, this
separation is very common amongst Cruciferæ and Umbelliferæ.
Generally speaking, the disunion is complicated with frondescence—but
not always so. I have, in my herbarium, specimens of Convallaria majalis,
Commelyna sp., and of Lilium auratum, in all of which the three carpels are
completely disjoined, and present three styles, three stigmas, &c., without
any other change. Engelmann[79] speaks of three classes of this
malformation. 1st, that in which the carpels separate one from the other
without opening, as in the lily just alluded to; 2nd, that in which the ovary
remains closed, but loses its internal partitions, as in a case mentioned by
Moquin in Stachys sylvatica, in which, owing to imperfect disjunction, the
two bi-lobed carpels were changed into a nearly one-celled capsule;[80] and
3rd, those cases in which the carpels are open and foliaceous.
Fig. 32.—Anomalous form of orange.
Page 125
Disjunction is more frequent in dry fruits than in fleshy ones. In the latter
instance it happens at an early stage of existence, and the pericarp becomes
more or less leafy, losing its faculty of becoming fleshy, as in Prunus
Cerasus and Amygdalus persica; nevertheless, fleshy fruits sometimes
become disunited. I have seen a case similar to that mentioned by M.
Alphonse de Candolle in Solanum esculentum, in which the pericarp
became ruptured, and the placentas protruded. A like occurrence has also
been observed in a species of Melastoma.[81] This is analogous to what
happens in Caulophyllum and Slateria. Disjunction of the carpels is not rare
in oranges. Sometimes this takes place regularly, at other times irregularly;
occasionally in such a manner as to give the appearance of a hand and
fingers to the fruit. Of one of these, Ferrari,[82] in the curious volume below
cited, speaks thus: "Arbor profusissima, quia dat utraque manu; imo quia
vere manus dat in poma conversis; utque magis munifica sit poma ipsa
convertit in manus."
M. Duchartre[83] mentions a semi-double flower of orange with eight to ten
distinct carpels in a whorl, and occasionally several whorls one above
another. De Candolle[84] considers the rind of the orange as a production
from the receptacle, and this view is confirmed by the specimens of
Duchartre, in which the carpels were quite naked or had a common
envelope truncated, and open above to allow of the passage of the styles
and stigmas.
instance it happens at an early stage of existence, and the pericarp becomes
more or less leafy, losing its faculty of becoming fleshy, as in Prunus
Cerasus and Amygdalus persica; nevertheless, fleshy fruits sometimes
become disunited. I have seen a case similar to that mentioned by M.
Alphonse de Candolle in Solanum esculentum, in which the pericarp
became ruptured, and the placentas protruded. A like occurrence has also
been observed in a species of Melastoma.[81] This is analogous to what
happens in Caulophyllum and Slateria. Disjunction of the carpels is not rare
in oranges. Sometimes this takes place regularly, at other times irregularly;
occasionally in such a manner as to give the appearance of a hand and
fingers to the fruit. Of one of these, Ferrari,[82] in the curious volume below
cited, speaks thus: "Arbor profusissima, quia dat utraque manu; imo quia
vere manus dat in poma conversis; utque magis munifica sit poma ipsa
convertit in manus."
M. Duchartre[83] mentions a semi-double flower of orange with eight to ten
distinct carpels in a whorl, and occasionally several whorls one above
another. De Candolle[84] considers the rind of the orange as a production
from the receptacle, and this view is confirmed by the specimens of
Duchartre, in which the carpels were quite naked or had a common
envelope truncated, and open above to allow of the passage of the styles
and stigmas.
Page 126
Fig. 33.—Orange. Showing disjunction of carpels, after Maout.
Page 127
Fig. 34.—Section of orange shown in fig. 33 after Maout.
It frequently happens in conjunction with this separation of the carpels one
from the other, that a lack of union manifests itself between the margins of
the individual carpels themselves. Very numerous cases of this kind have
been recorded, and the double tulips of gardens may be referred to as
showing this condition very frequently. In connection with this detachment
of the carpels, a change in the mode of placentation is often to be observed,
or two or more kinds may be seen in the same pistil, as in double-flowered
saponarias, many Crucifers, &c., as alluded to under the head of
displacements of the placenta.
FOOTNOTES:
[75] Loc. cit., p. 298.
[76] Masters in Seemann's 'Journal of Botany,' 1867, p. 158.
[77] Bull. Soc. Bot. France,' t. xiii, 1866, p. 234.
[78] 'Bull. Bot.,' pl. i, figs. 8–12.
[79] 'De Anthol.,' p. 37.
[80] Moquin, loc. cit., p. 305.
[81] 'Neue Denkschr. der Allg. Schweiz. Gesell.,' band v, pl. ii. p. 5.
[82] 'Hesperides,' auctore Ferrario. Rome, 1646, fig. 415, pp. 213 and 215. See also Michel,
'Traité du Citronnier.'
[83] 'Ann. des Science Nat.,' 3rd series, 1844, vol. i, p. 294.
[84] 'Org. Véget.,' vol. ii., p. 41.
CHAPTER III.
SOLUTION.
It frequently happens in conjunction with this separation of the carpels one
from the other, that a lack of union manifests itself between the margins of
the individual carpels themselves. Very numerous cases of this kind have
been recorded, and the double tulips of gardens may be referred to as
showing this condition very frequently. In connection with this detachment
of the carpels, a change in the mode of placentation is often to be observed,
or two or more kinds may be seen in the same pistil, as in double-flowered
saponarias, many Crucifers, &c., as alluded to under the head of
displacements of the placenta.
FOOTNOTES:
[75] Loc. cit., p. 298.
[76] Masters in Seemann's 'Journal of Botany,' 1867, p. 158.
[77] Bull. Soc. Bot. France,' t. xiii, 1866, p. 234.
[78] 'Bull. Bot.,' pl. i, figs. 8–12.
[79] 'De Anthol.,' p. 37.
[80] Moquin, loc. cit., p. 305.
[81] 'Neue Denkschr. der Allg. Schweiz. Gesell.,' band v, pl. ii. p. 5.
[82] 'Hesperides,' auctore Ferrario. Rome, 1646, fig. 415, pp. 213 and 215. See also Michel,
'Traité du Citronnier.'
[83] 'Ann. des Science Nat.,' 3rd series, 1844, vol. i, p. 294.
[84] 'Org. Véget.,' vol. ii., p. 41.
CHAPTER III.
SOLUTION.
Page 128
The isolation or separation of different whorls that are ordinarily adherent
together is by no means of rare occurrence. Were it not that the isolation is
often congenital, the word detachment would be an expressive one to apply
to these cases, but as the change in question occurs quite as often from a
want of union, an arrest or stasis of development, as from a bonâ fide
separation, the word solution seems to be, on the whole, the best. It
corresponds in application to the word liber (calyx liber, &c.), in general
use by descriptive botanists. As here employed, the term nearly corresponds
with the "adesmie hetérologue" of Morren. Moquin Tandon does not make
any special subdivision for the class of cases here grouped together, but
places them all under "Disjonctions qui isolent les organes." It seems,
however, desirable to have a separate word to express the converse
condition of adhesion, and for this purpose the term solution, as above
stated, is here employed. Diagrammatically, the condition may be expressed
by placing a dotted line at the side of the letters thus:
: S S S S S :
: C C C C C :
would indicate the disjunction of the sepals from the carpels (c), in
contradistinction to adhesion, which may be represented by the unbroken
line thus:
| S S S S S |
| C C C C C |
Solution of the calyx from the ovary.—Of all the instances of adhesion
which take place under ordinary circumstances, that between the calyx and
the ovary is perhaps the most common. The calyx adhærens or superus is a
structural characteristic to which all botanists attach considerable
importance; so that when exceptional cases occur in which the calyx
becomes detached from the ovary, becomes, that is, inferus or liber, a
together is by no means of rare occurrence. Were it not that the isolation is
often congenital, the word detachment would be an expressive one to apply
to these cases, but as the change in question occurs quite as often from a
want of union, an arrest or stasis of development, as from a bonâ fide
separation, the word solution seems to be, on the whole, the best. It
corresponds in application to the word liber (calyx liber, &c.), in general
use by descriptive botanists. As here employed, the term nearly corresponds
with the "adesmie hetérologue" of Morren. Moquin Tandon does not make
any special subdivision for the class of cases here grouped together, but
places them all under "Disjonctions qui isolent les organes." It seems,
however, desirable to have a separate word to express the converse
condition of adhesion, and for this purpose the term solution, as above
stated, is here employed. Diagrammatically, the condition may be expressed
by placing a dotted line at the side of the letters thus:
: S S S S S :
: C C C C C :
would indicate the disjunction of the sepals from the carpels (c), in
contradistinction to adhesion, which may be represented by the unbroken
line thus:
| S S S S S |
| C C C C C |
Solution of the calyx from the ovary.—Of all the instances of adhesion
which take place under ordinary circumstances, that between the calyx and
the ovary is perhaps the most common. The calyx adhærens or superus is a
structural characteristic to which all botanists attach considerable
importance; so that when exceptional cases occur in which the calyx
becomes detached from the ovary, becomes, that is, inferus or liber, a
Page 129
proportionate degree of interest attaches to the irregularity. It is not within
the scope of the present work to inquire whether this detachment be real or
merely apparent, arising from a want of union between parts ordinarily
united together. This point must be left to the organogenists to decide in
each particular case. So also the question as to what share, if any, the
expanded and dilated flower-stalk may take in what are usually called
inferior ovaries, can be here only incidentally touched upon.
Among Rosaceæ, the change in question is very common, especially in
conjunction with an elongation of the axis of the flower (apostasis) and with
prolification, though it is by no means always co-existent with these
malformations. When this alteration in the apparent relative position of
calyx and carpels occurs in roses (Rosa) the appearances are generally such
as to indicate that the "hip" of the rose is a dilatation of the peduncle,
continuous above with the coherent bases of the sepals; this inference seems
also to be borne out by what happens in the Pomaceæ. In some cases in this
sub-order, the calyx becomes detached from the carpels, so that the latter
organs become more or less "superior," and distinct one from the other. This
happens constantly in the double-flowered thorn, Cratægus Oxyacantha, in
some blossoms of which the hollowed end of the peduncle still invests the
base of the carpels, leaving the upper portions detached. In apples flowers
are occasionally met with of greater size than usual and on longer stalks, so
that the whole looks more like a rose than an apple blossom. In these cases
it will usually be found that the calyx consists of distinct sepals, without a
trace of the ordinary swelling beneath the flower. The petals are often more
numerous than usual; the stamens variously changed, and the carpels
sometimes absent; at other times, as in the instance figured in the adjacent
woodcuts, figs. 36, 37, consisting of separate, superior ovaries, sometimes
destitute of ovules, or, at other times, having two of these bodies.[85]
the scope of the present work to inquire whether this detachment be real or
merely apparent, arising from a want of union between parts ordinarily
united together. This point must be left to the organogenists to decide in
each particular case. So also the question as to what share, if any, the
expanded and dilated flower-stalk may take in what are usually called
inferior ovaries, can be here only incidentally touched upon.
Among Rosaceæ, the change in question is very common, especially in
conjunction with an elongation of the axis of the flower (apostasis) and with
prolification, though it is by no means always co-existent with these
malformations. When this alteration in the apparent relative position of
calyx and carpels occurs in roses (Rosa) the appearances are generally such
as to indicate that the "hip" of the rose is a dilatation of the peduncle,
continuous above with the coherent bases of the sepals; this inference seems
also to be borne out by what happens in the Pomaceæ. In some cases in this
sub-order, the calyx becomes detached from the carpels, so that the latter
organs become more or less "superior," and distinct one from the other. This
happens constantly in the double-flowered thorn, Cratægus Oxyacantha, in
some blossoms of which the hollowed end of the peduncle still invests the
base of the carpels, leaving the upper portions detached. In apples flowers
are occasionally met with of greater size than usual and on longer stalks, so
that the whole looks more like a rose than an apple blossom. In these cases
it will usually be found that the calyx consists of distinct sepals, without a
trace of the ordinary swelling beneath the flower. The petals are often more
numerous than usual; the stamens variously changed, and the carpels
sometimes absent; at other times, as in the instance figured in the adjacent
woodcuts, figs. 36, 37, consisting of separate, superior ovaries, sometimes
destitute of ovules, or, at other times, having two of these bodies.[85]
Page 130
Fig. 35.—Proliferous Rose. Showing an absence of the usual dilatation of the flower-stalk, and other
changes.
This condition accords precisely with the account of the development of the
flowers in Pomaceæ as given by Payer, Caspary, and others, so that the
flowers above described would owe their deficiency of the swollen
receptacle to an arrest of development. M. Germain de Saint Pierre, among
other malformations of the rose, presented to the Botanical Society of
France in 1854[86] two specimens which are of special interest as relating to
this contested point. In the one, the swollen portion beneath the flower was
surmounted by five perfect leaves, as, indeed, is not infrequent in such
malformations; here, then, the calyx could have had little or no share in the
production of the swelling in question. In the other, the swollen portion was
actually above the insertion of the sepals here represented by five perfect
leaves.
changes.
This condition accords precisely with the account of the development of the
flowers in Pomaceæ as given by Payer, Caspary, and others, so that the
flowers above described would owe their deficiency of the swollen
receptacle to an arrest of development. M. Germain de Saint Pierre, among
other malformations of the rose, presented to the Botanical Society of
France in 1854[86] two specimens which are of special interest as relating to
this contested point. In the one, the swollen portion beneath the flower was
surmounted by five perfect leaves, as, indeed, is not infrequent in such
malformations; here, then, the calyx could have had little or no share in the
production of the swelling in question. In the other, the swollen portion was
actually above the insertion of the sepals here represented by five perfect
leaves.
Page 131
Fig. 36.—Section through Apple blossom, showing detachment of calyx from ovaries, absence of
dilated flower-stalk, &c.
dilated flower-stalk, &c.
Page 132
Fig. 37.—Calyx detached from carpels in Apple.
On the other hand, M. Planchon's specimen of the Quince before alluded to,
not to mention other instances, tends to show that the bases of the sepals do
sometimes enter into the composition of the pome. And, indeed, in many of
these cases it would be impossible to say where the axial or receptacular
portion ended, and the foliar portion began. As both from normal
organogeny as well as from unusual conformation contradictory inferences
may be drawn, it would obviously be unsafe to attempt the explanation of
the so-called calyx-tube in general from any particular instances; so far as
Rosaceæ are concerned, there is so much variation in the relative position of
calyx and carpels under ordinary circumstances, that it is no matter for
surprise that similar diversities should exist in teratological cases. A similar
remark will apply to Saxifragaceæ, Cucurbitaceæ, Myrtaceæ, Bruniaceæ,
Rubiaceæ, and other families of like conformation.
On the other hand, M. Planchon's specimen of the Quince before alluded to,
not to mention other instances, tends to show that the bases of the sepals do
sometimes enter into the composition of the pome. And, indeed, in many of
these cases it would be impossible to say where the axial or receptacular
portion ended, and the foliar portion began. As both from normal
organogeny as well as from unusual conformation contradictory inferences
may be drawn, it would obviously be unsafe to attempt the explanation of
the so-called calyx-tube in general from any particular instances; so far as
Rosaceæ are concerned, there is so much variation in the relative position of
calyx and carpels under ordinary circumstances, that it is no matter for
surprise that similar diversities should exist in teratological cases. A similar
remark will apply to Saxifragaceæ, Cucurbitaceæ, Myrtaceæ, Bruniaceæ,
Rubiaceæ, and other families of like conformation.
Page 133
Fig. 38.—Flower of Œnanthe crocata, in which the five sepals were completely detached from the
ovaries, here three in number and destitute of stylopods.
In Umbelliferæ, a detachment of the calyx from the ovaries frequently
occurs, sometimes without any other change; at other times attended by
more serious alterations. So far as can be judged from exceptional
occurrences of this kind, it would appear that in this order the axis or
flower-stalk does not, in any material degree, enter into the composition of
the fruit.
In the Rubiaceous genus Bikkhia, as mentioned by Duchartre, the ovary is
completely inferior, but when the fruit arrives at maturity four small leaves
are detached from its surface which had previously adhered to it, and which
it seems reasonable to consider as the sepals.
In Campanulaceæ a similar separation of calyx from the ovary may be
occasionally met with. On the other hand, the occasional formation of a leaf
on the inferior ovary of those plants would indicate the axial nature of the
fruit. In Campanumæa and Cyclodon the calyx is inferior, while the corolla
ovaries, here three in number and destitute of stylopods.
In Umbelliferæ, a detachment of the calyx from the ovaries frequently
occurs, sometimes without any other change; at other times attended by
more serious alterations. So far as can be judged from exceptional
occurrences of this kind, it would appear that in this order the axis or
flower-stalk does not, in any material degree, enter into the composition of
the fruit.
In the Rubiaceous genus Bikkhia, as mentioned by Duchartre, the ovary is
completely inferior, but when the fruit arrives at maturity four small leaves
are detached from its surface which had previously adhered to it, and which
it seems reasonable to consider as the sepals.
In Campanulaceæ a similar separation of calyx from the ovary may be
occasionally met with. On the other hand, the occasional formation of a leaf
on the inferior ovary of those plants would indicate the axial nature of the
fruit. In Campanumæa and Cyclodon the calyx is inferior, while the corolla
Page 134
is superior. In the last-named genus this peculiarity "is carried to the highest
degree, the sepals being, in C. parviflorum, placed on the peduncle of the
flower far removed from the base of the corolla and ovary, whilst in C.
truncatum and in Campanumæa they adhere to the base of the tube of the
corolla."[87] In this order, then, as in Saxifragaceæ, Bruniaceæ, &c., no
hasty conclusion should be drawn as to the nature of the fruit. In Brunia
microphylla the ovary is superior, enclosed within but not adnate to the cup-
like calyx, to which latter, however, the petals and stamens are attached.
In Onagraceæ (Jussieua), as also in Cactaceæ (Opuntia), buds have been
observed on the surface and edges of the inferior ovary. Indeed, in the
former genus, they have been produced artificially, but as buds may be
formed on foliar as well as on axial organs, the fact cannot be made great
use of in support either of the foliar or axial nature of the inferior ovary. In
Epilobium, I have met with four perfect leaves at the summit of the ovary,
in the place usually occupied by the sepals. This would also favour the
notion that the axis entered into the constitution of the fruit in this genus.
Mr. B. Clarke, in his 'New Arrangement of Phanerogamous Plants,' p. 4,
cites a case wherein the perianth was completely detached from the surface
of the ovary in Cannabis sativa.
It must be borne in mind that some of the recorded instances of change in
the relative position of the calyx and pistil ought more properly to be
referred to a substitution of carpels for stamens, as in Begonia, Fuchsia, &c.
Among Cucurbitaceæ, examples have been recorded, both of the
detachment of the calyx from the ovary,[88] and of the partial conversion of
some of the anthers of the male flower to carpels.
The very singular mode of germination of Sechium edule in which the fruit,
instead of rotting, becomes thickened into a kind of rhizome or tuber, is a
degree, the sepals being, in C. parviflorum, placed on the peduncle of the
flower far removed from the base of the corolla and ovary, whilst in C.
truncatum and in Campanumæa they adhere to the base of the tube of the
corolla."[87] In this order, then, as in Saxifragaceæ, Bruniaceæ, &c., no
hasty conclusion should be drawn as to the nature of the fruit. In Brunia
microphylla the ovary is superior, enclosed within but not adnate to the cup-
like calyx, to which latter, however, the petals and stamens are attached.
In Onagraceæ (Jussieua), as also in Cactaceæ (Opuntia), buds have been
observed on the surface and edges of the inferior ovary. Indeed, in the
former genus, they have been produced artificially, but as buds may be
formed on foliar as well as on axial organs, the fact cannot be made great
use of in support either of the foliar or axial nature of the inferior ovary. In
Epilobium, I have met with four perfect leaves at the summit of the ovary,
in the place usually occupied by the sepals. This would also favour the
notion that the axis entered into the constitution of the fruit in this genus.
Mr. B. Clarke, in his 'New Arrangement of Phanerogamous Plants,' p. 4,
cites a case wherein the perianth was completely detached from the surface
of the ovary in Cannabis sativa.
It must be borne in mind that some of the recorded instances of change in
the relative position of the calyx and pistil ought more properly to be
referred to a substitution of carpels for stamens, as in Begonia, Fuchsia, &c.
Among Cucurbitaceæ, examples have been recorded, both of the
detachment of the calyx from the ovary,[88] and of the partial conversion of
some of the anthers of the male flower to carpels.
The very singular mode of germination of Sechium edule in which the fruit,
instead of rotting, becomes thickened into a kind of rhizome or tuber, is a
Page 135
fact that should not be overlooked in investigating the true nature of the
fruit in this order.
The following are the genera in which the change has been most frequently
observed:
*Rosa!
*Pyrus!
*Cratægus!
*Daucus!
Pastinaca.
Torilis.
Apium.
Œnanthe!
Heracleum,
Athamanta.
Selinum.
Carum.
Imperatoria.
Rudbeckia!
*Campanula!
Lonicera!
Cucumis!
Cannabis.
Solution of the stamens from the petals.—A separation of the stamens
from the petals in flowers, wherein those organs are usually adherent one to
the other does not often occur unattended by other changes. It has been
observed in Cobæa scandens (Turpin), in Antirrhinum majus, and in many
double flowers.
Partial detachment of the stamens from the styles occurs frequently in semi-
double flowers of Orchis.[89]
FOOTNOTES:
[85] 'Gard. Chron.,' 1865, p. 554; 1867, p. 599.
fruit in this order.
The following are the genera in which the change has been most frequently
observed:
*Rosa!
*Pyrus!
*Cratægus!
*Daucus!
Pastinaca.
Torilis.
Apium.
Œnanthe!
Heracleum,
Athamanta.
Selinum.
Carum.
Imperatoria.
Rudbeckia!
*Campanula!
Lonicera!
Cucumis!
Cannabis.
Solution of the stamens from the petals.—A separation of the stamens
from the petals in flowers, wherein those organs are usually adherent one to
the other does not often occur unattended by other changes. It has been
observed in Cobæa scandens (Turpin), in Antirrhinum majus, and in many
double flowers.
Partial detachment of the stamens from the styles occurs frequently in semi-
double flowers of Orchis.[89]
FOOTNOTES:
[85] 'Gard. Chron.,' 1865, p. 554; 1867, p. 599.
Page 136
[86] 'Bull. Soc. Bot. Fr.' 1854, p. 303.
[87] Hook et Thoms, 'Præcurs. ad Flor. Ind.,' Journ. Linn. Soc., vol. ii, 1858, p. 6.
[88] Lindley, 'Veget. Kingd.,' p. 315.
[89] Masters, 'Journal of Linnean Society,' 1866, vol. viii, p. 207. On the subject of this
chapter the reader should also consult Moquin-Tandon, 1. c., p. 298. Engelmann, 'De
Antholysi,' p. 37, tab. v. C. Morren, 'Bull. Acad. Belg.,' xix, part 3, p. 318. Cramer,
'Bildungsabweichungen,' p. 64. Fleischer, 'Missbild. Cultur. Pflanzen.' As to the nature of
inferior ovaries, see also Payer, 'Bull. Soc. Bot. Fr.' i, 1854, p. 283. Germain de Saint Pierre,
ibid., p. 302. Caspary, 'Bull. Soc. Bot. Fr.,' t. vi, 1859, p. 235. Schleiden, 'Principles of Botany,'
English translation, p. 368. Duchartre, 'Elements de Botanique,' p. 574. Le Maout et Decaisne,
'Traité général de Botanique,' p. 57. Bentham, 'Journ. Linn. Soc.,' vol. x, p. 104 (Structure of
Myrtaceæ), and other treatises on Organography.
PART III.
ALTERATIONS OF POSITION.
Necessarily connected with changes in the arrangement of organs are
similar alterations in their position; so closely, indeed, that but for
convenience sake, it would be unnecessary to treat them separately. There
are, however, some anomalous developments affecting the relative position
of organs that could hardly be treated of under any of the preceding
paragraphs. There are, also, certain rare instances where an organ is not so
much displaced as misplaced; that is to say, it is developed on or from a
portion of the plant, which under usual circumstances does not produce
such an organ. In the former instance, the altered position is due to or
coexistent with other changes, but in the latter case the new growth may
spring from organs otherwise in nowise different from ordinary. The word
Displacement is here used to signify the unusual position of an organ; while
Heterotaxy may serve to include those cases where a new growth makes its
[87] Hook et Thoms, 'Præcurs. ad Flor. Ind.,' Journ. Linn. Soc., vol. ii, 1858, p. 6.
[88] Lindley, 'Veget. Kingd.,' p. 315.
[89] Masters, 'Journal of Linnean Society,' 1866, vol. viii, p. 207. On the subject of this
chapter the reader should also consult Moquin-Tandon, 1. c., p. 298. Engelmann, 'De
Antholysi,' p. 37, tab. v. C. Morren, 'Bull. Acad. Belg.,' xix, part 3, p. 318. Cramer,
'Bildungsabweichungen,' p. 64. Fleischer, 'Missbild. Cultur. Pflanzen.' As to the nature of
inferior ovaries, see also Payer, 'Bull. Soc. Bot. Fr.' i, 1854, p. 283. Germain de Saint Pierre,
ibid., p. 302. Caspary, 'Bull. Soc. Bot. Fr.,' t. vi, 1859, p. 235. Schleiden, 'Principles of Botany,'
English translation, p. 368. Duchartre, 'Elements de Botanique,' p. 574. Le Maout et Decaisne,
'Traité général de Botanique,' p. 57. Bentham, 'Journ. Linn. Soc.,' vol. x, p. 104 (Structure of
Myrtaceæ), and other treatises on Organography.
PART III.
ALTERATIONS OF POSITION.
Necessarily connected with changes in the arrangement of organs are
similar alterations in their position; so closely, indeed, that but for
convenience sake, it would be unnecessary to treat them separately. There
are, however, some anomalous developments affecting the relative position
of organs that could hardly be treated of under any of the preceding
paragraphs. There are, also, certain rare instances where an organ is not so
much displaced as misplaced; that is to say, it is developed on or from a
portion of the plant, which under usual circumstances does not produce
such an organ. In the former instance, the altered position is due to or
coexistent with other changes, but in the latter case the new growth may
spring from organs otherwise in nowise different from ordinary. The word
Displacement is here used to signify the unusual position of an organ; while
Heterotaxy may serve to include those cases where a new growth makes its
Page 137
appearance in an unwonted situation, as, for instance, a leaf-bud on a root,
&c. Prolification is also included under this heading, the unusual position of
the buds in these cases being of graver import than the mere increase in
number. Alterations in the position of the sexual organs are spoken of under
the head of Heterogamy.
CHAPTER I.
DISPLACEMENT.
Real or apparent displacement of organs from their usual position is an
almost necessary consequence of, or is, at least, coexistent with a large
number of teratological phenomena. It is obvious that abnormal unions or
disunions, suppressions, hypertrophies, &c., are very liable to bring about
or to be accompanied with changes in the position, either of the parts
directly affected or of adjoining organs.
In this place, then, it is merely necessary to allude to some of the more
important displacements, and to refer for further details to the sections
relating to those irregularities of growth on which the displacement
depends.
Displacement of bulbs.—I owe to the kindness of Mr. James Salter a tulip
bulb which had been dug up after flowering, and from the base of which
were suspended several small bulbs; and I have since seen another
specimen showing the same unusual arrangement. The explanation of these
formations seems to be that they correspond to the bulbils ordinarily found
in the axils of the scales of the parent organ, and which, in some way or
another, have been displaced and thrust into the ground. Professor de Vriese
figures something of the same kind in Ixia carminosa.[90]
&c. Prolification is also included under this heading, the unusual position of
the buds in these cases being of graver import than the mere increase in
number. Alterations in the position of the sexual organs are spoken of under
the head of Heterogamy.
CHAPTER I.
DISPLACEMENT.
Real or apparent displacement of organs from their usual position is an
almost necessary consequence of, or is, at least, coexistent with a large
number of teratological phenomena. It is obvious that abnormal unions or
disunions, suppressions, hypertrophies, &c., are very liable to bring about
or to be accompanied with changes in the position, either of the parts
directly affected or of adjoining organs.
In this place, then, it is merely necessary to allude to some of the more
important displacements, and to refer for further details to the sections
relating to those irregularities of growth on which the displacement
depends.
Displacement of bulbs.—I owe to the kindness of Mr. James Salter a tulip
bulb which had been dug up after flowering, and from the base of which
were suspended several small bulbs; and I have since seen another
specimen showing the same unusual arrangement. The explanation of these
formations seems to be that they correspond to the bulbils ordinarily found
in the axils of the scales of the parent organ, and which, in some way or
another, have been displaced and thrust into the ground. Professor de Vriese
figures something of the same kind in Ixia carminosa.[90]
Page 138
Of somewhat different nature to those above described was an anomaly
described by M. Gay at a meeting of the Botanical Society of France, April
8th, 1859. The plant affected was Leucoium æstivum, and the changes
observed were apparently attributable to a simple separation of two leaves
that are usually contiguous. "Suppose," says M. Gay in describing this
malformation, "the first leaf of the terminal bud separated by a long
internode from the other leaves, which remain closely packed; and further,
suppose an evident thickening of the upper portion of the lengthened
internode, and there will be not only a single bulb, bearing with the leaves
of the present year all the remnants of the leaves of the two preceding years,
but two bulbs placed one above another, on the same axis, separated by the
length of the internode."
Fig. 39.—Unusual position of bulbs of tulip; the parent-bulb cut open.
The formation of bulbs in the axils of the leaves, as happens occasionally in
tulips, is further alluded to under the head of hypertrophy.
described by M. Gay at a meeting of the Botanical Society of France, April
8th, 1859. The plant affected was Leucoium æstivum, and the changes
observed were apparently attributable to a simple separation of two leaves
that are usually contiguous. "Suppose," says M. Gay in describing this
malformation, "the first leaf of the terminal bud separated by a long
internode from the other leaves, which remain closely packed; and further,
suppose an evident thickening of the upper portion of the lengthened
internode, and there will be not only a single bulb, bearing with the leaves
of the present year all the remnants of the leaves of the two preceding years,
but two bulbs placed one above another, on the same axis, separated by the
length of the internode."
Fig. 39.—Unusual position of bulbs of tulip; the parent-bulb cut open.
The formation of bulbs in the axils of the leaves, as happens occasionally in
tulips, is further alluded to under the head of hypertrophy.
Page 139
Displacements affecting the inflorescence.—These are, for the most part,
dependent on hypertrophy, elongation, atrophy, spiral torsion, &c., but there
are a few instances of a different nature, which may here be alluded to as
not being coincident with any of the phenomena just mentioned. Sometimes
these deviations from the ordinary position have the more interest as
affecting characters used to distinguish genera; thus one of the distinctions
between rye-grass (Lolium) and wheat (Triticum) resides in the relative
position of the spikelets and the main stem; in Triticum the spikelets are
placed with their backs against the rachis, in Lolium with one edge against
it; but in a specimen of rye-grass that has come under my own observation,
the arrangement was that of Triticum.
M. Kirschleger relates having found a specimen of Leucanthemum
pratense, in which the ligulate female flowers were growing singly in the
axils of the upper leaves of the stem.[91] The ordinary capitulum would here
seem to have been replaced by a spike or a raceme. A less degree of this
change wherein a few flowers may be found, as it were, detached from the
ordinary capitulum may often be observed in Compositæ, Dipsacaceæ, &c.
I have also met with specimens of Lamium album in which some of the
fascicles or clusters of flowers in place of being placed at the same level on
opposite sides of the stem were placed alternately one above another.
Caspary[92] mentions a flower of Aldrovanda vesiculosa, which was
elevated on a stalk that was adherent to the stem for a certain distance, and
then separated from it. This flower, with the leaf to which it was axillary,
evidently belonged to the whorl beneath, where there was a corresponding
deficiency. Another flower of the same plant bore on its pedicel a small leaf,
which was doubtless the bract raised above its ordinary position.
M. Fournier mentions an instance in Pelargonium grandiflorum, where,
owing to the lengthening of the axis, the pedicels, instead of being
dependent on hypertrophy, elongation, atrophy, spiral torsion, &c., but there
are a few instances of a different nature, which may here be alluded to as
not being coincident with any of the phenomena just mentioned. Sometimes
these deviations from the ordinary position have the more interest as
affecting characters used to distinguish genera; thus one of the distinctions
between rye-grass (Lolium) and wheat (Triticum) resides in the relative
position of the spikelets and the main stem; in Triticum the spikelets are
placed with their backs against the rachis, in Lolium with one edge against
it; but in a specimen of rye-grass that has come under my own observation,
the arrangement was that of Triticum.
M. Kirschleger relates having found a specimen of Leucanthemum
pratense, in which the ligulate female flowers were growing singly in the
axils of the upper leaves of the stem.[91] The ordinary capitulum would here
seem to have been replaced by a spike or a raceme. A less degree of this
change wherein a few flowers may be found, as it were, detached from the
ordinary capitulum may often be observed in Compositæ, Dipsacaceæ, &c.
I have also met with specimens of Lamium album in which some of the
fascicles or clusters of flowers in place of being placed at the same level on
opposite sides of the stem were placed alternately one above another.
Caspary[92] mentions a flower of Aldrovanda vesiculosa, which was
elevated on a stalk that was adherent to the stem for a certain distance, and
then separated from it. This flower, with the leaf to which it was axillary,
evidently belonged to the whorl beneath, where there was a corresponding
deficiency. Another flower of the same plant bore on its pedicel a small leaf,
which was doubtless the bract raised above its ordinary position.
M. Fournier mentions an instance in Pelargonium grandiflorum, where,
owing to the lengthening of the axis, the pedicels, instead of being
Page 140
umbellate, had become racemose; and I owe to the kindness of Dr. Sankey a
somewhat similar specimen, but in a less perfect condition. Here there was
but a single flower, and that rudimentary, placed at the extremity of the axis.
There were several bracts beneath this flower disposed spirally in the 1/3
arrangement, all being empty, excepting the terminal one. In like manner, a
head of flowers becomes sometimes converted into an umbel.
Displacement of leaves.—A cohesion of parts will sometimes give rise to
an apparent displacement, but the true nature of the malformation can, in
general, be readily made out.
Steinheil[93] found a specimen of Salvia Verbenaca, the leaves of which
presented very curious examples of displacement arising from cohesion.
Two of these leaves placed at the base of a branch were completely fused in
their lower thirds, and divided into two distinct lobes at the upper part; each
of these lobes seemed to be as large as the limb of an ordinary leaf. Above
these was another very broad one, apparently entire, but evidently produced
by a complete cohesion of two. This completely fused leaf alternated in
position with the imperfectly fused one below it; the alternation is explained
by supposing that the opposite leaves of each pair were directed one
towards the other, and became fused, and that thus resulted the
displacement. The dislocation of the organs took place in one direction for
one pair of leaves, and in another direction for the other pair, hence the
alternation. Thus, leaves normally opposite and decussate may, by fusion,
become alternate. A similar instance occurred to the writer in Lysimachia
vulgaris, wherein the changes arising from fusion and suppression of parts,
&c., were very considerable; as far as the leaves were concerned they
presented the following arrangement in succession from below upwards:—
first verticillate, then opposite, then spirally alternate, lastly opposite.[94]
The term "diremption" has sometimes been applied to cases where leaves
are thus apparently dragged out of position.
somewhat similar specimen, but in a less perfect condition. Here there was
but a single flower, and that rudimentary, placed at the extremity of the axis.
There were several bracts beneath this flower disposed spirally in the 1/3
arrangement, all being empty, excepting the terminal one. In like manner, a
head of flowers becomes sometimes converted into an umbel.
Displacement of leaves.—A cohesion of parts will sometimes give rise to
an apparent displacement, but the true nature of the malformation can, in
general, be readily made out.
Steinheil[93] found a specimen of Salvia Verbenaca, the leaves of which
presented very curious examples of displacement arising from cohesion.
Two of these leaves placed at the base of a branch were completely fused in
their lower thirds, and divided into two distinct lobes at the upper part; each
of these lobes seemed to be as large as the limb of an ordinary leaf. Above
these was another very broad one, apparently entire, but evidently produced
by a complete cohesion of two. This completely fused leaf alternated in
position with the imperfectly fused one below it; the alternation is explained
by supposing that the opposite leaves of each pair were directed one
towards the other, and became fused, and that thus resulted the
displacement. The dislocation of the organs took place in one direction for
one pair of leaves, and in another direction for the other pair, hence the
alternation. Thus, leaves normally opposite and decussate may, by fusion,
become alternate. A similar instance occurred to the writer in Lysimachia
vulgaris, wherein the changes arising from fusion and suppression of parts,
&c., were very considerable; as far as the leaves were concerned they
presented the following arrangement in succession from below upwards:—
first verticillate, then opposite, then spirally alternate, lastly opposite.[94]
The term "diremption" has sometimes been applied to cases where leaves
are thus apparently dragged out of position.
Page 141
In Tradescantia virginica I have met with opposite connate leaves; the
altered position, however, being due to the union of two stems.
Fig. 40.—Large-coloured leaf occupying the position of the inflorescence in Gesnera, after Morren.
altered position, however, being due to the union of two stems.
Fig. 40.—Large-coloured leaf occupying the position of the inflorescence in Gesnera, after Morren.
Page 142
Fig. 41.—Ordinary arrangement of leaves in fascicles of three in Pinus pinea and unusual
arrangement of leaves of same plant in spires.
Twisting of the stem is a frequent cause of the displacement of leaves (see
spiral torsion), as also hypertrophy, whether that excess of development
take place laterally or lengthwise (see elongation). Atrophy or suppression
will also frequently bring about an alteration in the position of leaves;
sometimes in such a manner that the place of the suppressed organ is
occupied by another one. One of the most curious instances of displacement
of leaves arising from suppression is that mentioned by Morren,[95] where,
in Gesnera Geroltiana, a large leaf apparently occupied the extremity of the
axis, a position which, under ordinary circumstances, no leaf could assume.
The explanation given by the Belgian professor is, that the axis in this case,
instead of throwing off a pair of leaves, one on each side, had from some
cause or another produced only one; this one not only being much larger
than ordinary, but brightly coloured, thus assuming some of the
characteristics as well as the position of the inflorescence.
arrangement of leaves of same plant in spires.
Twisting of the stem is a frequent cause of the displacement of leaves (see
spiral torsion), as also hypertrophy, whether that excess of development
take place laterally or lengthwise (see elongation). Atrophy or suppression
will also frequently bring about an alteration in the position of leaves;
sometimes in such a manner that the place of the suppressed organ is
occupied by another one. One of the most curious instances of displacement
of leaves arising from suppression is that mentioned by Morren,[95] where,
in Gesnera Geroltiana, a large leaf apparently occupied the extremity of the
axis, a position which, under ordinary circumstances, no leaf could assume.
The explanation given by the Belgian professor is, that the axis in this case,
instead of throwing off a pair of leaves, one on each side, had from some
cause or another produced only one; this one not only being much larger
than ordinary, but brightly coloured, thus assuming some of the
characteristics as well as the position of the inflorescence.
Page 143
Alterations in the usual arrangement of leaves, however, are not always
dependent on or coexistent with other teratological changes, but may simply
depend on a natural elongation of the internodes, or on fission or
multiplication; for instance, in some conifers, such as the Larch, (Abies
Larix) or Pinus pinea, there may be found at different stages in the growth
of the branches leaves in crowded fascicles or tufts; while, when growth is
more rapid, the leaves may be disposed in a spiral or alternate manner.
In the yew (Taxus) the leaves at the ends of the shoots not unfrequently lose
their usual distichous arrangement and become arranged in a close spiral
manner, the elongation of the shoot being arrested. This appears to be the
result of the injury effected by some insect.
Fig. 42.—Altered arrangement of leaves of yew, Taxus baccata.
So, too, the alteration from verticillate to spiral, or vice versâ, may take
place without any other notable change.[96] This may frequently be seen in
dependent on or coexistent with other teratological changes, but may simply
depend on a natural elongation of the internodes, or on fission or
multiplication; for instance, in some conifers, such as the Larch, (Abies
Larix) or Pinus pinea, there may be found at different stages in the growth
of the branches leaves in crowded fascicles or tufts; while, when growth is
more rapid, the leaves may be disposed in a spiral or alternate manner.
In the yew (Taxus) the leaves at the ends of the shoots not unfrequently lose
their usual distichous arrangement and become arranged in a close spiral
manner, the elongation of the shoot being arrested. This appears to be the
result of the injury effected by some insect.
Fig. 42.—Altered arrangement of leaves of yew, Taxus baccata.
So, too, the alteration from verticillate to spiral, or vice versâ, may take
place without any other notable change.[96] This may frequently be seen in
Page 144
Rhododendrons.
Displacement of the parts of the flower.—This subject is partly touched
on in the chapters on solution, adhesion, and in those on hypertrophy,
elongation, prolification, &c., so that in this place it is only requisite to offer
a few general remarks, and to refer to other sections for further details.
Morren, in referring to displacement of the floral organs, mentions an
instance in a Fuchsia, wherein the four petals in place of being alternate
with the sepals were placed in front of them, owing to the adhesion that had
taken place between the petals and the stamens. He speaks of this
transposition as metaphery.[97] The same author also gives an account of the
displacement of several of the organs of the flower in Cypripedium insigne,
the displacement being consequent, apparently, on a spiral torsion
proceeding from right to left, and involving the complete or partial
suppression of several of the organs of the flower. The dislocation of organs
in a spiral direction led Morren to apply the term "speiranthie" to similar
deviations from the usual construction. Changes of this kind among
Orchidaceæ are by no means uncommon; the following may be cited by
way of illustration. In a specimen of Oncidium cucullatum furnished me by
Mr. Anderson, well known for his success as a cultivator of these plants,
there was, associated with a cohesion of one sepal with another, and
probably dependent on the same cause, a displacement of the sepals and
petals—so that all were dragged out of place. This dislocation may be better
appreciated by the accompanying formula than even by the woodcut. Let
the usual arrangement be thus represented:
S
P ST P
L
S S
Displacement of the parts of the flower.—This subject is partly touched
on in the chapters on solution, adhesion, and in those on hypertrophy,
elongation, prolification, &c., so that in this place it is only requisite to offer
a few general remarks, and to refer to other sections for further details.
Morren, in referring to displacement of the floral organs, mentions an
instance in a Fuchsia, wherein the four petals in place of being alternate
with the sepals were placed in front of them, owing to the adhesion that had
taken place between the petals and the stamens. He speaks of this
transposition as metaphery.[97] The same author also gives an account of the
displacement of several of the organs of the flower in Cypripedium insigne,
the displacement being consequent, apparently, on a spiral torsion
proceeding from right to left, and involving the complete or partial
suppression of several of the organs of the flower. The dislocation of organs
in a spiral direction led Morren to apply the term "speiranthie" to similar
deviations from the usual construction. Changes of this kind among
Orchidaceæ are by no means uncommon; the following may be cited by
way of illustration. In a specimen of Oncidium cucullatum furnished me by
Mr. Anderson, well known for his success as a cultivator of these plants,
there was, associated with a cohesion of one sepal with another, and
probably dependent on the same cause, a displacement of the sepals and
petals—so that all were dragged out of place. This dislocation may be better
appreciated by the accompanying formula than even by the woodcut. Let
the usual arrangement be thus represented:
S
P ST P
L
S S
Page 145
S standing for sepal, P for petal, L for lip, ST for stamen; then the dislocated
form may be represented thus:
S
P P
T
S
S_S
L
Fig. 43.—Flower of Oncidium cucullatum, showing union of two lower sepals, displacement of
column and lip, &c.
In a specimen of Cypripedium also furnished by Mr. Anderson the
appearance was as represented in the accompanying figure and diagrams,
figs. 44, 45. Referring to the plan of the natural arrangement at fig. 46, it
will be seen that an explanation of the peculiar appearance of the flower
form may be represented thus:
S
P P
T
S
S_S
L
Fig. 43.—Flower of Oncidium cucullatum, showing union of two lower sepals, displacement of
column and lip, &c.
In a specimen of Cypripedium also furnished by Mr. Anderson the
appearance was as represented in the accompanying figure and diagrams,
figs. 44, 45. Referring to the plan of the natural arrangement at fig. 46, it
will be seen that an explanation of the peculiar appearance of the flower
Page 146
may be arrived at by supposing a disunion and lateral displacement of the
upper segment of the outer perianth together with the complete absence of
the lower one. In the second or inner whorl of the perianth the lip is merely
a little oblique on one side, but the lateral petals are distorted, displaced,
and adherent one to the other and to the column, while the posterior shield-
like rudimentary anther is completely wanting.
Fig. 44.—Malformed flower of Cypripedium.
upper segment of the outer perianth together with the complete absence of
the lower one. In the second or inner whorl of the perianth the lip is merely
a little oblique on one side, but the lateral petals are distorted, displaced,
and adherent one to the other and to the column, while the posterior shield-
like rudimentary anther is completely wanting.
Fig. 44.—Malformed flower of Cypripedium.
Page 147
Fig. 45.—Diagram of malformed Cypripedium. o, outer segments; i, inner segments of perianth; e,
lip; s, stigma; a, anther.
Fig. 46—- Diagram showing ordinary arrangement in Cypripedium. o, outer, i, inner segments of
perianth; e, lip, a, anther, a', abortive stamen; s, stigma.
lip; s, stigma; a, anther.
Fig. 46—- Diagram showing ordinary arrangement in Cypripedium. o, outer, i, inner segments of
perianth; e, lip, a, anther, a', abortive stamen; s, stigma.
Page 148
Fig. 47.—Plan of flower of Lycaste Skinneri showing displacement of organs.
In a specimen of Lycaste Skinneri similar changes were observed, as shown
in the plan, fig. 47. Here the posterior sepal was deficient, the two lateral
ones were present, one of them with a long tubular spur, o o; of the two
lateral petals, i i, one was twisted out of place, so as partially to occupy the
place of the deficient sepal; the lip was represented by two three-lobed
segments, l, one above and within the other. The column and ovary of this
flower were in their normal condition.
Cohesion of two or more segments of the perianth is frequently associated
with displacements of this nature: thus, in a flower of Dendrobium nobile, a
diagram of which is given at fig. 48, the uppermost sepal was coherent with
one of the lateral ones, and at the same time diminished in size, and, as it
were, dragged out of position. All the other organs of the flower are also
more or less displaced, forming a minor degree of the change already
alluded to, and which Morren termed speiranthy. The changes will be better
appreciated by comparing them with fig. 49, a diagram showing the natural
arrangement of parts in this species.
In a specimen of Lycaste Skinneri similar changes were observed, as shown
in the plan, fig. 47. Here the posterior sepal was deficient, the two lateral
ones were present, one of them with a long tubular spur, o o; of the two
lateral petals, i i, one was twisted out of place, so as partially to occupy the
place of the deficient sepal; the lip was represented by two three-lobed
segments, l, one above and within the other. The column and ovary of this
flower were in their normal condition.
Cohesion of two or more segments of the perianth is frequently associated
with displacements of this nature: thus, in a flower of Dendrobium nobile, a
diagram of which is given at fig. 48, the uppermost sepal was coherent with
one of the lateral ones, and at the same time diminished in size, and, as it
were, dragged out of position. All the other organs of the flower are also
more or less displaced, forming a minor degree of the change already
alluded to, and which Morren termed speiranthy. The changes will be better
appreciated by comparing them with fig. 49, a diagram showing the natural
arrangement of parts in this species.
Page 149
Fig. 48.—Plan of malformed flower of Dendrobium nobile.
Fig. 49.—Plan of natural arrangement in Dendrobium nobile. The x x represent processes of the
column, perhaps rudiments of stamens.
Sometimes the displacement seems consequent on hypertrophy of one of
the parts of the flower, the disproportionate size of one organ pushing the
Fig. 49.—Plan of natural arrangement in Dendrobium nobile. The x x represent processes of the
column, perhaps rudiments of stamens.
Sometimes the displacement seems consequent on hypertrophy of one of
the parts of the flower, the disproportionate size of one organ pushing the
Page 150
others out of place. This was the case in a violet, fig. 50, in which one of the
sepals s was greatly thickened, and the petals and stamens were displaced in
consequence.
Fig. 50.—Plan of flower of violet showing displacement of petals, &c. At b was a rudiment of a
stamen.
It is curious to observe in many of these cases that the transposed organ not
only occupies the place of a suppressed or abortive organ, but frequently
assumes its colour, and, to some extent, its function. This has been alluded
to in the case of the leaf of Gesnera (see p. 88) and in Orchids this
replacement seems to be very common; thus, in addition to the cases before
mentioned, in a flower of an Odontoglossum, for which I am indebted to
Professor Oliver, the two lateral sepals were united together and occupied
the position of the labellum, which was absent. A similar occurrence
happens occasionally in Lycaste Skinneri, thus recalling the structure of
Masdevallia, where the labellum is normally very small. The arrangement
in Lycaste may thus be symbolised:
S
P st P
+
sepals s was greatly thickened, and the petals and stamens were displaced in
consequence.
Fig. 50.—Plan of flower of violet showing displacement of petals, &c. At b was a rudiment of a
stamen.
It is curious to observe in many of these cases that the transposed organ not
only occupies the place of a suppressed or abortive organ, but frequently
assumes its colour, and, to some extent, its function. This has been alluded
to in the case of the leaf of Gesnera (see p. 88) and in Orchids this
replacement seems to be very common; thus, in addition to the cases before
mentioned, in a flower of an Odontoglossum, for which I am indebted to
Professor Oliver, the two lateral sepals were united together and occupied
the position of the labellum, which was absent. A similar occurrence
happens occasionally in Lycaste Skinneri, thus recalling the structure of
Masdevallia, where the labellum is normally very small. The arrangement
in Lycaste may thus be symbolised:
S
P st P
+
Page 151
S S
---
[Transcriber's note: The underscores represent a horizontal curly brace in
the original.]
the + indicating the position of the absent labellum.
Cases of this kind are the more interesting from their relation to the
fertilization of these flowers by insects; it seems as though, when the
labellum, which performs so important an office in attracting and guiding
insects, is deficient, its place is supplied by other means.
Displacement of the parts of the flower from elongation of the receptacle is
a not infrequent teratological occurrence, resulting sometimes in the
conversion of the verticillate into the spiral arrangement. Instances of this
are cited under Elongation, Prolification, &c. In this place it is merely
necessary to refer to a curious circumstance that is met with in some double
flowers, owing to this separation of some parts of the flower and the
cohesion or adhesion of others. Thus, in some double flowers of Primula
sinensis and in the Pea (Pisum sativum), I have seen a gradual passage of
sepals to petals, so that the calyx and corolla formed one continuous sheet,
winding spirally around the central axis of the flower, after the fashion of a
spiral tube.[98]
Displacement of the carpels arises from one or other of the causes above
alluded to, and when suppression takes place in this whorl it generally
happens that the place of the suppressed organ is occupied by one of the
remaining ones, which thus becomes partially dislocated.
Displacement of the placentas and ovules is a necessary result of many of
the changes to which the carpels are subject. The disjunction or dialysis of
the carpels, for instance, frequently renders axile placentation marginal.
---
[Transcriber's note: The underscores represent a horizontal curly brace in
the original.]
the + indicating the position of the absent labellum.
Cases of this kind are the more interesting from their relation to the
fertilization of these flowers by insects; it seems as though, when the
labellum, which performs so important an office in attracting and guiding
insects, is deficient, its place is supplied by other means.
Displacement of the parts of the flower from elongation of the receptacle is
a not infrequent teratological occurrence, resulting sometimes in the
conversion of the verticillate into the spiral arrangement. Instances of this
are cited under Elongation, Prolification, &c. In this place it is merely
necessary to refer to a curious circumstance that is met with in some double
flowers, owing to this separation of some parts of the flower and the
cohesion or adhesion of others. Thus, in some double flowers of Primula
sinensis and in the Pea (Pisum sativum), I have seen a gradual passage of
sepals to petals, so that the calyx and corolla formed one continuous sheet,
winding spirally around the central axis of the flower, after the fashion of a
spiral tube.[98]
Displacement of the carpels arises from one or other of the causes above
alluded to, and when suppression takes place in this whorl it generally
happens that the place of the suppressed organ is occupied by one of the
remaining ones, which thus becomes partially dislocated.
Displacement of the placentas and ovules is a necessary result of many of
the changes to which the carpels are subject. The disjunction or dialysis of
the carpels, for instance, frequently renders axile placentation marginal.
Page 152
Moreover, it frequently happens, when the carpels become foliaceous and
their margins are disconnected, that the ovules, in place of being placed on
the suture, or rather on the margins of the altered carpel, are placed on the
surface of the expanded carpel. Thus, in some double flowers of
Ranunculus Ficaria that came under the writer's notice the carpels were
open, i.e. disunited at the margins, and each bore two imperfect ovules upon
its inner surface a little way above the base, and midway between the edges
of the carpel and the midrib, the ovules being partly enclosed within a little
depression or pouch, similar to the pit on the petals. On closer examination
the ovules were found to spring from the two lateral divisions of the midrib,
the vascular cords of which were prolonged under the form of barred or
spiral fusiform tubes into the outer coating of the ovule. In this instance,
then, the ovules did not originate from the margins of the leaf, nor from a
prolonged axis, but they seemed to spring, in the guise of little buds, from
the inner surface of the carpellary leaf.[99]
The occurrence, also, of different forms of placentation in different flowers
on the same plant is no unusual thing in malformed flowers; thus, in double
flowers of Saponaria officinalis I have met with sutural, parietal, and free
central placentation in the same plant.[100]
Professor Babington describes in the 'Gardeners' Chronicle,' 1844, p. 557, a
curious flower of Cerastium, in which, in addition to other changes, the five
carpellary leaves "were partially turned in without touching the placenta,
which bears a cluster of ovules, and is perfectly clear of all connection with
those partitions" (fig. 51). See also Lindley, 'Veg. Kingdom,' p. 497.
their margins are disconnected, that the ovules, in place of being placed on
the suture, or rather on the margins of the altered carpel, are placed on the
surface of the expanded carpel. Thus, in some double flowers of
Ranunculus Ficaria that came under the writer's notice the carpels were
open, i.e. disunited at the margins, and each bore two imperfect ovules upon
its inner surface a little way above the base, and midway between the edges
of the carpel and the midrib, the ovules being partly enclosed within a little
depression or pouch, similar to the pit on the petals. On closer examination
the ovules were found to spring from the two lateral divisions of the midrib,
the vascular cords of which were prolonged under the form of barred or
spiral fusiform tubes into the outer coating of the ovule. In this instance,
then, the ovules did not originate from the margins of the leaf, nor from a
prolonged axis, but they seemed to spring, in the guise of little buds, from
the inner surface of the carpellary leaf.[99]
The occurrence, also, of different forms of placentation in different flowers
on the same plant is no unusual thing in malformed flowers; thus, in double
flowers of Saponaria officinalis I have met with sutural, parietal, and free
central placentation in the same plant.[100]
Professor Babington describes in the 'Gardeners' Chronicle,' 1844, p. 557, a
curious flower of Cerastium, in which, in addition to other changes, the five
carpellary leaves "were partially turned in without touching the placenta,
which bears a cluster of ovules, and is perfectly clear of all connection with
those partitions" (fig. 51). See also Lindley, 'Veg. Kingdom,' p. 497.
Page 153
Fig. 51.—1. Monstrous flower of a Cerastium; sepals and petals leafy. 2. Stamens and pistils
separate. 3. Ovary cut open to show the imperfect dissepiments and the attachment of the ovules. 4. A
deformed ovule.
M. Baillon[101] records flowers of Bunias, some with ovules on the margins
of the carpels, others with a central branch bearing the ovules; hence he
concludes very justly that no fair inference can be drawn from these facts as
to the normal placentation of Cruciferæ.
The same excellent observer has recorded the occurrence of free central
placentation in malformed flowers of Trifolium repens.[102]
In malformed flowers of Digitalis the change from axile to parietal
placentation may often be seen. Mr. Berkeley describes an instance of this
nature where the placentas were strictly parietal, and therefore receded from
the distinctive characters of the order, and approximated to those of
Gesneraceæ.
separate. 3. Ovary cut open to show the imperfect dissepiments and the attachment of the ovules. 4. A
deformed ovule.
M. Baillon[101] records flowers of Bunias, some with ovules on the margins
of the carpels, others with a central branch bearing the ovules; hence he
concludes very justly that no fair inference can be drawn from these facts as
to the normal placentation of Cruciferæ.
The same excellent observer has recorded the occurrence of free central
placentation in malformed flowers of Trifolium repens.[102]
In malformed flowers of Digitalis the change from axile to parietal
placentation may often be seen. Mr. Berkeley describes an instance of this
nature where the placentas were strictly parietal, and therefore receded from
the distinctive characters of the order, and approximated to those of
Gesneraceæ.
Page 154
The same author alludes to certain changes in the same flower where two
open carpels "were soldered together laterally, as was clear by the
rudiments of two styles, the placenta being produced only at the two united
edges, the outer margins remaining in the normal condition. This may
possibly tend to the explanation of some cases of anomalous placentation,
for the only indication of the true nature of the placentation is afforded by
the two rudimentary styles, in the absence of which the spongy receptacle
of the seeds must have been supposed to spring from the medial nerve."
In other cases the placentas were parietal above, but axile at the base of the
capsule, a striking instance of the facility with which axile placentation
becomes parietal, the change being here effected by the prolongation of the
axis, and the formation on it of a second whorl of carpellary leaves.
In double flowers of Primulaceæ similar alterations in the placentation may
often be observed. I have seen in Primula sinensis sutural, parietal, axile,
and free central placentation all on the same plant; nay, even in the same
capsule the ovules may be attached in various ways, and transitions from
one form of placentation to another are not infrequent. The late Professor E.
Forbes describes[103] an instance of true foliar and true axile placentation in
the same flower in Vinca minor.
These and many similar changes, which it is not necessary further to allude
to, are not so much to be wondered at when it is borne in mind how slight
an alteration suffices to produce a change in the mode of placentation, and
how frequent is the production of adventitious buds or of foliar outgrowths,
as may be seen in the sections relating to those subjects and to
Substitutions.
It will be remembered, also, how, in certain natural orders, under ordinary
circumstances, considerable diversity in placentation exists, according as
open carpels "were soldered together laterally, as was clear by the
rudiments of two styles, the placenta being produced only at the two united
edges, the outer margins remaining in the normal condition. This may
possibly tend to the explanation of some cases of anomalous placentation,
for the only indication of the true nature of the placentation is afforded by
the two rudimentary styles, in the absence of which the spongy receptacle
of the seeds must have been supposed to spring from the medial nerve."
In other cases the placentas were parietal above, but axile at the base of the
capsule, a striking instance of the facility with which axile placentation
becomes parietal, the change being here effected by the prolongation of the
axis, and the formation on it of a second whorl of carpellary leaves.
In double flowers of Primulaceæ similar alterations in the placentation may
often be observed. I have seen in Primula sinensis sutural, parietal, axile,
and free central placentation all on the same plant; nay, even in the same
capsule the ovules may be attached in various ways, and transitions from
one form of placentation to another are not infrequent. The late Professor E.
Forbes describes[103] an instance of true foliar and true axile placentation in
the same flower in Vinca minor.
These and many similar changes, which it is not necessary further to allude
to, are not so much to be wondered at when it is borne in mind how slight
an alteration suffices to produce a change in the mode of placentation, and
how frequent is the production of adventitious buds or of foliar outgrowths,
as may be seen in the sections relating to those subjects and to
Substitutions.
It will be remembered, also, how, in certain natural orders, under ordinary
circumstances, considerable diversity in placentation exists, according as
Page 155
the margins of the carpels are merely valvate or are infolded so as to reach
the centre. Often this diversity is due merely to the changes that take place
during growth; thus, the placentation of Caryophylleæ, Cucurbitaceæ,
Papaveraceæ, and many other orders, varies according to the age of the
carpel, and if any stasis or arrest of development occurs the placentation
becomes altered accordingly.
It is not necessary, in this place, to enter into the question whether the
placenta is, in all cases whatsoever, a dependence of the axis, as Payer,
Schleiden, and others, have maintained, or whether it be foliar in some
cases, axial in others. This question must be decided by the organogenists;
teratologically, however, there can be no doubt that ovules may be formed
from both foliar and axial organs, and, moreover, that, owing to the
variability above referred to, both in what are called natural and in what are
deemed abnormal conditions, it can rarely happen that any safe inferences
as to the normal or typical placentation of any family of plants can be
drawn from exceptional or monstrous formations.
On the subject of placentation the following authors may be consulted:
R. Brown, 'Ann. Nat. Hist.,' 1843, vol. xi, 35. Brongniart, 'Ann. Sc. Nat.,' 1834, sér. 2. i, p.
308. Alph. De Candolle, 'Neue Denkschrift der Allg. Schweizer Gesellsch.,' Band v. 1841,
p. 9. Duchartre, 'Ann. Sc. Nat.,' 3rd ser., 1844, vol. ii, p. 290. Ibid., 'Elem. Bot.,' p. 574;
'Rev. Bot.,' 1846–7, p. 213. Babington, 'Gard. Chron.,' 1844, p. 557. Lindley, 'Elements,' p.
89; 'Veg. King.,' pp. 313, 497, &c. Berkeley, 'Gard. Chron.,' 1850, p. 612. Unger, 'Nov. Act.
Acad. Nat. Cur.,' 1850; and in Henfrey's, 'Bot. Gazette,' 1851, p. 70. Schleiden, 'Principles,'
English edit., p. 385. Payer, 'Elem. Bot.,' pp. 196, 211, 224. Baillon, 'Adansonia.' iii, p. 310.
tab. iv. Cramer, 'Bildungsabweichungen,' p. 20, &c. Clos, 'Ann. Sc. Nat.,' 5th ser., iii, 313,
as well as any of the general treatises on botany. Reference may also be made to the
chapters on Prolification and Substitutions (in the case of the carpels and ovules), and to
the authorities therein cited.
FOOTNOTES:
[90] 'Tijdschr. voor. nat. Gesch.,' viii, 1841. tab. ii, p. 178.
the centre. Often this diversity is due merely to the changes that take place
during growth; thus, the placentation of Caryophylleæ, Cucurbitaceæ,
Papaveraceæ, and many other orders, varies according to the age of the
carpel, and if any stasis or arrest of development occurs the placentation
becomes altered accordingly.
It is not necessary, in this place, to enter into the question whether the
placenta is, in all cases whatsoever, a dependence of the axis, as Payer,
Schleiden, and others, have maintained, or whether it be foliar in some
cases, axial in others. This question must be decided by the organogenists;
teratologically, however, there can be no doubt that ovules may be formed
from both foliar and axial organs, and, moreover, that, owing to the
variability above referred to, both in what are called natural and in what are
deemed abnormal conditions, it can rarely happen that any safe inferences
as to the normal or typical placentation of any family of plants can be
drawn from exceptional or monstrous formations.
On the subject of placentation the following authors may be consulted:
R. Brown, 'Ann. Nat. Hist.,' 1843, vol. xi, 35. Brongniart, 'Ann. Sc. Nat.,' 1834, sér. 2. i, p.
308. Alph. De Candolle, 'Neue Denkschrift der Allg. Schweizer Gesellsch.,' Band v. 1841,
p. 9. Duchartre, 'Ann. Sc. Nat.,' 3rd ser., 1844, vol. ii, p. 290. Ibid., 'Elem. Bot.,' p. 574;
'Rev. Bot.,' 1846–7, p. 213. Babington, 'Gard. Chron.,' 1844, p. 557. Lindley, 'Elements,' p.
89; 'Veg. King.,' pp. 313, 497, &c. Berkeley, 'Gard. Chron.,' 1850, p. 612. Unger, 'Nov. Act.
Acad. Nat. Cur.,' 1850; and in Henfrey's, 'Bot. Gazette,' 1851, p. 70. Schleiden, 'Principles,'
English edit., p. 385. Payer, 'Elem. Bot.,' pp. 196, 211, 224. Baillon, 'Adansonia.' iii, p. 310.
tab. iv. Cramer, 'Bildungsabweichungen,' p. 20, &c. Clos, 'Ann. Sc. Nat.,' 5th ser., iii, 313,
as well as any of the general treatises on botany. Reference may also be made to the
chapters on Prolification and Substitutions (in the case of the carpels and ovules), and to
the authorities therein cited.
FOOTNOTES:
[90] 'Tijdschr. voor. nat. Gesch.,' viii, 1841. tab. ii, p. 178.
Page 156
[91] Communication to the Internat. Bot. Congress, Paris, 1867.
[92] 'Bot. Zeit.,' 1859, p. 117, tab. v.
[93] 'Ann. Sc. Nat.,' ser. 2, vol. iv, 1835, p. 143. tab. v.
[94] See Kirschleger, 'Flora.' 1844. p. 566 (Scabiosa).
[95] 'Bull. Acad. Belg.,' t. xvii. part ii, p. 387.
[96] 'Clos. Mem. Acad. Toulouse,' 5th ser., t. vi. pp. 51, 70.
[97] 'Bull. Acad. Roy. Belg.,' xviii. part ii, p. 505, and vol. xvii, part i, p. 196, and vol. xix.
part i. p. 260.
[98] See also Schlechtendal, 'Bot. Zeit.,' iv, p. 804. Primula veris, partibus perigonii spiræ
in modum confluentibus.
[99] Seemann's 'Journal of Botany,' vol. v, 1867, p. 158.
[100] 'Journ. Linn. Soc.,' i, 1857, p. 161. c. xylog.
[101] 'Adansonia,' ii, 306.
[102] 'Adansonia,' iv, p. 70, t. i.
[103] Henfrey's 'Bot. Gazette,' i, 265.
CHAPTER II.
PROLIFICATION.
Moquin-Tandon and other writers have classed the production of buds in
unwonted situations under the head of multiplication, but, as the altered
arrangement is of graver import than the mere increase in number, it seems
preferable to place these cases under this heading rather than under that of
alterations of number.
[92] 'Bot. Zeit.,' 1859, p. 117, tab. v.
[93] 'Ann. Sc. Nat.,' ser. 2, vol. iv, 1835, p. 143. tab. v.
[94] See Kirschleger, 'Flora.' 1844. p. 566 (Scabiosa).
[95] 'Bull. Acad. Belg.,' t. xvii. part ii, p. 387.
[96] 'Clos. Mem. Acad. Toulouse,' 5th ser., t. vi. pp. 51, 70.
[97] 'Bull. Acad. Roy. Belg.,' xviii. part ii, p. 505, and vol. xvii, part i, p. 196, and vol. xix.
part i. p. 260.
[98] See also Schlechtendal, 'Bot. Zeit.,' iv, p. 804. Primula veris, partibus perigonii spiræ
in modum confluentibus.
[99] Seemann's 'Journal of Botany,' vol. v, 1867, p. 158.
[100] 'Journ. Linn. Soc.,' i, 1857, p. 161. c. xylog.
[101] 'Adansonia,' ii, 306.
[102] 'Adansonia,' iv, p. 70, t. i.
[103] Henfrey's 'Bot. Gazette,' i, 265.
CHAPTER II.
PROLIFICATION.
Moquin-Tandon and other writers have classed the production of buds in
unwonted situations under the head of multiplication, but, as the altered
arrangement is of graver import than the mere increase in number, it seems
preferable to place these cases under this heading rather than under that of
alterations of number.
Page 157
The adventitious bud may be a leaf-bud or a flower-bud; it may occupy the
centre of a flower, thus terminating the axis, or it may be axillary to some or
other of its component parts, or, again, it may be extra-floral. In this last
case the prolification is of the inflorescence, and is hardly distinguishable
from multiplication or subdivision of the common flower-stalk. In
accordance with these differences we have median, axillary, and extra-floral
prolification, each admitting of subdivision into a leafy or a floral variety,
according to the nature of the adventitious bud. Under the head of each
variety certain special peculiarities are noticed, but it may here be advisable
to add a few general remarks on the subject.
Axillary prolification is a much less frequent malformation than the median
form. If only the number of orders and genera be reckoned, the truth of this
statement will be scarcely recognised; but if individual cases could be
estimated, the difference in frequency between the two would be very much
more obvious. This may, perhaps, be explained by the fact that the branch
has a greater tendency to grow in length than it has to develop buds from
the axils of the leaves. The flower is admitted to be homologous with the
branch, and it is also known that, up to a certain time, the branch-bud or
leaf-bud and the flower-bud do not essentially differ.[104] At a later stage the
difference between the two is manifested, not only in the altered form of the
lateral organs in the flower-bud, but in the tendency to an arrest of growth,
thus limiting the length of the central axial portion. Now, in prolified
flowers the functions and, to a considerable extent, the appearance of a leaf-
bud or of a branch are assumed, and with them the tendency to grow in
length is developed. Median prolification, therefore, in this sense, is a
further step in retrograde metamorphosis than is the axillary form. To grow
in length, and to produce axillary buds, are alike attributes of the branch;
but the former is much more frequently called into play than the latter; for
the same reason, median prolification is more common than the axillary
form. This is borne out by the frequency with which apostasis, or the
centre of a flower, thus terminating the axis, or it may be axillary to some or
other of its component parts, or, again, it may be extra-floral. In this last
case the prolification is of the inflorescence, and is hardly distinguishable
from multiplication or subdivision of the common flower-stalk. In
accordance with these differences we have median, axillary, and extra-floral
prolification, each admitting of subdivision into a leafy or a floral variety,
according to the nature of the adventitious bud. Under the head of each
variety certain special peculiarities are noticed, but it may here be advisable
to add a few general remarks on the subject.
Axillary prolification is a much less frequent malformation than the median
form. If only the number of orders and genera be reckoned, the truth of this
statement will be scarcely recognised; but if individual cases could be
estimated, the difference in frequency between the two would be very much
more obvious. This may, perhaps, be explained by the fact that the branch
has a greater tendency to grow in length than it has to develop buds from
the axils of the leaves. The flower is admitted to be homologous with the
branch, and it is also known that, up to a certain time, the branch-bud or
leaf-bud and the flower-bud do not essentially differ.[104] At a later stage the
difference between the two is manifested, not only in the altered form of the
lateral organs in the flower-bud, but in the tendency to an arrest of growth,
thus limiting the length of the central axial portion. Now, in prolified
flowers the functions and, to a considerable extent, the appearance of a leaf-
bud or of a branch are assumed, and with them the tendency to grow in
length is developed. Median prolification, therefore, in this sense, is a
further step in retrograde metamorphosis than is the axillary form. To grow
in length, and to produce axillary buds, are alike attributes of the branch;
but the former is much more frequently called into play than the latter; for
the same reason, median prolification is more common than the axillary
form. This is borne out by the frequency with which apostasis, or the
Page 158
separation of the floral whorls one from another, to a greater degree than
usual, is met with in prolified flowers.
In both forms the adventitious growth is much more frequently a flower-
bud or an inflorescence than a leaf-bud or a branch. This may be due to the
position of the flowers on a portion of the stem of the plant especially
devoted to the formation of flower-buds, to the more or less complete
exclusion of leaf-buds, i.e. on the inflorescence. This conjecture is borne
out by the comparative rarity with which prolification has been observed in
flowers that are solitary in the axils of the ordinary leaves of the plant. If the
lists of genera appended hereto be perused, it will be seen that nearly all the
cases occur in genera where the inflorescence is distinctly separated from
the other branches of the stem. In direct proportion, then, to the degree in
which one region of the axis or certain branches of a plant are devoted to
the formation of flower-buds to the exclusion of leaf-buds, is the frequency
with which those flowers become affected with floral prolification.
Flowers produced upon indefinite inflorescences are liable to be affected
with either form of prolification more frequently than those borne upon
definite inflorescences. Prolification in both varieties is also more
frequently met with in branched inflorescences than in those in which the
flowers are sessile; but the degree of branching seems less material,
inasmuch as this malformation is more commonly recorded as occurring in
racemes than in the more branched panicles, &c. From the similar arrest of
growth in length, in the case of the flower, to that which occurs in the stem
in the case of definite inflorescence, it might have been expected that
axillary prolification would be more frequent in plants having a cymose
arrangement of their flowers than in those whose inflorescence is indefinite;
such, however, is not the case. The reason for this may be sought for in the
lengthening of the floral axis, so common in prolified flowers—a condition
the reverse of that which happens in the case of definite inflorescence.
usual, is met with in prolified flowers.
In both forms the adventitious growth is much more frequently a flower-
bud or an inflorescence than a leaf-bud or a branch. This may be due to the
position of the flowers on a portion of the stem of the plant especially
devoted to the formation of flower-buds, to the more or less complete
exclusion of leaf-buds, i.e. on the inflorescence. This conjecture is borne
out by the comparative rarity with which prolification has been observed in
flowers that are solitary in the axils of the ordinary leaves of the plant. If the
lists of genera appended hereto be perused, it will be seen that nearly all the
cases occur in genera where the inflorescence is distinctly separated from
the other branches of the stem. In direct proportion, then, to the degree in
which one region of the axis or certain branches of a plant are devoted to
the formation of flower-buds to the exclusion of leaf-buds, is the frequency
with which those flowers become affected with floral prolification.
Flowers produced upon indefinite inflorescences are liable to be affected
with either form of prolification more frequently than those borne upon
definite inflorescences. Prolification in both varieties is also more
frequently met with in branched inflorescences than in those in which the
flowers are sessile; but the degree of branching seems less material,
inasmuch as this malformation is more commonly recorded as occurring in
racemes than in the more branched panicles, &c. From the similar arrest of
growth in length, in the case of the flower, to that which occurs in the stem
in the case of definite inflorescence, it might have been expected that
axillary prolification would be more frequent in plants having a cymose
arrangement of their flowers than in those whose inflorescence is indefinite;
such, however, is not the case. The reason for this may be sought for in the
lengthening of the floral axis, so common in prolified flowers—a condition
the reverse of that which happens in the case of definite inflorescence.
Page 159
Median prolification occurs frequently in double flowers; the axillary
variety, on the other hand, is most common in flowers whose lateral organs
have assumed more or less of the condition of leaves. The other coincident
changes are alluded to elsewhere or do not present useful points of
comparison, and may therefore be passed over.
Prolification of the inflorescence.—This consists in the formation of leaf-
buds or of an undue number of flower-buds on the inflorescence. It must be
distinguished from virescence, or the mere green colour of the floral organs,
and from chloranthy, in which all or the greater portion of the parts of the
flower are replaced by leaves. Prolification is, in fact, a formation of
supernumerary buds, leafy or floral, as the case may be, these buds being
sessile or stalked, the ordinary buds being not necessarily changed.
Prolification of the inflorescence, like the other varieties, admits of
subdivision, not only according to the foliar or floral nature of the bud, but
according to its position, terminal or median and lateral.
Terminal prolification of the inflorescence, whether leafy or floral, is hardly
to be looked upon in the light of a malformation[105] seeing that a similar
condition is so commonly met with normally, as in Epacris, Metrosideros,
Bromelia, Eucomis, &c., wherein the leafy axis projects beyond the
inflorescence proper; or as in Primula imperialis, in which plant, as also in
luxuriant forms of P. sinensis, tier after tier of flowers are placed in
succession above the primary umbel. Nevertheless, when we meet with
such conditions in plants which, under ordinary circumstances, do not
manifest them, we must consider them as coming under the domain of
teratology.
Median foliar prolification of the inflorescence is frequently met with in
Coniferæ, and has of late attracted unwonted attention from the researches
of Caspary, Baillon, and others, on the morphology of these plants. The
variety, on the other hand, is most common in flowers whose lateral organs
have assumed more or less of the condition of leaves. The other coincident
changes are alluded to elsewhere or do not present useful points of
comparison, and may therefore be passed over.
Prolification of the inflorescence.—This consists in the formation of leaf-
buds or of an undue number of flower-buds on the inflorescence. It must be
distinguished from virescence, or the mere green colour of the floral organs,
and from chloranthy, in which all or the greater portion of the parts of the
flower are replaced by leaves. Prolification is, in fact, a formation of
supernumerary buds, leafy or floral, as the case may be, these buds being
sessile or stalked, the ordinary buds being not necessarily changed.
Prolification of the inflorescence, like the other varieties, admits of
subdivision, not only according to the foliar or floral nature of the bud, but
according to its position, terminal or median and lateral.
Terminal prolification of the inflorescence, whether leafy or floral, is hardly
to be looked upon in the light of a malformation[105] seeing that a similar
condition is so commonly met with normally, as in Epacris, Metrosideros,
Bromelia, Eucomis, &c., wherein the leafy axis projects beyond the
inflorescence proper; or as in Primula imperialis, in which plant, as also in
luxuriant forms of P. sinensis, tier after tier of flowers are placed in
succession above the primary umbel. Nevertheless, when we meet with
such conditions in plants which, under ordinary circumstances, do not
manifest them, we must consider them as coming under the domain of
teratology.
Median foliar prolification of the inflorescence is frequently met with in
Coniferæ, and has of late attracted unwonted attention from the researches
of Caspary, Baillon, and others, on the morphology of these plants. The
Page 160
scales and bracts of the cone in these abnormal specimens frequently afford
transitional forms of the greatest value in enabling morphologists to
comprehend the real nature of the floral structure. It would be irrelevant
here to enter into this subject; suffice it merely to say that an examination of
very numerous specimens of this kind, in the common larch and in
Cryptomeria Japonica, has enabled me to verify nearly the whole of
Caspary's observations. A similar prolongation of the axis occurred in some
of the male catkins of Castanea vesca, each of which had a tuft of small
leaves at their extremity. In the common marigold and in Lotus corniculatus
I have also seen instances of this kind. Kirschleger[106] describes a tuft of
leaves as occurring on the apex of the flowering spike after the maturation
of the fruit in Plantago, and a similar growth frequently takes place in the
common wallflower, in Antirrhinum majus, &c. In cases where a renewal of
growth in the axis of inflorescence has taken place after the ripening of the
fruit, the French botanists use the term recrudescence, but the growth in
question by no means always occurs after the ripening of the fruit, but
frequently before. Professor Braun cites the case of a specimen of Plantago
lanceolata, in which the spike was surmounted by a tuft of leaves and roots,
as well as a still more singular instance in Eryngium viviparum, in which
not only did particular branches terminate in rosettes of leaves provided
with roots, but similar growths proceeded from the heads of flowers
themselves. Baron de Mélicoq[107] gives a case in Primula variabilis, in
which at the top of the flower-stalk, in the centre of six flowers, was placed
a complete plant in miniature, having three leaves, from the axil of one of
which proceeded a rudimentary flower. Mr. W. B. Jeffries also forwarded
me a polyanthus (fig. 52) in which the peduncle was surmounted by a small
plant, forming a crown above the ordinary flower-stalk, just as the crown of
the pineapple surmounts that fruit. A similar instance was exhibited at the
Scientific Committee of the Horticultural Society on July 11th, 1868, by
Mr. Wilson Saunders; the species in this case was P. cortusoides. To Mr. R.
transitional forms of the greatest value in enabling morphologists to
comprehend the real nature of the floral structure. It would be irrelevant
here to enter into this subject; suffice it merely to say that an examination of
very numerous specimens of this kind, in the common larch and in
Cryptomeria Japonica, has enabled me to verify nearly the whole of
Caspary's observations. A similar prolongation of the axis occurred in some
of the male catkins of Castanea vesca, each of which had a tuft of small
leaves at their extremity. In the common marigold and in Lotus corniculatus
I have also seen instances of this kind. Kirschleger[106] describes a tuft of
leaves as occurring on the apex of the flowering spike after the maturation
of the fruit in Plantago, and a similar growth frequently takes place in the
common wallflower, in Antirrhinum majus, &c. In cases where a renewal of
growth in the axis of inflorescence has taken place after the ripening of the
fruit, the French botanists use the term recrudescence, but the growth in
question by no means always occurs after the ripening of the fruit, but
frequently before. Professor Braun cites the case of a specimen of Plantago
lanceolata, in which the spike was surmounted by a tuft of leaves and roots,
as well as a still more singular instance in Eryngium viviparum, in which
not only did particular branches terminate in rosettes of leaves provided
with roots, but similar growths proceeded from the heads of flowers
themselves. Baron de Mélicoq[107] gives a case in Primula variabilis, in
which at the top of the flower-stalk, in the centre of six flowers, was placed
a complete plant in miniature, having three leaves, from the axil of one of
which proceeded a rudimentary flower. Mr. W. B. Jeffries also forwarded
me a polyanthus (fig. 52) in which the peduncle was surmounted by a small
plant, forming a crown above the ordinary flower-stalk, just as the crown of
the pineapple surmounts that fruit. A similar instance was exhibited at the
Scientific Committee of the Horticultural Society on July 11th, 1868, by
Mr. Wilson Saunders; the species in this case was P. cortusoides. To Mr. R.
Page 161
Dean I am indebted for a similar proliferous cyclamen, which seems similar
to one mentioned by Schlechtendal.[108] This author alludes to an analogous
circumstance in the inflorescence of Cytisus nigricans, where, however, the
change was not so great as in the preceding cases. The instances just cited
all occur in plants having an indefinite form of inflorescence; but the
production of a tuft of leaves or of a leafy shoot above or beyond the
inflorescence is not confined to plants with this habit of growth, for Jacquin
figures and describes an instance of this nature in the cymose flower-stems
of a Sempervivum. "Hi racemi," says he, "ultra flores producuntur in
ramos, foliosos duo bifidos qui tandem trium unciarum longitudinem adepti
fuerunt."[109]
Fig. 52.—Inflorescence of Polyanthus, bearing a tuft of leaves at the top of the scape intermixed with
the flowers.
Median floral prolification of the inflorescence, wherein a new
inflorescence projects beyond the primary one, is not uncommon in plants
to one mentioned by Schlechtendal.[108] This author alludes to an analogous
circumstance in the inflorescence of Cytisus nigricans, where, however, the
change was not so great as in the preceding cases. The instances just cited
all occur in plants having an indefinite form of inflorescence; but the
production of a tuft of leaves or of a leafy shoot above or beyond the
inflorescence is not confined to plants with this habit of growth, for Jacquin
figures and describes an instance of this nature in the cymose flower-stems
of a Sempervivum. "Hi racemi," says he, "ultra flores producuntur in
ramos, foliosos duo bifidos qui tandem trium unciarum longitudinem adepti
fuerunt."[109]
Fig. 52.—Inflorescence of Polyanthus, bearing a tuft of leaves at the top of the scape intermixed with
the flowers.
Median floral prolification of the inflorescence, wherein a new
inflorescence projects beyond the primary one, is not uncommon in plants
Page 162
having their flowers arranged in close heads or umbels, as in the common
wild celery and other Umbelliferæ.[110] I have also met with it in Trifolium
repens, in the umbellate variety of the common primrose, and in the scarlet
geranium. Engelmann cites it in Triticum repens, Roëper in Euphorbia
palustris.[111]
Lateral foliar prolification of the inflorescence is of more common
occurrence than the preceding. I have met with it, amongst other plants,
frequently in Brassica oleracea, Pelargonium zonale, Scabiosa, Bellis, and
many other composites, also in Leguminosæ, e.g. Lupinus, Trifolium,
Coronilla, &c. Prof. Oliver forwarded me a specimen of Euphorbia
geniculata in which, in addition to other changes, there was a series of
stalked buds bearing tufts of green scales, but without any trace of stamens
or pistil; these adventitious buds occurred within the ordinary involucre of
the plant, between it and the stamens. The pistil was unaffected in some
cases, while in some others it was entirely wanting, the gynophore being
surmounted by a cup-like involucre, divided into three acutely pointed
lobes, each with a midrib; these encircled a series of stalked involucels, as
before, and among which were scattered a few stamens, some perfect,
others partially frondescent.
In a specimen of Scrophularia nodosa examined by me one of the lateral
buds on each of the cymes was represented, not by a flower, but by a tuft of
leaves, the other buds being unchanged. As the inflorescence was much
contracted in size, the appearance of the whole plant was greatly changed.
Many of the instances of so-called viviparous plants, e.g., Polygonum
viviparum, may be cited under this head.[112] Many species of Allium,
Lilium, Saxifraga, Begonia, Achimenes, normally produce leaf-buds or
bulbs in the inflorescence; so, too, leafy shoots are sometimes found in
Alisma natans, Juncus uliginosus, Chlorophytum Sternbergianum, &c. As
wild celery and other Umbelliferæ.[110] I have also met with it in Trifolium
repens, in the umbellate variety of the common primrose, and in the scarlet
geranium. Engelmann cites it in Triticum repens, Roëper in Euphorbia
palustris.[111]
Lateral foliar prolification of the inflorescence is of more common
occurrence than the preceding. I have met with it, amongst other plants,
frequently in Brassica oleracea, Pelargonium zonale, Scabiosa, Bellis, and
many other composites, also in Leguminosæ, e.g. Lupinus, Trifolium,
Coronilla, &c. Prof. Oliver forwarded me a specimen of Euphorbia
geniculata in which, in addition to other changes, there was a series of
stalked buds bearing tufts of green scales, but without any trace of stamens
or pistil; these adventitious buds occurred within the ordinary involucre of
the plant, between it and the stamens. The pistil was unaffected in some
cases, while in some others it was entirely wanting, the gynophore being
surmounted by a cup-like involucre, divided into three acutely pointed
lobes, each with a midrib; these encircled a series of stalked involucels, as
before, and among which were scattered a few stamens, some perfect,
others partially frondescent.
In a specimen of Scrophularia nodosa examined by me one of the lateral
buds on each of the cymes was represented, not by a flower, but by a tuft of
leaves, the other buds being unchanged. As the inflorescence was much
contracted in size, the appearance of the whole plant was greatly changed.
Many of the instances of so-called viviparous plants, e.g., Polygonum
viviparum, may be cited under this head.[112] Many species of Allium,
Lilium, Saxifraga, Begonia, Achimenes, normally produce leaf-buds or
bulbs in the inflorescence; so, too, leafy shoots are sometimes found in
Alisma natans, Juncus uliginosus, Chlorophytum Sternbergianum, &c. As
Page 163
an accidental occurrence, a similar thing has been noticed in Lychnis
coronaria, Phaius grandifolius, Oncidium cebolleta, Epidendrum
elongatum,[113] &c. &c.
Here, too, may be mentioned those cases wherein a leaf-bud is found upon
the surface of the so-called inferior ovary; generally a leaf only is found,
but a leaf-bud may also originate in this situation, and in either case the
inference is that the ovary is, in part at least, made of the dilated and
hollowed axis. Leaves may occasionally be found in this way on the so-
called calyx-tube or on the inferior ovaries of roses, pears, apples, Pereskia,
Cratægus tanacetifolia, &c.
The fruits of Opuntia Salmania and of O. fragilis ('Bull. Soc. Bot. France,'
vol. i, p. 306; vol. v, p. 115) have been observed to form small fruit-like
branches around their summits. This circumstance is more fully treated of in
the succeeding chapter relating to Heterotaxy.
Lateral floral prolification of the inflorescence.—This, which is termed
by Engelmann Ecblastesis foliorum sub floralium,[114] is much the most
common of all these deviations, and it is met with in every degree, from the
presence of a single supernumerary flower in the axil of a bract to the
existence of a small cluster or panicle of such flowers.
coronaria, Phaius grandifolius, Oncidium cebolleta, Epidendrum
elongatum,[113] &c. &c.
Here, too, may be mentioned those cases wherein a leaf-bud is found upon
the surface of the so-called inferior ovary; generally a leaf only is found,
but a leaf-bud may also originate in this situation, and in either case the
inference is that the ovary is, in part at least, made of the dilated and
hollowed axis. Leaves may occasionally be found in this way on the so-
called calyx-tube or on the inferior ovaries of roses, pears, apples, Pereskia,
Cratægus tanacetifolia, &c.
The fruits of Opuntia Salmania and of O. fragilis ('Bull. Soc. Bot. France,'
vol. i, p. 306; vol. v, p. 115) have been observed to form small fruit-like
branches around their summits. This circumstance is more fully treated of in
the succeeding chapter relating to Heterotaxy.
Lateral floral prolification of the inflorescence.—This, which is termed
by Engelmann Ecblastesis foliorum sub floralium,[114] is much the most
common of all these deviations, and it is met with in every degree, from the
presence of a single supernumerary flower in the axil of a bract to the
existence of a small cluster or panicle of such flowers.
Page 164
Fig. 53.—Lateral prolification in inflorescence of Pelargonium.
It is common in the Anemone coronaria and hortensis, also in the common
scarlet Pelargonium (fig. 53). It has been frequently recorded in Poterium
sanguisorba, and in Sanguisorba officinalis, and is especially common in
Umbelliferæ, Dipsacaceæ, and Compositæ; a familiar illustration in the
latter order is afforded by the hen-and-chicken daisy. In some species of
Compositæ, indeed, it is a normal and constant occurrence, while in other
cases, such as Filago germanica, usually described as proliferous, there is
not, strictly speaking, any prolification, for the branching of the stalk takes
place below the inflorescence, and the branches originate from the axils of
ordinary leaves, not from the floral leaves or bracts. Convolvulus Sepium is
very commonly subject to the production of flower-buds from the axils of
the floral leaves. The several species of Plantain (Plantago) seem very
liable to this and similar changes. Schlechtendal[115] gives a summary of the
various kinds of malformation affecting the inflorescence in Plantago, and
It is common in the Anemone coronaria and hortensis, also in the common
scarlet Pelargonium (fig. 53). It has been frequently recorded in Poterium
sanguisorba, and in Sanguisorba officinalis, and is especially common in
Umbelliferæ, Dipsacaceæ, and Compositæ; a familiar illustration in the
latter order is afforded by the hen-and-chicken daisy. In some species of
Compositæ, indeed, it is a normal and constant occurrence, while in other
cases, such as Filago germanica, usually described as proliferous, there is
not, strictly speaking, any prolification, for the branching of the stalk takes
place below the inflorescence, and the branches originate from the axils of
ordinary leaves, not from the floral leaves or bracts. Convolvulus Sepium is
very commonly subject to the production of flower-buds from the axils of
the floral leaves. The several species of Plantain (Plantago) seem very
liable to this and similar changes. Schlechtendal[115] gives a summary of the
various kinds of malformation affecting the inflorescence in Plantago, and
Page 165
divides them into five groups, as follows:—1st, bracteate, wherein the
inferior bracts are quite leaf-like, as is frequently seen in Plantago major.
2nd, roseate; bracts leafy in tufts or rosettes, without flowers, as in the so-
called rose plantain, common in old-fashioned gardens in this country. 3rd,
polystachyate; spike-branched, bearing other spikes in the axils of the
bracts, as in P. lanceolata, P. maritima, &c. 4th, proliferous, where the
flower-stalk bears a rosette, a spike, or a head with other rosettes. 5th,
paniculate, in which the inflorescence has become a much-branched
pyramidal panicle, covered with little bracts, and with very rudimentary
flowers.[116] The first two groups belong rather to frondescence of the
bracts; but with regard to the whole of them it will easily be surmised that
intermediate forms occur, linking one group to the other, and defying exact
allocation in either. Thus, in the borders of richly cultivated fields in the
neighbourhood of London I have frequently gathered specimens of
Plantago major with a branched spike provided with large leafy bracts, the
branches of the spike being but little less in diameter than the ordinary
single spike. These specimens would therefore seem to be intermediate
between Schlechtendal's bracteate and polystachyate divisions. Wigand[117]
also describes an anomalous specimen of Plantago major similar to those
just mentioned, but having small lateral spikes in place of large ones. The
instance quoted from Professor Braun would fall under the roseate section,
as would also that of Kirschleger, though we are expressly told that the tuft
of leaves in this last case was not developed until after the ripening of the
seed-vessel. One of the characters of the roseate group, according to
Schlechtendal, is the absence of flowers, but most persons who have had
the opportunity of watching the growth of the rose plantain must have
observed the occasional production of flowers, sometimes stalked, in the
axils of the leafy bracts, and at the same time have noticed that the
internodes become elongated, so that an approach is made to the ordinary
spike-like form of the inflorescence. The proliferous group would include
inferior bracts are quite leaf-like, as is frequently seen in Plantago major.
2nd, roseate; bracts leafy in tufts or rosettes, without flowers, as in the so-
called rose plantain, common in old-fashioned gardens in this country. 3rd,
polystachyate; spike-branched, bearing other spikes in the axils of the
bracts, as in P. lanceolata, P. maritima, &c. 4th, proliferous, where the
flower-stalk bears a rosette, a spike, or a head with other rosettes. 5th,
paniculate, in which the inflorescence has become a much-branched
pyramidal panicle, covered with little bracts, and with very rudimentary
flowers.[116] The first two groups belong rather to frondescence of the
bracts; but with regard to the whole of them it will easily be surmised that
intermediate forms occur, linking one group to the other, and defying exact
allocation in either. Thus, in the borders of richly cultivated fields in the
neighbourhood of London I have frequently gathered specimens of
Plantago major with a branched spike provided with large leafy bracts, the
branches of the spike being but little less in diameter than the ordinary
single spike. These specimens would therefore seem to be intermediate
between Schlechtendal's bracteate and polystachyate divisions. Wigand[117]
also describes an anomalous specimen of Plantago major similar to those
just mentioned, but having small lateral spikes in place of large ones. The
instance quoted from Professor Braun would fall under the roseate section,
as would also that of Kirschleger, though we are expressly told that the tuft
of leaves in this last case was not developed until after the ripening of the
seed-vessel. One of the characters of the roseate group, according to
Schlechtendal, is the absence of flowers, but most persons who have had
the opportunity of watching the growth of the rose plantain must have
observed the occasional production of flowers, sometimes stalked, in the
axils of the leafy bracts, and at the same time have noticed that the
internodes become elongated, so that an approach is made to the ordinary
spike-like form of the inflorescence. The proliferous group would include
Page 166
such specimens as that of P. lanceolata mentioned by Dr. Johnston,[118]
wherein were several spikes, some sessile, others stalked and pendent, the
whole intermixed with leaves and disposed in a rose-like manner. I have
myself gathered specimens of this nature, occurring in the same plant, at
Shanklin, Isle of Wight (fig. 56).
Fig. 54.—Plantago major, with panicled inflorescence.
wherein were several spikes, some sessile, others stalked and pendent, the
whole intermixed with leaves and disposed in a rose-like manner. I have
myself gathered specimens of this nature, occurring in the same plant, at
Shanklin, Isle of Wight (fig. 56).
Fig. 54.—Plantago major, with panicled inflorescence.
Page 167
Fig. 55.—Inflorescence of Plantago major, with bracts partly replaced by leaves and spike branched.
Page 168
Fig. 56.—Inflorescence of Plantago lanceolata, bearing a tuft of leaves and flowers at the end of the
flower-scape.
It is rather singular that each species of Plantago seems to have its own
perverse mode of growth; for instance, the bracteate, polystachyate and
paniculate forms are almost exclusively confined to P. major, the roseate
form to P. media, the proliferous form to P. lanceolata.
The instances wherein flower-buds originate from the surface of an inferior
ovary, as in those cases where the top of the stem is dilated so as to form
part of the fruit, would be properly classed under the head of prolification
of the inflorescence. As, however, there is still some difference of opinion
as to the correct morphological interpretation to be put on some of these
cases, it has been thought better to include them under the head of
heterotaxy than of prolification.
Fig. 57.—Branched inflorescence of Reseda luteola.
flower-scape.
It is rather singular that each species of Plantago seems to have its own
perverse mode of growth; for instance, the bracteate, polystachyate and
paniculate forms are almost exclusively confined to P. major, the roseate
form to P. media, the proliferous form to P. lanceolata.
The instances wherein flower-buds originate from the surface of an inferior
ovary, as in those cases where the top of the stem is dilated so as to form
part of the fruit, would be properly classed under the head of prolification
of the inflorescence. As, however, there is still some difference of opinion
as to the correct morphological interpretation to be put on some of these
cases, it has been thought better to include them under the head of
heterotaxy than of prolification.
Fig. 57.—Branched inflorescence of Reseda luteola.
Page 169
Some of the cases of prolification of the inflorescence resulting in a
branching of an ordinarily simple inflorescence, as in Reseda luteola (fig.
57), might equally well be placed with fission or multiplication of the axile
organs. Branched spikes of this character are not so common among
Orchids as might be expected. Professor Reichenbach enumerates a few
instances in the Report of the International Botanical Congress of London,
1866, p. 121, and the same author gives an illustration in his
'Orchidographia Europœa,' tab. 150.
In Grasses, as indeed in other plants with a spicate inflorescence, this
change occurs not unfrequently. The common Ray Grass (Lolium) is
especially subject to the change in question, and among cultivated cereals,
maize and wheat occasionally show this tendency to subdivision. One
variety of the latter grain is cultivated in hot countries under the name of
Egyptian wheat—Triticum vulgare, var. compositum.
Prolification of the inflorescence has been most frequently observed in the
following genera:
Leafy. Floral.
Ranunculaceæ Ranunculus. Ranunculus!
Anemone. Anemone.
Cruciferæ. *Brassica!
Caryophyllaceæ. Lychnis!
Dianthus!
Geraniaceæ. *Pelargonium! *Pelargonium!
Leguminosæ. *Trifolium! Trifolium!
Lotus! Lotus!
Coronilla!
Cytisus. Cytisus.
Rosaceæ. Poterium.
*Pyrus! *Pyrus!
*Cratægus! Cratægus!
*Rosa. Rosa!
Sanguisorba.
Philadelphaceæ. Philadelphus.
Crassulaceæ. Sempervivum.
branching of an ordinarily simple inflorescence, as in Reseda luteola (fig.
57), might equally well be placed with fission or multiplication of the axile
organs. Branched spikes of this character are not so common among
Orchids as might be expected. Professor Reichenbach enumerates a few
instances in the Report of the International Botanical Congress of London,
1866, p. 121, and the same author gives an illustration in his
'Orchidographia Europœa,' tab. 150.
In Grasses, as indeed in other plants with a spicate inflorescence, this
change occurs not unfrequently. The common Ray Grass (Lolium) is
especially subject to the change in question, and among cultivated cereals,
maize and wheat occasionally show this tendency to subdivision. One
variety of the latter grain is cultivated in hot countries under the name of
Egyptian wheat—Triticum vulgare, var. compositum.
Prolification of the inflorescence has been most frequently observed in the
following genera:
Leafy. Floral.
Ranunculaceæ Ranunculus. Ranunculus!
Anemone. Anemone.
Cruciferæ. *Brassica!
Caryophyllaceæ. Lychnis!
Dianthus!
Geraniaceæ. *Pelargonium! *Pelargonium!
Leguminosæ. *Trifolium! Trifolium!
Lotus! Lotus!
Coronilla!
Cytisus. Cytisus.
Rosaceæ. Poterium.
*Pyrus! *Pyrus!
*Cratægus! Cratægus!
*Rosa. Rosa!
Sanguisorba.
Philadelphaceæ. Philadelphus.
Crassulaceæ. Sempervivum.
Page 170
Echeveria.
Crassula.
Ficoideæ. ?Tetragonia.
Cactaceæ. Opuntia. Opuntia.
Pereskia.
Saxifragaceæ. Saxifraga!
Umbelliferæ. Seseli.
*Apium!
Cnidium.
Chærophyllum.
Eryngium. Eryngium.
Silaus.
Heracleum! Heracleum!
Hydrocotyle. Hydrocotyle.
Daucus.
Carum.
Selinum.
Angelica!
Conium.
Astrantia.
Œnanthe. Œnanthe.
Begoniaceæ. Begonia!
Valerianaceæ. Valeriana.
Dipsacaceæ. *Scabiosa! *Scabiosa!
Knautia! Knautia!
Compositæ. *Bellis!
Centaurea.
Calendula. Calendula.
Anthemis.
Coreopsis.
Apargia.
Lampsana.
Carlina.
Arnoseris.
Tragopogon! Tragopogon!
Rudbeckia!
Senecio!
Carlina.
Bidens! Pyrethrum.
Filago.
Hedypnois.
Cirsium.
Lactuca.
Campanulaceæ. Prismatocarpus.
Lobeliaceæ. Jasione.
Ericaceæ. Azalea!
Crassula.
Ficoideæ. ?Tetragonia.
Cactaceæ. Opuntia. Opuntia.
Pereskia.
Saxifragaceæ. Saxifraga!
Umbelliferæ. Seseli.
*Apium!
Cnidium.
Chærophyllum.
Eryngium. Eryngium.
Silaus.
Heracleum! Heracleum!
Hydrocotyle. Hydrocotyle.
Daucus.
Carum.
Selinum.
Angelica!
Conium.
Astrantia.
Œnanthe. Œnanthe.
Begoniaceæ. Begonia!
Valerianaceæ. Valeriana.
Dipsacaceæ. *Scabiosa! *Scabiosa!
Knautia! Knautia!
Compositæ. *Bellis!
Centaurea.
Calendula. Calendula.
Anthemis.
Coreopsis.
Apargia.
Lampsana.
Carlina.
Arnoseris.
Tragopogon! Tragopogon!
Rudbeckia!
Senecio!
Carlina.
Bidens! Pyrethrum.
Filago.
Hedypnois.
Cirsium.
Lactuca.
Campanulaceæ. Prismatocarpus.
Lobeliaceæ. Jasione.
Ericaceæ. Azalea!
Page 171
Convolvulaceæ. Convolvulus! Convolvulus!
Calystegia!
Scrophulariaceæ. Scrophularia!
Antirrhinum!
Gesneraceæ. Achimenes!
Primulaceæ. Primula! Primula!
Cyclamen! Cyclamen!
Plumbaginaceæ. Armeria.
Plantaginaceæ. *Plantago! *Plantago!
Polygonaceæ. Polygonum!
Euphorbiaceæ. Euphorbia!
Urticaceæ. Ficus.
Amentaceæ. Corylus!
Castanea! Castanea.
Coniferæ. *Larix!
*Cryptomeria!
Taxodium! Pinus.
Orchidaceæ. Phaius! Ophrys!
Epidendrum!
Oncidium!
Liliaceæ. *Allium!
*Ornithogalum!
*Lilium!
Amaryllidaceæ. Fourcroya
Alismaceæ. Alisma!
Palmaceæ. Cocos.
Juncaceæ. *Juncus!
Restiaceæ. Restio! Restio!
Elegia! Elegia!
Willdenovia! Willdenovia!
Cyperaceæ. Carex.
Graminaceæ. Dactylis.
*Lolium!
Festuca.
*Zea!
*Triticum!
*Hordeum!
Secale.
Phleum.
In addition to the papers already cited the following works may be
consulted with reference to prolification of the inflorescence:
Moquin-Tandon. 'El. Ter. Veg.,' p. 376. Engelmann, 'De Antholysi,' §§ 85–87. Fleischer,
'Missbild. Versch. Cultur. Pflanz.' For figures of Hen and Chicken Daisy (Bellis prolifera).
Calystegia!
Scrophulariaceæ. Scrophularia!
Antirrhinum!
Gesneraceæ. Achimenes!
Primulaceæ. Primula! Primula!
Cyclamen! Cyclamen!
Plumbaginaceæ. Armeria.
Plantaginaceæ. *Plantago! *Plantago!
Polygonaceæ. Polygonum!
Euphorbiaceæ. Euphorbia!
Urticaceæ. Ficus.
Amentaceæ. Corylus!
Castanea! Castanea.
Coniferæ. *Larix!
*Cryptomeria!
Taxodium! Pinus.
Orchidaceæ. Phaius! Ophrys!
Epidendrum!
Oncidium!
Liliaceæ. *Allium!
*Ornithogalum!
*Lilium!
Amaryllidaceæ. Fourcroya
Alismaceæ. Alisma!
Palmaceæ. Cocos.
Juncaceæ. *Juncus!
Restiaceæ. Restio! Restio!
Elegia! Elegia!
Willdenovia! Willdenovia!
Cyperaceæ. Carex.
Graminaceæ. Dactylis.
*Lolium!
Festuca.
*Zea!
*Triticum!
*Hordeum!
Secale.
Phleum.
In addition to the papers already cited the following works may be
consulted with reference to prolification of the inflorescence:
Moquin-Tandon. 'El. Ter. Veg.,' p. 376. Engelmann, 'De Antholysi,' §§ 85–87. Fleischer,
'Missbild. Versch. Cultur. Pflanz.' For figures of Hen and Chicken Daisy (Bellis prolifera).
Page 172
see Lobel, 'Ic.,' 477. Sweert, 'Florileg.,' pl. 98, f. 5. 'Hort. Eystett. Plant. Vern.,' fol. iv, f. i.
&c. For similar malformations in marigold (Calendula), see Lobel, 'Ic.,' 553. 'Act. Acad.
Nat. Cur.,' vol. x, p. 208. Jaeger, 'Missbilld.,' 192–195. 'Hort. Eystett.,' pl. æstiv. fol. iii, f. i.
Klinsmann, 'Linnæa,' t. x, p. 607.
For monstrous plantains, in addition to previous citations, see Camerarius, 'Epist.,' p. 261,
P. rosea. Matthioli, 'Krauterb,' 245. Lobel, 'Stirp. Advers. Nov.,' p. 128, P. major
paniculata. J. Bauhin, 'Hist. Plant.,' i, p. 503 b. Ibid., p. 503, a, c, P. major rosea, bracteata
paniculata, prolifera, &c. 'Hort. Eystett.,' pl. æstiv., t. vii, f. 2, P. rosea et P. bracteata.
Lobel, 'Stirp. Hist.,' p. 162. Dodonæus, 'Pempt.,' 1–4, cap. xxiii, P. major spica multiplex,
i.e. paniculata. Gerard, 'Herbal.' Clusius, 'Plant. Rar. Hist.,' lib. v, p. 109–10, Plantago
augustifolia Gareti prolifera. Marchand, 'Adansonia,' iv, p. 156.
Coniferæ.—Richard, 'Mem. Conif.,' tab. xiii, f. 9. A. Braun, 'Das Individ.,' 1853, p. 65. De
Cand., 'Organogr.,' tab. xxxvi. Wigand, 'Bot. Untersuch.,' 154. Schlechtendal, 'Bot. Zeit.,'
1859, p. 239. Caspary, 'De Abiet. flor. fem. struct. morphol.' Parlatore, 'Ann. Sc. Nat.,'
1862, vol. xvi, p. 215. Cramer, 'Bildungsabweich.,' p. 4, &c., &c.
Gramineæ.—Bauhin, 'Pinax.,' 21. Morison, 'Hist. Plant.,' t. i. Winckler, 'Ephem. Nat. Cur.,'
dec. i, ann. 7, 8, p. 151. Irmisch, 'Flora,' 1858, p. 40, &c.
See also under Chloranthy, Viviparous plants, &c.
Prolification of the flower.—In the preceding sections the formation of
adventitious buds of a leafy or floral nature on the inflorescence has been
considered. A similar production of buds may take place in the flower itself,
either from its centre or from the axil of some of its constituent parts.
Prolification of the flower is therefore median or axillary, and the
adventitious bud itself may be of a leafy or a floral nature.
Median leafy prolification.—In this malformation the centre of the flower
is occupied by a bud or a branch; the growing point or termination of the
axis which ordinarily ceases to grow after the formation of the carpels,
takes on new growth. This is well shown in the accompanying illustration
(fig. 58), representing the thalamus of a strawberry prolonged beyond the
fruits into a small leaf-bearing branch.
&c. For similar malformations in marigold (Calendula), see Lobel, 'Ic.,' 553. 'Act. Acad.
Nat. Cur.,' vol. x, p. 208. Jaeger, 'Missbilld.,' 192–195. 'Hort. Eystett.,' pl. æstiv. fol. iii, f. i.
Klinsmann, 'Linnæa,' t. x, p. 607.
For monstrous plantains, in addition to previous citations, see Camerarius, 'Epist.,' p. 261,
P. rosea. Matthioli, 'Krauterb,' 245. Lobel, 'Stirp. Advers. Nov.,' p. 128, P. major
paniculata. J. Bauhin, 'Hist. Plant.,' i, p. 503 b. Ibid., p. 503, a, c, P. major rosea, bracteata
paniculata, prolifera, &c. 'Hort. Eystett.,' pl. æstiv., t. vii, f. 2, P. rosea et P. bracteata.
Lobel, 'Stirp. Hist.,' p. 162. Dodonæus, 'Pempt.,' 1–4, cap. xxiii, P. major spica multiplex,
i.e. paniculata. Gerard, 'Herbal.' Clusius, 'Plant. Rar. Hist.,' lib. v, p. 109–10, Plantago
augustifolia Gareti prolifera. Marchand, 'Adansonia,' iv, p. 156.
Coniferæ.—Richard, 'Mem. Conif.,' tab. xiii, f. 9. A. Braun, 'Das Individ.,' 1853, p. 65. De
Cand., 'Organogr.,' tab. xxxvi. Wigand, 'Bot. Untersuch.,' 154. Schlechtendal, 'Bot. Zeit.,'
1859, p. 239. Caspary, 'De Abiet. flor. fem. struct. morphol.' Parlatore, 'Ann. Sc. Nat.,'
1862, vol. xvi, p. 215. Cramer, 'Bildungsabweich.,' p. 4, &c., &c.
Gramineæ.—Bauhin, 'Pinax.,' 21. Morison, 'Hist. Plant.,' t. i. Winckler, 'Ephem. Nat. Cur.,'
dec. i, ann. 7, 8, p. 151. Irmisch, 'Flora,' 1858, p. 40, &c.
See also under Chloranthy, Viviparous plants, &c.
Prolification of the flower.—In the preceding sections the formation of
adventitious buds of a leafy or floral nature on the inflorescence has been
considered. A similar production of buds may take place in the flower itself,
either from its centre or from the axil of some of its constituent parts.
Prolification of the flower is therefore median or axillary, and the
adventitious bud itself may be of a leafy or a floral nature.
Median leafy prolification.—In this malformation the centre of the flower
is occupied by a bud or a branch; the growing point or termination of the
axis which ordinarily ceases to grow after the formation of the carpels,
takes on new growth. This is well shown in the accompanying illustration
(fig. 58), representing the thalamus of a strawberry prolonged beyond the
fruits into a small leaf-bearing branch.
Page 173
Fig. 58.—Receptacle of strawberry prolonged into a leafy branch. From the 'American Agriculturist.'
Page 174
Fig. 59.—Flower of Verbascum with five disunited sepals, five similar green petals, and a prolonged
branch in the centre of the flower.
In other cases the carpels are entirely absent and their place is supplied by a
leafy shoot as in a species of Verbascum, which came under my own
observation. In this case the petals were virescent, and the stamens and
pistils were entirely absent, hence in truth, the so-called flower more nearly
resembled a branch. In a flower of a May Duke cherry, for which I am
indebted to Mr. Salter, there was a gradual change from the floral to the
foliar condition; thus there were five distinct lanceolate sepals, the
arrangement of whose veins betokened that they were leaf-sheaths rather
than perfect leaves, ten petals partly foliaceous and sheath-like as to their
venation, one of them funnel-shaped, but whether from dilatation or
cohesion of the margins could not be determined. The stamens were eight
or ten in number, their connectives prolonged into foliaceous or petaloid
appendages, so that the filament represented the stalk of the leaf. The pistil
was entirely absent and its place was supplied by a branch with numerous
perfectly formed stipulate leaves.
Some flowers of Anagallis arvensis described by Dr. Marchand[119] are so
interesting and show so well the gradual stages by which this malformation
is arrived at, that it is desirable to cite the summary of Dr. Marchand's
researches as given in the 'Gardeners' Chronicle' by Mr. Berkeley, taking
that instance first in which the parts of the flower departed least from the
normal condition, and then the others in their proper order. In all the parts
there was a greater or less tendency to assume a green tint; in some they
were entirely green, in others the brighter colours were confined to the more
recently developed parts.
"1. In the first case then, the sepals and petals were in their normal position,
though rather more dilated than usual; the anthers were fertile, the principal
branch in the centre of the flower.
In other cases the carpels are entirely absent and their place is supplied by a
leafy shoot as in a species of Verbascum, which came under my own
observation. In this case the petals were virescent, and the stamens and
pistils were entirely absent, hence in truth, the so-called flower more nearly
resembled a branch. In a flower of a May Duke cherry, for which I am
indebted to Mr. Salter, there was a gradual change from the floral to the
foliar condition; thus there were five distinct lanceolate sepals, the
arrangement of whose veins betokened that they were leaf-sheaths rather
than perfect leaves, ten petals partly foliaceous and sheath-like as to their
venation, one of them funnel-shaped, but whether from dilatation or
cohesion of the margins could not be determined. The stamens were eight
or ten in number, their connectives prolonged into foliaceous or petaloid
appendages, so that the filament represented the stalk of the leaf. The pistil
was entirely absent and its place was supplied by a branch with numerous
perfectly formed stipulate leaves.
Some flowers of Anagallis arvensis described by Dr. Marchand[119] are so
interesting and show so well the gradual stages by which this malformation
is arrived at, that it is desirable to cite the summary of Dr. Marchand's
researches as given in the 'Gardeners' Chronicle' by Mr. Berkeley, taking
that instance first in which the parts of the flower departed least from the
normal condition, and then the others in their proper order. In all the parts
there was a greater or less tendency to assume a green tint; in some they
were entirely green, in others the brighter colours were confined to the more
recently developed parts.
"1. In the first case then, the sepals and petals were in their normal position,
though rather more dilated than usual; the anthers were fertile, the principal
Page 175
change existing in the ovary, the upper part of which was wanting, so that
the ovules were exposed seated on the central placenta.
2. In the next step the calyx, more developed than usual, was separated
from the corolla by a long peduncle, and the ovary, which was ovate,
contained instead of a placenta a sort of plumule or young shoot.
3. In this case the corolla and calyx were distant from each other; there was
no trace of stamens, but the axis was continued from the centre of the
corolla, and ended in a leaf-bud.
4. The calyx and corolla nearly as before, but instead of stamens a whorl of
little leaves was developed, in the centre of which the axis was continued,
bearing at its tip two whorls of leaflets, alternately three and three.
5. In this case two out of the five stamens were normal, the other three
changed into leaves, showing clearly the origin of the leaflets, in the last
case, which took the place of the stamens.
6. The ovary varied in different flowers. In some the placenta was crowned
with ovules; in others the ovules were replaced by a single whorl of leaflets;
in others there was every shade of change from ordinary ovules to perfect
leaflets; while in others, again, every ovule was converted into a leaf with a
long petiole.
7. In these flowers shoots were developed in the axils of the sepals, or on
the face of the petals between the point of their insertion and that of the
stamens, and, what is most curious, in the interior of the ovaries round the
foot of the placenta.
8. Here, again, a very singular condition presented itself: the calyx and
corolla separated from each other, the stamens partly developed, the axis
the ovules were exposed seated on the central placenta.
2. In the next step the calyx, more developed than usual, was separated
from the corolla by a long peduncle, and the ovary, which was ovate,
contained instead of a placenta a sort of plumule or young shoot.
3. In this case the corolla and calyx were distant from each other; there was
no trace of stamens, but the axis was continued from the centre of the
corolla, and ended in a leaf-bud.
4. The calyx and corolla nearly as before, but instead of stamens a whorl of
little leaves was developed, in the centre of which the axis was continued,
bearing at its tip two whorls of leaflets, alternately three and three.
5. In this case two out of the five stamens were normal, the other three
changed into leaves, showing clearly the origin of the leaflets, in the last
case, which took the place of the stamens.
6. The ovary varied in different flowers. In some the placenta was crowned
with ovules; in others the ovules were replaced by a single whorl of leaflets;
in others there was every shade of change from ordinary ovules to perfect
leaflets; while in others, again, every ovule was converted into a leaf with a
long petiole.
7. In these flowers shoots were developed in the axils of the sepals, or on
the face of the petals between the point of their insertion and that of the
stamens, and, what is most curious, in the interior of the ovaries round the
foot of the placenta.
8. Here, again, a very singular condition presented itself: the calyx and
corolla separated from each other, the stamens partly developed, the axis
Page 176
continued beyond the corolla, branched and bearing normal leaves so as
exactly to resemble an ordinary stem, while in consequence of the calyx and
corolla being bent down to the ground, adventitious roots were developed
from the axis on the under side above each of them. In another case, where
the calyx and corolla were approximated, the ovary was open above, and
sent out six shoots from within, perfectly developed, clearly representing
the central placenta and five axile buds, and each giving out a number of
adventitious roots at its base."
In other genera of the same order (Primulaceæ) an extension of the placenta
into a leafy branch has been observed, as in Lysimachia, where in one case
the prolonged placenta was removed and struck as a cutting.[120]
In Ericaceæ too, the axile placenta has been seen ovuliferous at the base
and prolonged above into a leafy branch.[121]
Median floral prolification.—This is of more frequent occurrence than the
preceding. The prolonged axis is more frequently terminated by a flower-
bud than by a leaf-bud, though it must be remarked, that the lengthened and
protruded stem frequently bears leaves upon its sides, even if it terminate in
a flower, and thus the new growth partakes of a mixed leafy and floral
nature. Instances of this kind have long been familiar to observers, and have
always excited attention from the singularity of their appearance. In one of
the old stained-glass windows, apparently of Dutch manufacture, in the
Bodleian Picture Gallery at Oxford, is a representation of a Ranunculus
affected with median floral prolification.[122] In pinks the affection is not
unfrequently met with. Fig. 60 shows an instance of the kind copied from
Schotterbec.
A singular instance of prolification in the central flower of one of the
verticillasters of Phlomis fruticosa fell under my own notice; it was a case
exactly to resemble an ordinary stem, while in consequence of the calyx and
corolla being bent down to the ground, adventitious roots were developed
from the axis on the under side above each of them. In another case, where
the calyx and corolla were approximated, the ovary was open above, and
sent out six shoots from within, perfectly developed, clearly representing
the central placenta and five axile buds, and each giving out a number of
adventitious roots at its base."
In other genera of the same order (Primulaceæ) an extension of the placenta
into a leafy branch has been observed, as in Lysimachia, where in one case
the prolonged placenta was removed and struck as a cutting.[120]
In Ericaceæ too, the axile placenta has been seen ovuliferous at the base
and prolonged above into a leafy branch.[121]
Median floral prolification.—This is of more frequent occurrence than the
preceding. The prolonged axis is more frequently terminated by a flower-
bud than by a leaf-bud, though it must be remarked, that the lengthened and
protruded stem frequently bears leaves upon its sides, even if it terminate in
a flower, and thus the new growth partakes of a mixed leafy and floral
nature. Instances of this kind have long been familiar to observers, and have
always excited attention from the singularity of their appearance. In one of
the old stained-glass windows, apparently of Dutch manufacture, in the
Bodleian Picture Gallery at Oxford, is a representation of a Ranunculus
affected with median floral prolification.[122] In pinks the affection is not
unfrequently met with. Fig. 60 shows an instance of the kind copied from
Schotterbec.
A singular instance of prolification in the central flower of one of the
verticillasters of Phlomis fruticosa fell under my own notice; it was a case
Page 177
wherein the calyx was torn on one side, and one of its lobes had become
petaloid. Between the calyx and the corolla were three or four spathulate,
hairy, bract-like organs; the corolla and stamens were unchanged; but in
place of the usual four-lobed ovary there was a single carpel with a basilar
style, terminated by a forked stigma. Occupying the place of the other lobes
of the pistil was an oblong woolly flower-bud, consisting of calyx, corolla,
and stamens, but with no trace of pistil. I have been unable to find recorded
any instance of malformation among Labiates or Borages at all similar to
this. It differed from most other examples of prolification in that the axis
was not prolonged, the adventitious bud occupying precisely the position of
the three lobes of the ovary that were absent. The sole remaining carpel had
a style and a stigma as perfect in appearance as though the pistil had been
complete.
Fig. 60.—Flower of Dianthus affected with median floral prolification.
petaloid. Between the calyx and the corolla were three or four spathulate,
hairy, bract-like organs; the corolla and stamens were unchanged; but in
place of the usual four-lobed ovary there was a single carpel with a basilar
style, terminated by a forked stigma. Occupying the place of the other lobes
of the pistil was an oblong woolly flower-bud, consisting of calyx, corolla,
and stamens, but with no trace of pistil. I have been unable to find recorded
any instance of malformation among Labiates or Borages at all similar to
this. It differed from most other examples of prolification in that the axis
was not prolonged, the adventitious bud occupying precisely the position of
the three lobes of the ovary that were absent. The sole remaining carpel had
a style and a stigma as perfect in appearance as though the pistil had been
complete.
Fig. 60.—Flower of Dianthus affected with median floral prolification.
Page 178
In a flower of Conostephium (Epacridaceæ) forwarded to me by Mr.
Bentham, there was a similar adventitious bud placed by the side of the
pistil, but as the latter contained the usual number of cells it is probable that
the supernumerary bud in this case originated rather from the side than the
end of the axis.
Certain families of plants present this deviation from their ordinary
structure with greater frequency than others: the following orders seem to
be the most frequently affected by it: Ranunculaceæ, Caryophyllaceæ,
Rosaceæ; while it is commonly met with in Scrophulariaceæ, Primulaceæ
and Umbelliferæ. Of genera which seem peculiarly liable to it may be
mentioned the following: Anemone, Ranunculus, Cheiranthus, Dianthus,
Dictamnus, Daucus, Rosa, Geum, Pyrus, Trifolium, Antirrhinum, Digitalis,
Primula.
A reference to the subjoined list of genera affected by this malformation,
and the knowledge of its comparatively greater frequency in some than in
others of them, will show that it is more often met with in plants having an
indefinite form of inflorescence than in those having a definite one. The
change may affect some only, or the whole of the flowers constituting an
inflorescence; and though it is by no means a constant occurrence, it very
frequently happens that the central or terminal flower in a definite
inflorescence is alone affected, the others remaining in their ordinary
condition, as in pinks (Dianthus); and in the indefinite forms of
inflorescence, it is equally common that the uppermost flower or flowers
are the most liable to be thus affected.
In those plants which present this deviation from the ordinary condition
with the greatest frequency, it often happens that the axis is normally more
or less prolonged, either between the various whorls of the flower, as in the
case of the gynophore, &c., or into the cavity of the carpels, as in the
Bentham, there was a similar adventitious bud placed by the side of the
pistil, but as the latter contained the usual number of cells it is probable that
the supernumerary bud in this case originated rather from the side than the
end of the axis.
Certain families of plants present this deviation from their ordinary
structure with greater frequency than others: the following orders seem to
be the most frequently affected by it: Ranunculaceæ, Caryophyllaceæ,
Rosaceæ; while it is commonly met with in Scrophulariaceæ, Primulaceæ
and Umbelliferæ. Of genera which seem peculiarly liable to it may be
mentioned the following: Anemone, Ranunculus, Cheiranthus, Dianthus,
Dictamnus, Daucus, Rosa, Geum, Pyrus, Trifolium, Antirrhinum, Digitalis,
Primula.
A reference to the subjoined list of genera affected by this malformation,
and the knowledge of its comparatively greater frequency in some than in
others of them, will show that it is more often met with in plants having an
indefinite form of inflorescence than in those having a definite one. The
change may affect some only, or the whole of the flowers constituting an
inflorescence; and though it is by no means a constant occurrence, it very
frequently happens that the central or terminal flower in a definite
inflorescence is alone affected, the others remaining in their ordinary
condition, as in pinks (Dianthus); and in the indefinite forms of
inflorescence, it is equally common that the uppermost flower or flowers
are the most liable to be thus affected.
In those plants which present this deviation from the ordinary condition
with the greatest frequency, it often happens that the axis is normally more
or less prolonged, either between the various whorls of the flower, as in the
case of the gynophore, &c., or into the cavity of the carpels, as in the
Page 179
instances of free central placentation. To bear out this assertion, the
following instances taken from those genera having definite inflorescence,
and which are very commonly affected with prolification, may be cited;
thus, in Anemone and Ranunculus the thalamus is prolonged to bear the
numerous carpels; in Dianthus there is a marked internode separating the
carpels from the other parts of the flower; in Primulaceæ central
prolification is very common, and this is one of the orders where the
placenta seems from the researches of Duchartre and others, to be truly a
production of the axis within the carpels;[123] in Thesium also, another
genus with free central placenta, this malformation has been found.
So also among plants with indefinite inflorescence, prolification seems very
frequently to affect those wherein the axis is normally prolonged; thus it is
common in Dictamnus, which plant has an internode supporting the pistil; it
is frequent among Umbelliferæ, where the carpophore may be truly
considered an axile production; it is common among Rosaceæ and
Ranunculaceæ, in many of which the axis or thalamus is well-marked, and
it is by no means infrequent in the flowers of the Orange, where the floral
internodes are also slightly elongated; on the other hand, there is no case on
record in Magnoliaceæ, and some other orders where the floral part of the
axis is at some point or other elongated; still, on the whole, there can be but
little doubt that there is a real relation between prolification and the normal
extension of the floral internodes.
Under these circumstances, those instances wherein the parts of the flower
become separated one from the other by the elongation of the internodes
(apostatis), constitute a lesser degree of the same change, which operates
most completely in the formation of a new bud at the extremity of the
prolonged axis. Some specimens of Geum rivale (a plant very liable to
become prolified) in my possession show this very clearly. In the wild plant
the thalamus is elevated on a short stalk; in the abnormal ones the thalamus
following instances taken from those genera having definite inflorescence,
and which are very commonly affected with prolification, may be cited;
thus, in Anemone and Ranunculus the thalamus is prolonged to bear the
numerous carpels; in Dianthus there is a marked internode separating the
carpels from the other parts of the flower; in Primulaceæ central
prolification is very common, and this is one of the orders where the
placenta seems from the researches of Duchartre and others, to be truly a
production of the axis within the carpels;[123] in Thesium also, another
genus with free central placenta, this malformation has been found.
So also among plants with indefinite inflorescence, prolification seems very
frequently to affect those wherein the axis is normally prolonged; thus it is
common in Dictamnus, which plant has an internode supporting the pistil; it
is frequent among Umbelliferæ, where the carpophore may be truly
considered an axile production; it is common among Rosaceæ and
Ranunculaceæ, in many of which the axis or thalamus is well-marked, and
it is by no means infrequent in the flowers of the Orange, where the floral
internodes are also slightly elongated; on the other hand, there is no case on
record in Magnoliaceæ, and some other orders where the floral part of the
axis is at some point or other elongated; still, on the whole, there can be but
little doubt that there is a real relation between prolification and the normal
extension of the floral internodes.
Under these circumstances, those instances wherein the parts of the flower
become separated one from the other by the elongation of the internodes
(apostatis), constitute a lesser degree of the same change, which operates
most completely in the formation of a new bud at the extremity of the
prolonged axis. Some specimens of Geum rivale (a plant very liable to
become prolified) in my possession show this very clearly. In the wild plant
the thalamus is elevated on a short stalk; in the abnormal ones the thalamus
Page 180
is simply upon a longer stalk than usual, or in a more advanced stage of the
deviation the lengthened thalamus takes the form of a branch provided with
leaves and terminated by a flower; it is noticeable, also, in these specimens,
that the sepals of the lower flower have assumed entirely the dimensions
and appearance of leaves.
Median prolification has occasionally been recorded in flowers that have, in
their ordinary condition, but one carpel, as in Leguminosæ and in
Santalaceæ. In Leguminosæ, as also in Amygdalus, it would seem as if the
adventitious bud were strictly a lateral and axillary production, and
moreover that the carpel itself is not strictly terminal but lateral in position,
though apparently terminal from the abortion of other carpels. In the only
recorded instance that I am aware of, of this malformation affecting the
genus Thesium, the pistil was altogether absent, and occupying its place was
the new bud or branch.[124]
deviation the lengthened thalamus takes the form of a branch provided with
leaves and terminated by a flower; it is noticeable, also, in these specimens,
that the sepals of the lower flower have assumed entirely the dimensions
and appearance of leaves.
Median prolification has occasionally been recorded in flowers that have, in
their ordinary condition, but one carpel, as in Leguminosæ and in
Santalaceæ. In Leguminosæ, as also in Amygdalus, it would seem as if the
adventitious bud were strictly a lateral and axillary production, and
moreover that the carpel itself is not strictly terminal but lateral in position,
though apparently terminal from the abortion of other carpels. In the only
recorded instance that I am aware of, of this malformation affecting the
genus Thesium, the pistil was altogether absent, and occupying its place was
the new bud or branch.[124]
Page 181
Fig. 61.—Daucus Carota, showing leafly carpels, prolification, &c.
As the carpels are not unfrequently absent in cases of median prolification,
it has been thought that the pistil in such cases was metamorphosed into a
stem bearing leaves or flowers. Setting aside the physiological difficulties
in the way of accepting such an opinion, an examination of any number of
cases is sufficient to refute it; for, as Moquin well remarks, the carpels may
frequently be found either in an unaltered condition or more or less
modified.
If the pistil be normally syncarpous, its constituent carpels, if present at all
in the prolified flower, become disjoined one from the other to allow of the
passage between them of the prolonged axis; thus in some malformed
flowers of Daucus Carota gathered in Switzerland (fig. 61), not only was
the calyx partially detached from the pistil, but the carpels themselves were
leaf-like, disjoined, and unprovided with ovules; between them rose a
central prolongation of the axis, which almost immediately divided into two
branches, each terminated by a small umbel of perfect flowers, surrounded
by minute bracts.[125]
Not only are the carpels thus frequently separated one from the other by the
prolonged axis, but they undergo commonly a still further change in
becoming more or less completely foliaceous, as in the Daucus just
mentioned, where the carpels were prolonged into two lance-shaped leaves,
whose margins in some cases were slightly incurved at the apex, forcibly
calling to mind the long "beaks" that some Umbelliferous genera have
terminating their fruits—for instance, Scandix. Dr. Norman, in the fourth
series of the 'Annales des Sciences,' vol. ix, has described a prolification of
the flower of Anchusa ochroleuca, in which the pistil consisted of two
leaves, situated antero-posteriorly on a long internode, with a small terminal
flower-bud between them; and numerous similar instances might be cited.
As the carpels are not unfrequently absent in cases of median prolification,
it has been thought that the pistil in such cases was metamorphosed into a
stem bearing leaves or flowers. Setting aside the physiological difficulties
in the way of accepting such an opinion, an examination of any number of
cases is sufficient to refute it; for, as Moquin well remarks, the carpels may
frequently be found either in an unaltered condition or more or less
modified.
If the pistil be normally syncarpous, its constituent carpels, if present at all
in the prolified flower, become disjoined one from the other to allow of the
passage between them of the prolonged axis; thus in some malformed
flowers of Daucus Carota gathered in Switzerland (fig. 61), not only was
the calyx partially detached from the pistil, but the carpels themselves were
leaf-like, disjoined, and unprovided with ovules; between them rose a
central prolongation of the axis, which almost immediately divided into two
branches, each terminated by a small umbel of perfect flowers, surrounded
by minute bracts.[125]
Not only are the carpels thus frequently separated one from the other by the
prolonged axis, but they undergo commonly a still further change in
becoming more or less completely foliaceous, as in the Daucus just
mentioned, where the carpels were prolonged into two lance-shaped leaves,
whose margins in some cases were slightly incurved at the apex, forcibly
calling to mind the long "beaks" that some Umbelliferous genera have
terminating their fruits—for instance, Scandix. Dr. Norman, in the fourth
series of the 'Annales des Sciences,' vol. ix, has described a prolification of
the flower of Anchusa ochroleuca, in which the pistil consisted of two
leaves, situated antero-posteriorly on a long internode, with a small terminal
flower-bud between them; and numerous similar instances might be cited.
Page 182
In this place may also be noticed those instances wherein the placenta
elongates so much that the pericarp becomes ruptured to allow of the
protrusion of the placenta, although this prolongation is not attended by the
formation of new buds. Cases of this kind occurring in Melastoma and
Solanum have been put on record by M. Alph. de Candolle.[126] This is a
change analogous with that which occurs in some species of Leontice or
Caulophyllum, as commented on by Robert Brown. See 'Miscellaneous
Botanical Works' of this author, Ray Society, vol. i, p. 359.
If the pistil be apocarpous, and the carpels arranged spirally on an elevated
thalamus, it then frequently happens that the carpels, especially the upper
ones, become carried up with the prolonged axis, more widely separated
one from the other than below, and particularly liable to undergo various
petalloid or foliaceous changes as in proliferous Roses, Potentilla, &c.
Fig. 62.—Median floral prolification, &c., in flower of Delphinium.
elongates so much that the pericarp becomes ruptured to allow of the
protrusion of the placenta, although this prolongation is not attended by the
formation of new buds. Cases of this kind occurring in Melastoma and
Solanum have been put on record by M. Alph. de Candolle.[126] This is a
change analogous with that which occurs in some species of Leontice or
Caulophyllum, as commented on by Robert Brown. See 'Miscellaneous
Botanical Works' of this author, Ray Society, vol. i, p. 359.
If the pistil be apocarpous, and the carpels arranged spirally on an elevated
thalamus, it then frequently happens that the carpels, especially the upper
ones, become carried up with the prolonged axis, more widely separated
one from the other than below, and particularly liable to undergo various
petalloid or foliaceous changes as in proliferous Roses, Potentilla, &c.
Fig. 62.—Median floral prolification, &c., in flower of Delphinium.
Page 183
Fig. 62, copied from Cramer, shows an instance of this kind in Delphinium
elatum, where not only is the thalamus prolonged, and the carpels
separated, but from the axils of some of the latter which have assumed from
the disunion of their margins somewhat of the appearance of leaves, other
flowering branches proceed—axillary prolification. If, on the other hand,
the carpels be few in number, and placed in a verticillate manner, the axis
then generally passes upwards without any change in the form or position
of the carpels being apparent, as in a proliferous columbine, figured in the
'Linnean Transactions,' vol. xxiii, tab. 34, fig. 5.
When a flower with the ovary naturally inferior or adherent to the calyx
becomes prolified, a change in the relative position of the calyx and ovary
almost necessarily takes place, the latter becoming superior or detached
from the calyx; this has been already alluded to in Umbelliferæ. In a species
of Campanula examined by me, the calyx was free, the corolla double, the
stamens with petaloid filaments, and in the place of the pistil there was a
bud consisting of several series of green bracts, arranged in threes, and
enclosing quite in the centre three carpellary leaves detached from one
another and the other parts of the flower, and open along their margins,
where the ovules were placed. In other similar instances in the same species
of Campanula, the styles were present, forming below an imperfect tube
which surrounded the adventitious bud; in another, contrary to what occurs
usually in such cases, the ovary was present in its usual position, but
surmounted by a bud of leafy scales, enclosed within the base of a tube
formed by the union of the styles. A similar relative change in the position
of the calyx and the ovary takes place when the Compositæ are affected
with central prolification, or even in that lesser degree of change which
merely consists in the separation and disunion of the parts of the flower, but
which in these flowers appear to be, as it were, the first stage towards
prolification. I owe to the kindness of Professor Oliver a sketch of a species
of Rudbeckia? showing this detachment of the calyx from the ovary. In a
elatum, where not only is the thalamus prolonged, and the carpels
separated, but from the axils of some of the latter which have assumed from
the disunion of their margins somewhat of the appearance of leaves, other
flowering branches proceed—axillary prolification. If, on the other hand,
the carpels be few in number, and placed in a verticillate manner, the axis
then generally passes upwards without any change in the form or position
of the carpels being apparent, as in a proliferous columbine, figured in the
'Linnean Transactions,' vol. xxiii, tab. 34, fig. 5.
When a flower with the ovary naturally inferior or adherent to the calyx
becomes prolified, a change in the relative position of the calyx and ovary
almost necessarily takes place, the latter becoming superior or detached
from the calyx; this has been already alluded to in Umbelliferæ. In a species
of Campanula examined by me, the calyx was free, the corolla double, the
stamens with petaloid filaments, and in the place of the pistil there was a
bud consisting of several series of green bracts, arranged in threes, and
enclosing quite in the centre three carpellary leaves detached from one
another and the other parts of the flower, and open along their margins,
where the ovules were placed. In other similar instances in the same species
of Campanula, the styles were present, forming below an imperfect tube
which surrounded the adventitious bud; in another, contrary to what occurs
usually in such cases, the ovary was present in its usual position, but
surmounted by a bud of leafy scales, enclosed within the base of a tube
formed by the union of the styles. A similar relative change in the position
of the calyx and the ovary takes place when the Compositæ are affected
with central prolification, or even in that lesser degree of change which
merely consists in the separation and disunion of the parts of the flower, but
which in these flowers appear to be, as it were, the first stage towards
prolification. I owe to the kindness of Professor Oliver a sketch of a species
of Rudbeckia? showing this detachment of the calyx from the ovary. In a
Page 184
monstrous Fuchsia that I have had the opportunity of recently examining,
the calyx was similarly detached from the ovary simultaneously with the
extension of the axis. Here the petals were increased in number and
variously modified, the stamens also; while in the centre and at the top of
the flower, conjoined at the base with some imperfect stamens, was a carpel
open along its ovuliferous margins. Such instances as these seem to be the
first stages of a change which, carried out more perfectly, would result in
the formation of a new bud on the extremity of the prolonged axis.
In Orchidaceæ, among which family I have now met with several instances
of prolification, the ovary seems usually to be absent. Fig. 63 shows a
prolified flower of Orchis pyramidalis in which the perianth was nearly
regular, the central portions of the flower absent, and their place supplied by
a new miniature raceme. This specimen was forwarded to me by Dr. Moore,
of Glasnevin.
the calyx was similarly detached from the ovary simultaneously with the
extension of the axis. Here the petals were increased in number and
variously modified, the stamens also; while in the centre and at the top of
the flower, conjoined at the base with some imperfect stamens, was a carpel
open along its ovuliferous margins. Such instances as these seem to be the
first stages of a change which, carried out more perfectly, would result in
the formation of a new bud on the extremity of the prolonged axis.
In Orchidaceæ, among which family I have now met with several instances
of prolification, the ovary seems usually to be absent. Fig. 63 shows a
prolified flower of Orchis pyramidalis in which the perianth was nearly
regular, the central portions of the flower absent, and their place supplied by
a new miniature raceme. This specimen was forwarded to me by Dr. Moore,
of Glasnevin.
Page 185
Fig. 63.—Median prolification in Orchis pyramidalis, the outer segments of the perianth regular and
reflexed.
As might be expected, it very rarely happens that median prolification
occurs without some other deviation in one or more parts of the flower
being simultaneously manifested. Some of these changes have been already
mentioned, but others are commonly met with, as, for instance, the
multiplication or doubling, as it is termed, of the petals; others, though less
frequent, are of more interest. Fusion of two or more flowers in association
with prolification is especially common in cultivated specimens of Digitalis
purpurea; the uppermost flowers of the raceme become fused together so as
to form one large, regular, erect, cup-shaped corolla, to the tube of which
the stamens are attached, in greater number than ordinary, and all of equal
length; the bracts and sepals are confusedly arranged on the exterior of the
flower; while in the centre, in the place usually occupied by the pistil, there
rises a conical prolongation of the axis, bearing at its outer or lower portion
a number of open carpels, provided, it may be, with styles and ovules; these
enclose an inner series of scale-like bracts, from whose axils proceed more
or less perfect florets; so that in the most highly developed stage a perfect
raceme of flowers may be seen to spring from the centre of a cup-shaped
regular flower, whose lobes show its compound character. All intermediate
stages of this malformation may be found from cases where there is a
simple fusion of two flowers with a second verticil of carpels within the
outer, up to such cases as those which have been just mentioned. It is
worthy of special remark, that in all these cases the flowers at the
uppermost part of the raceme are alone affected, and that, in addition to the
prolification, there is fusion of two or more flowers, and regularity in the
form of the compound corolla and stamens.
The calyx of a prolified flower is either unchanged, or it is modified in
harmony with the changes in the central part of the flower. If the ovary be
normally superior or free from the calyx, then the latter is comparatively
reflexed.
As might be expected, it very rarely happens that median prolification
occurs without some other deviation in one or more parts of the flower
being simultaneously manifested. Some of these changes have been already
mentioned, but others are commonly met with, as, for instance, the
multiplication or doubling, as it is termed, of the petals; others, though less
frequent, are of more interest. Fusion of two or more flowers in association
with prolification is especially common in cultivated specimens of Digitalis
purpurea; the uppermost flowers of the raceme become fused together so as
to form one large, regular, erect, cup-shaped corolla, to the tube of which
the stamens are attached, in greater number than ordinary, and all of equal
length; the bracts and sepals are confusedly arranged on the exterior of the
flower; while in the centre, in the place usually occupied by the pistil, there
rises a conical prolongation of the axis, bearing at its outer or lower portion
a number of open carpels, provided, it may be, with styles and ovules; these
enclose an inner series of scale-like bracts, from whose axils proceed more
or less perfect florets; so that in the most highly developed stage a perfect
raceme of flowers may be seen to spring from the centre of a cup-shaped
regular flower, whose lobes show its compound character. All intermediate
stages of this malformation may be found from cases where there is a
simple fusion of two flowers with a second verticil of carpels within the
outer, up to such cases as those which have been just mentioned. It is
worthy of special remark, that in all these cases the flowers at the
uppermost part of the raceme are alone affected, and that, in addition to the
prolification, there is fusion of two or more flowers, and regularity in the
form of the compound corolla and stamens.
The calyx of a prolified flower is either unchanged, or it is modified in
harmony with the changes in the central part of the flower. If the ovary be
normally superior or free from the calyx, then the latter is comparatively
Page 186
rarely altered; for instance, in proliferous pinks (Dianthus) the calyx is
seldom affected, except, indeed, in those instances where the floral axis is
prolonged, and produces from its side a successive series of sepals, as in
what is called the wheat-ear carnation; but though these instances may be,
as I believe, an imperfect degree of prolification, they do not affect the
general truth of the above opinion, that the calyx, if it be free from the
ovary, is but rarely changed in a prolified flower; but that this is not a
universal rule is shown by proliferous flowers of Geum rivale, where the
sepals are usually large and leaf-like, as they likewise are frequently in
proliferous roses and pears.
Fig. 64.—Proliferous rose. Hip absent, sepals leafy, stamens wanting, axis prolonged bearing
supplementary flower, &c. (Bell Salter).
Proliferous roses have a special interest, inasmuch as they show very
conclusively that the so-called calyx-tube of these plants is merely a
concave and inverted thalamus, which, in prolified specimens, becomes
seldom affected, except, indeed, in those instances where the floral axis is
prolonged, and produces from its side a successive series of sepals, as in
what is called the wheat-ear carnation; but though these instances may be,
as I believe, an imperfect degree of prolification, they do not affect the
general truth of the above opinion, that the calyx, if it be free from the
ovary, is but rarely changed in a prolified flower; but that this is not a
universal rule is shown by proliferous flowers of Geum rivale, where the
sepals are usually large and leaf-like, as they likewise are frequently in
proliferous roses and pears.
Fig. 64.—Proliferous rose. Hip absent, sepals leafy, stamens wanting, axis prolonged bearing
supplementary flower, &c. (Bell Salter).
Proliferous roses have a special interest, inasmuch as they show very
conclusively that the so-called calyx-tube of these plants is merely a
concave and inverted thalamus, which, in prolified specimens, becomes
Page 187
elongated (fig. 64) after the fashion of Geum rivale, &c.[127] Occasionally
from the middle of the outer surface of the urn-shaped thalamus proceeds a
perfect leaf, which could hardly be produced from the united sepals or
calyx-tube; a similar occurrence in a pear is figured in Keith's
'Physiological Botany,' plate ix, fig. 12.
The change which the calyx undergoes when flowers with an habitually
adherent ovary become prolified, and wherein the calyx is disjoined from
the ovary, has been before mentioned, but it may also be stated that, under
such circumstances, the constituent sepals are frequently separated one
from the other, and not rarely assume more or less of the appearance of
leaves, as in proliferous flowers of Umbelliferæ, Campanulaceæ,
Compositæ, &c.
As to the corolla, it was long since noticed that prolification was especially
liable to occur in double flowers; indeed, Dr. Hill, who published a treatise
on this subject, setting forth the method of artificially producing prolified
flowers, deemed the doubling to be an almost necessary precursor of
prolification;[128] but, though frequently so, it is not invariably the case that
the flower so affected is double—e.g. Geum. If double, the doubling may
arise from actual multiplication of the petals, or from the substitution of
petals for stamens and pistils, according to the particular plant affected.
Occasionally in prolified flowers the parts of the corolla, like those of the
calyx, become foliaceous, and in the case of proliferous pears fleshy and
succulent. There is in cultivation a kind of Cheiranthus? in which there is a
constant repetition of the calyx and corolla, conjoined with an entire
absence of the stamens and pistils; a short internode separates each flower
from the one above it, and thus frequently ten or a dozen of these imperfect
flowers may be seen on the end of a flower-stalk, giving an appearance as if
they were strung like beads, at regular intervals, on a common stalk. I have
seen a similar instance in a less degree in a species of Helianthemum.
from the middle of the outer surface of the urn-shaped thalamus proceeds a
perfect leaf, which could hardly be produced from the united sepals or
calyx-tube; a similar occurrence in a pear is figured in Keith's
'Physiological Botany,' plate ix, fig. 12.
The change which the calyx undergoes when flowers with an habitually
adherent ovary become prolified, and wherein the calyx is disjoined from
the ovary, has been before mentioned, but it may also be stated that, under
such circumstances, the constituent sepals are frequently separated one
from the other, and not rarely assume more or less of the appearance of
leaves, as in proliferous flowers of Umbelliferæ, Campanulaceæ,
Compositæ, &c.
As to the corolla, it was long since noticed that prolification was especially
liable to occur in double flowers; indeed, Dr. Hill, who published a treatise
on this subject, setting forth the method of artificially producing prolified
flowers, deemed the doubling to be an almost necessary precursor of
prolification;[128] but, though frequently so, it is not invariably the case that
the flower so affected is double—e.g. Geum. If double, the doubling may
arise from actual multiplication of the petals, or from the substitution of
petals for stamens and pistils, according to the particular plant affected.
Occasionally in prolified flowers the parts of the corolla, like those of the
calyx, become foliaceous, and in the case of proliferous pears fleshy and
succulent. There is in cultivation a kind of Cheiranthus? in which there is a
constant repetition of the calyx and corolla, conjoined with an entire
absence of the stamens and pistils; a short internode separates each flower
from the one above it, and thus frequently ten or a dozen of these imperfect
flowers may be seen on the end of a flower-stalk, giving an appearance as if
they were strung like beads, at regular intervals, on a common stalk. I have
seen a similar instance in a less degree in a species of Helianthemum.
Page 188
The stamens are subject to various changes in prolified flowers; they
assume, for instance, a leaf-like or petal-like condition, or take on them
more or less of a carpellary form, or they may be entirely absent; but none
of these changes seem to be at all necessarily connected with the proliferous
state of the flower. Of more interest is the alteration in the position of these
organs which sometimes necessarily accrues from the elongation of the axis
and the disjunction of the calyx; thus, in proliferous roses the stamens
become strictly hypogynous, instead of remaining perigynous. In
Umbelliferæ the epigynous condition is changed for the perigynous, &c.
The condition of the pistillary organs in prolified flowers has already been
alluded to. Hitherto those instances have been considered in which either
the carpels were absent, or the new bud proceeded from between the
carpels. There is also an interesting class of cases where the prolification is
strictly intra-carpellary; the axis is so slightly prolonged that it does not
protrude beyond the carpels, does not separate them in any way, but is
wholly enclosed within their cavity. Doubtless, in many cases, this is
merely a less perfect development of that change in which the axis
protrudes beyond the carpels. This intra-carpellary prolification occurs most
frequently in plants having a free central placenta, though it is not confined
to them, as it is recorded among Boragineæ. A remarkable instance of this is
described by Mr. H. C. Watson in the first volume of Henfrey's 'Botanical
Gazette,' p. 88. In this specimen a raceme of small flowers was included
within the enlarged pericarp of a species of Anchusa. But the most curious
instances of this form of prolification are, no doubt, those which are met
with among Primulaceæ and other orders with free central placentation.
Duchartre, in his memoir on the organogeny of plants with a free central
placenta, in the 'Ann. des Sc. Nat.,' 3 sér., 1844, p. 290, among other similar
instances, mentions two flowers of Cortusa Matthioli, wherein the placenta
was ovuliferous at the base; but the upper portion, instead of simply
assume, for instance, a leaf-like or petal-like condition, or take on them
more or less of a carpellary form, or they may be entirely absent; but none
of these changes seem to be at all necessarily connected with the proliferous
state of the flower. Of more interest is the alteration in the position of these
organs which sometimes necessarily accrues from the elongation of the axis
and the disjunction of the calyx; thus, in proliferous roses the stamens
become strictly hypogynous, instead of remaining perigynous. In
Umbelliferæ the epigynous condition is changed for the perigynous, &c.
The condition of the pistillary organs in prolified flowers has already been
alluded to. Hitherto those instances have been considered in which either
the carpels were absent, or the new bud proceeded from between the
carpels. There is also an interesting class of cases where the prolification is
strictly intra-carpellary; the axis is so slightly prolonged that it does not
protrude beyond the carpels, does not separate them in any way, but is
wholly enclosed within their cavity. Doubtless, in many cases, this is
merely a less perfect development of that change in which the axis
protrudes beyond the carpels. This intra-carpellary prolification occurs most
frequently in plants having a free central placenta, though it is not confined
to them, as it is recorded among Boragineæ. A remarkable instance of this is
described by Mr. H. C. Watson in the first volume of Henfrey's 'Botanical
Gazette,' p. 88. In this specimen a raceme of small flowers was included
within the enlarged pericarp of a species of Anchusa. But the most curious
instances of this form of prolification are, no doubt, those which are met
with among Primulaceæ and other orders with free central placentation.
Duchartre, in his memoir on the organogeny of plants with a free central
placenta, in the 'Ann. des Sc. Nat.,' 3 sér., 1844, p. 290, among other similar
instances, mentions two flowers of Cortusa Matthioli, wherein the placenta
was ovuliferous at the base; but the upper portion, instead of simply
Page 189
elongating itself into a sterile cone, had produced a little flower with its
parts slightly different from those of the normal flowers. M. Alph. de
Candolle has likewise described somewhat similar deviations, and one in
particular in Primula Auricula, where the elongated placenta gave off long
and dilated funiculi bearing ovules, while other funiculi were destitute of
these bodies, but were much dilated and foliaceous in appearance.[129] In
some flowers of Rhododendron I have observed a similar condition of the
ovules, which, moreover, in the primary flowers, were attached to the walls
of the carpels—parietal placentation.
In speaking of these as cases of intra-carpellary prolification, it is, of
course, impossible to overlook the fact that they differ in degree only from
those cases where the lengthened axis projects beyond the cavity of the
carpels; nevertheless they seem to demand special notice, because in these
particular plants the placenta or its prolongation appears never to protrude
beyond the carpels, or at least very rarely. There are, however, numerous
instances of such an extension of the placenta and of prolification occurring
among Primulaceæ in conjunction with the more or less complete arrest of
growth of the carpels.[130] An instance of this kind has come under my own
notice in a monstrosity of the chinese primrose, in which the carpels were
reduced to a hardly discernible rim surrounding an umbel of five rays, each
terminated by a small normally constituted flower-bud.
The ovules of a prolified flower are either unaffected, or they occur in a
rudimentary form, or, lastly, they may be present in the guise of small
leaves.
Under the term prolification of the fruit two or three distinct kinds of
malformation appear to have been included. The term seems usually to be
applied to those cases where from the centre of one fruit a branch bearing
leaves, flowers, or another fruit, is seen to project, as happens occasionally
parts slightly different from those of the normal flowers. M. Alph. de
Candolle has likewise described somewhat similar deviations, and one in
particular in Primula Auricula, where the elongated placenta gave off long
and dilated funiculi bearing ovules, while other funiculi were destitute of
these bodies, but were much dilated and foliaceous in appearance.[129] In
some flowers of Rhododendron I have observed a similar condition of the
ovules, which, moreover, in the primary flowers, were attached to the walls
of the carpels—parietal placentation.
In speaking of these as cases of intra-carpellary prolification, it is, of
course, impossible to overlook the fact that they differ in degree only from
those cases where the lengthened axis projects beyond the cavity of the
carpels; nevertheless they seem to demand special notice, because in these
particular plants the placenta or its prolongation appears never to protrude
beyond the carpels, or at least very rarely. There are, however, numerous
instances of such an extension of the placenta and of prolification occurring
among Primulaceæ in conjunction with the more or less complete arrest of
growth of the carpels.[130] An instance of this kind has come under my own
notice in a monstrosity of the chinese primrose, in which the carpels were
reduced to a hardly discernible rim surrounding an umbel of five rays, each
terminated by a small normally constituted flower-bud.
The ovules of a prolified flower are either unaffected, or they occur in a
rudimentary form, or, lastly, they may be present in the guise of small
leaves.
Under the term prolification of the fruit two or three distinct kinds of
malformation appear to have been included. The term seems usually to be
applied to those cases where from the centre of one fruit a branch bearing
leaves, flowers, or another fruit, is seen to project, as happens occasionally
Page 190
in pears. Now, in many instances, not only the fruit, is repeated, but also the
outer portions of the flower, which wither and fall away as the adventitious
fruit ripens; so that at length the phenomenon of one fruit projecting from
another is produced. It is obvious that this form of prolification in no wise
differs from ordinary central prolification. Sometimes some of the whorls of
the adventitious flower are suppressed; thus, M. Duchartre describes some
orange blossoms as presenting alternating series of stamens and pistils one
above another, while the calyces and corollas belonging to each series of
stamens and pistils were entirely suppressed.[131] In other cases, doubtless,
the carpellary whorl is alone repeated, the other whorls of the adventitious
flower being completely absent.
Another condition, apparently sometimes mistaken for prolification of the
fruit, is that in which the carpellary whorl becomes multiplied; so that there
is a second or even a third series within the outer whorl of carpels. If the
axis be at all prolonged, then these whorls are separated one from the other,
and produce in this way an appearance of prolification. This happens
frequently in oranges, as in the variety called Mellarose.[132]
Moquin has given an explanation of the St. Valery Apples, wherein the
petals are sepaloid, the stamens absent, and where there is a double row of
carpels, by supposing these peculiarities to be due to "a prolification
combined with penetration and fusion of two or more flowers," but it is
surely more reasonable to conceive a second row of carpels placed above
the first by the prolongation of the central part of the axis. Supposing this
view to be correct, the inner calyx-like whorl might be considered either as
a repetition of the calycine whorl, or it might be inferred that the corolla
was present in the guise of a second calyx.
Moquin-Tandon suggests another explanation—namely, that though the
stamens are absent in these curious flowers, at least in their ordinary shape,
outer portions of the flower, which wither and fall away as the adventitious
fruit ripens; so that at length the phenomenon of one fruit projecting from
another is produced. It is obvious that this form of prolification in no wise
differs from ordinary central prolification. Sometimes some of the whorls of
the adventitious flower are suppressed; thus, M. Duchartre describes some
orange blossoms as presenting alternating series of stamens and pistils one
above another, while the calyces and corollas belonging to each series of
stamens and pistils were entirely suppressed.[131] In other cases, doubtless,
the carpellary whorl is alone repeated, the other whorls of the adventitious
flower being completely absent.
Another condition, apparently sometimes mistaken for prolification of the
fruit, is that in which the carpellary whorl becomes multiplied; so that there
is a second or even a third series within the outer whorl of carpels. If the
axis be at all prolonged, then these whorls are separated one from the other,
and produce in this way an appearance of prolification. This happens
frequently in oranges, as in the variety called Mellarose.[132]
Moquin has given an explanation of the St. Valery Apples, wherein the
petals are sepaloid, the stamens absent, and where there is a double row of
carpels, by supposing these peculiarities to be due to "a prolification
combined with penetration and fusion of two or more flowers," but it is
surely more reasonable to conceive a second row of carpels placed above
the first by the prolongation of the central part of the axis. Supposing this
view to be correct, the inner calyx-like whorl might be considered either as
a repetition of the calycine whorl, or it might be inferred that the corolla
was present in the guise of a second calyx.
Moquin-Tandon suggests another explanation—namely, that though the
stamens are absent in these curious flowers, at least in their ordinary shape,
Page 191
they are represented by the lower row of carpels, which become, in process
of development, fused with the upper or true carpels. If this were so, surely
some intermediate conditions between stamen and carpel would
occasionally be present; but such does not appear to be the case.[133]
In some of the instances of so-called proliferous pears the carpels would
seem to be entirely absent, and the dilated portion of the axis to be alone
repeated. Thus, the axis dilates to form the lower fruit without any true
carpels being produced, but at its summit a whorl of leaves (sepals) is
formed; above these another swelling of the axis takes place also without
the formation of carpels, and this, it may be, is terminated in its turn by a
branch producing leaves. In these cases there is no true prolification, but
simply an extension of the axis. That the outer portion (so-called calyx-
tube) of these fruits is really an axile product there can now be little doubt;
and, as if to show their axile nature, they occasionally produce leaves from
their sides, as before mentioned. Moquin, in the tenth volume of the
'Bulletin of the Botanical Society of France,' p. 73, says that when the case
is one of prolification the lower fruit is larger and is formed of a fleshy
mass; moreover, the line of demarcation between the fruits is more distinct,
and there are traces of the seed-bearing cavity in the interior, and of
calycine lobes at the top. On the other hand, if the case be one of
hypertrophy merely, the lowermost fruit is the smallest, and there is no trace
of seed-bearing cavity nor of sepals. See also under Hypertrophy.
Some other malformations usually referred to prolification of the fruit seem
due to branching of the inflorescence, as in Plantago, wheat, maize; or to a
simple extension of the axis beyond its ordinary limit, as in some cones of
firs, &c. It is obvious that the true fruits in these cases are in no wise
affected.
of development, fused with the upper or true carpels. If this were so, surely
some intermediate conditions between stamen and carpel would
occasionally be present; but such does not appear to be the case.[133]
In some of the instances of so-called proliferous pears the carpels would
seem to be entirely absent, and the dilated portion of the axis to be alone
repeated. Thus, the axis dilates to form the lower fruit without any true
carpels being produced, but at its summit a whorl of leaves (sepals) is
formed; above these another swelling of the axis takes place also without
the formation of carpels, and this, it may be, is terminated in its turn by a
branch producing leaves. In these cases there is no true prolification, but
simply an extension of the axis. That the outer portion (so-called calyx-
tube) of these fruits is really an axile product there can now be little doubt;
and, as if to show their axile nature, they occasionally produce leaves from
their sides, as before mentioned. Moquin, in the tenth volume of the
'Bulletin of the Botanical Society of France,' p. 73, says that when the case
is one of prolification the lower fruit is larger and is formed of a fleshy
mass; moreover, the line of demarcation between the fruits is more distinct,
and there are traces of the seed-bearing cavity in the interior, and of
calycine lobes at the top. On the other hand, if the case be one of
hypertrophy merely, the lowermost fruit is the smallest, and there is no trace
of seed-bearing cavity nor of sepals. See also under Hypertrophy.
Some other malformations usually referred to prolification of the fruit seem
due to branching of the inflorescence, as in Plantago, wheat, maize; or to a
simple extension of the axis beyond its ordinary limit, as in some cones of
firs, &c. It is obvious that the true fruits in these cases are in no wise
affected.
Page 192
From these considerations it would appear better to abandon the use of the
expression prolification of the fruit, as unnecessary where it is really
applicable, and as delusive in the numerous other cases where it is
employed.
Median prolification of one or other kind has been met with in the
following genera:
Leafy. Floral.
Ranunculaceæ. Clematis.
Anemone! *Anemone!
Ranunculus! *Ranunculus!
Delphinium.
Caltha.
Aquilegia!
Cruciferæ. Bunias.
*Cheiranthus!
Erucago.
*Matthiola!
Sisymbrium!
Brassica!
Nasturtium.
Hesperis.
Sinapis!
Diplotaxis.
Lunaria.
Erysimum.
Alyssum.
Peltaria.
Cardamine!
Cleome.
Cistaceæ. Helianthemum!
Caryophylleæ. Dianthus! *Dianthus!
Silene!
Lychnis!
Violaceæ. Viola!
Tiliaceæ. Triumfetta!
Geraniaceæ. Geranium!
Sapindaceæ. Pavia! Pavia!
Malvaceæ. Paritium.
Hibiscus!
Malpighiaceæ. Byrsonima!
Rutaceæ. Genera not specified. *Dictamnus!
expression prolification of the fruit, as unnecessary where it is really
applicable, and as delusive in the numerous other cases where it is
employed.
Median prolification of one or other kind has been met with in the
following genera:
Leafy. Floral.
Ranunculaceæ. Clematis.
Anemone! *Anemone!
Ranunculus! *Ranunculus!
Delphinium.
Caltha.
Aquilegia!
Cruciferæ. Bunias.
*Cheiranthus!
Erucago.
*Matthiola!
Sisymbrium!
Brassica!
Nasturtium.
Hesperis.
Sinapis!
Diplotaxis.
Lunaria.
Erysimum.
Alyssum.
Peltaria.
Cardamine!
Cleome.
Cistaceæ. Helianthemum!
Caryophylleæ. Dianthus! *Dianthus!
Silene!
Lychnis!
Violaceæ. Viola!
Tiliaceæ. Triumfetta!
Geraniaceæ. Geranium!
Sapindaceæ. Pavia! Pavia!
Malvaceæ. Paritium.
Hibiscus!
Malpighiaceæ. Byrsonima!
Rutaceæ. Genera not specified. *Dictamnus!
Page 193
Resedaceæ. Reseda.
Caylussa!
Aurantiaceæ. *Citrus!
Vitaceæ. Vitis. Vitis.
Umbelliferæ. Heracleum.
Angelica.
Thysselinum.
*Athamanta.
*Daucus!
*Torilis.
Rosaceæ. *Rosa! *Rosa!
*Geum! *Geum!
Agrimonia. Amygdalus.
Prunus!
Spiræa! Spiræa!
Rubus.
*Pyrus! *Pyrus!
?Leguminosæ Trifolium!
Medicago!
Melilotus.
Pisum!
Cucurbitaceæ. Cucumis.
Passifloraceæ. Passiflora.
Philadelphaceæ. Philadelphus.
Onagraceæ. Epilobium!
Epacridaceæ. Epacris!
Ericaceæ. *Erica.
Rhododendron!
Convolvulaceæ. Convolvulus.
Gentianaceæ. Gentiana. Gentiana.
Apocynaceæ. Vinca.
Jasminaceæ. Jasminum!
Scrophulariaceæ. Verbascum! Antirrhinum!
*Digitalis!
*Linaria!
Veronica.
Orobanchaceæ. Orobanche.
Labiatæ. Genera not specified. Stachys.
Phlomis!
Hydrophyllaceæ. Hydrophyllum.
Boraginaceæ. Anchusa.
Symphytum.
Primulaceæ. *Dodecatheon. *Cortusa.
*Anagallis! *Anagallis!
*Primula.
Dipsacaceæ. Scabiosa.
Caylussa!
Aurantiaceæ. *Citrus!
Vitaceæ. Vitis. Vitis.
Umbelliferæ. Heracleum.
Angelica.
Thysselinum.
*Athamanta.
*Daucus!
*Torilis.
Rosaceæ. *Rosa! *Rosa!
*Geum! *Geum!
Agrimonia. Amygdalus.
Prunus!
Spiræa! Spiræa!
Rubus.
*Pyrus! *Pyrus!
?Leguminosæ Trifolium!
Medicago!
Melilotus.
Pisum!
Cucurbitaceæ. Cucumis.
Passifloraceæ. Passiflora.
Philadelphaceæ. Philadelphus.
Onagraceæ. Epilobium!
Epacridaceæ. Epacris!
Ericaceæ. *Erica.
Rhododendron!
Convolvulaceæ. Convolvulus.
Gentianaceæ. Gentiana. Gentiana.
Apocynaceæ. Vinca.
Jasminaceæ. Jasminum!
Scrophulariaceæ. Verbascum! Antirrhinum!
*Digitalis!
*Linaria!
Veronica.
Orobanchaceæ. Orobanche.
Labiatæ. Genera not specified. Stachys.
Phlomis!
Hydrophyllaceæ. Hydrophyllum.
Boraginaceæ. Anchusa.
Symphytum.
Primulaceæ. *Dodecatheon. *Cortusa.
*Anagallis! *Anagallis!
*Primula.
Dipsacaceæ. Scabiosa.
Page 194
Compositæ. Hieracium! Hieracium!
Cirsium. Cirsium.
Hypochæris. Calendula!
Spilanthes.
Carthamus.
Coreopsis.
Campanulaceæ. Campanula. *Campanula!
Polygonaceæ. Genera not specified. Rumex.
Santalaceæ. Thesium.
Liliaceæ. Genera not specified. Tulipa!
Hemerocallis!
Asphodelus.
Hyacinthus!
Iridaceæ. Iris.
Amaryllidaceæ. Narcissus!
Leucojum.
Orchidaceæ. Orchis!
Habenaria.
Cyperaceæ. Carex.
Gramineæ. Phleum.
Axillary prolification is the term applied to those cases wherein one or
more adventitious buds spring from the axils of one or more of the parts of
the flower. Engelmann makes use of the word ecblastesis to denote the
same condition. Both terms are open to the objection that they do not
clearly enable us to distinguish prolification occurring within the flower
from a similar state originating outside the flower, within the bracts of the
inflorescence. This latter condition, called by Moquin-Tandon lateral
prolification (see Prolification of the Inflorescence), is as truly axillary as
that to which the name is restricted. In consequence of certain peculiarities
in the structure of some flowers, to be hereafter alluded to, it is not in all
cases easy to decide whether the new growth springs from the interior of the
flower, or from the inflorescence beneath the flower.
The accessory bud presents itself as a leaf-bud, a branch, a flower-bud, or a
miniature inflorescence; it may be sessile, but is far more frequently
stalked, and in more than half the number of cases it is a flower-bud or an
inflorescence. There may be one or more of these buds; if two only, then
Cirsium. Cirsium.
Hypochæris. Calendula!
Spilanthes.
Carthamus.
Coreopsis.
Campanulaceæ. Campanula. *Campanula!
Polygonaceæ. Genera not specified. Rumex.
Santalaceæ. Thesium.
Liliaceæ. Genera not specified. Tulipa!
Hemerocallis!
Asphodelus.
Hyacinthus!
Iridaceæ. Iris.
Amaryllidaceæ. Narcissus!
Leucojum.
Orchidaceæ. Orchis!
Habenaria.
Cyperaceæ. Carex.
Gramineæ. Phleum.
Axillary prolification is the term applied to those cases wherein one or
more adventitious buds spring from the axils of one or more of the parts of
the flower. Engelmann makes use of the word ecblastesis to denote the
same condition. Both terms are open to the objection that they do not
clearly enable us to distinguish prolification occurring within the flower
from a similar state originating outside the flower, within the bracts of the
inflorescence. This latter condition, called by Moquin-Tandon lateral
prolification (see Prolification of the Inflorescence), is as truly axillary as
that to which the name is restricted. In consequence of certain peculiarities
in the structure of some flowers, to be hereafter alluded to, it is not in all
cases easy to decide whether the new growth springs from the interior of the
flower, or from the inflorescence beneath the flower.
The accessory bud presents itself as a leaf-bud, a branch, a flower-bud, or a
miniature inflorescence; it may be sessile, but is far more frequently
stalked, and in more than half the number of cases it is a flower-bud or an
inflorescence. There may be one or more of these buds; if two only, then
Page 195
they are usually placed directly opposite one to the other, on the opposite
sides of the flower.
It will be seen, from the appended list, that the orders and genera in which
this description of adventitious growth occurs most frequently are the
following:—Cruciferæ, especially the genus Brassica; Caryophyllaceæ,
e.g. Dianthus; Resedaceæ; Leguminosæ, e.g. Melilotus, Trifolium, &c.;
Rosaceæ, e.g. Rosa, Potentilla, &c.; Umbelliferæ, and Campanulaceæ. For
the most part, these are groups also peculiarly liable to central prolification.
All the parts of the flower may be thus affected; but, as might have been
anticipated from the foliaceous nature of the sepals, the new bud usually
arises from within the axil of one of those organs. Next in frequency to the
calyx, the pistil is subjected to this change—the carpels in such a case being
disunited and leaf-like. The petals rank next, and lastly the stamens; these
latter, indeed, are usually, but not invariably, absent, the new growth
occupying their position. Hence it may well be that when such is the case,
there is no real axillary prolification, but rather the substitution of a bud for
a stamen. Generally, however, the position of the accessory bud is such that
it may properly be referred to the axil of an undeveloped or rudimentary
stamen.
The largest number of instances of this malformation, not merely
generically, but also individually, occurs in plants the members of whose
floral whorls are not united one to the other; thus, it is far more common in
polypetalous plants than in gamopetalous ones. In the prolified flowers
belonging to the latter group, the sepals, if not actually uncombined, are
only united for a short distance. The same relationship, but in a much less
degree, exists in the case of median prolification, as that aberration is
likewise most commonly met with in polypetalous flowers. Another feature
of interest is the rarity with which axillary prolification is found in irregular
sides of the flower.
It will be seen, from the appended list, that the orders and genera in which
this description of adventitious growth occurs most frequently are the
following:—Cruciferæ, especially the genus Brassica; Caryophyllaceæ,
e.g. Dianthus; Resedaceæ; Leguminosæ, e.g. Melilotus, Trifolium, &c.;
Rosaceæ, e.g. Rosa, Potentilla, &c.; Umbelliferæ, and Campanulaceæ. For
the most part, these are groups also peculiarly liable to central prolification.
All the parts of the flower may be thus affected; but, as might have been
anticipated from the foliaceous nature of the sepals, the new bud usually
arises from within the axil of one of those organs. Next in frequency to the
calyx, the pistil is subjected to this change—the carpels in such a case being
disunited and leaf-like. The petals rank next, and lastly the stamens; these
latter, indeed, are usually, but not invariably, absent, the new growth
occupying their position. Hence it may well be that when such is the case,
there is no real axillary prolification, but rather the substitution of a bud for
a stamen. Generally, however, the position of the accessory bud is such that
it may properly be referred to the axil of an undeveloped or rudimentary
stamen.
The largest number of instances of this malformation, not merely
generically, but also individually, occurs in plants the members of whose
floral whorls are not united one to the other; thus, it is far more common in
polypetalous plants than in gamopetalous ones. In the prolified flowers
belonging to the latter group, the sepals, if not actually uncombined, are
only united for a short distance. The same relationship, but in a much less
degree, exists in the case of median prolification, as that aberration is
likewise most commonly met with in polypetalous flowers. Another feature
of interest is the rarity with which axillary prolification is found in irregular
Page 196
gamopetalous blooms. It may be that the irregular and comparatively
excessive growth in some parts of these flowers, as compared with others,
may operate in checking any luxuriant tendency in other directions.
As in the case of median prolification, plants having an indefinite
inflorescence are more liable to be affected with ecblastesis than those
having a definite one. The degree of branching of the inflorescence may be
noticed, as this deformity is far more common in plants whose peduncles
are branched than in those which have either a solitary flower or an
unbranched flower-stalk. More than two thirds of the entire number of
genera cited as the subjects of this malformation have a branched
inflorescence of some form or other; and about two thirds of the cases occur
in genera having some form of indefinite inflorescence. If individual
instances could be accurately computed, the proportion would be even
higher.
Fully three fourths of the entire number of genera recorded as occasionally
the subjects of this irregularity possess in their usual state some peculiarity
of the thalamus; for instance, in about a third of the whole number of genera
the thalamus is more or less prolonged between some or other of the floral
whorl, e.g. Caryophyllaceæ, Potentilla, Anemone, Dictamnus, Umbelliferæ,
&c. About one fourth of the genera have numerous stamens or numerous
carpels, or both, springing naturally from the thalamus. In others (about one
sixth) the thalamus is enlarged into a disc, or else presents one or more
glandular swellings, e.g. Reseda, Nymphæa, Cruciferæ. In the last-named
family, as has been already remarked, prolification is very common. It
would be interesting to ascertain precisely what part of an inflorescence is
most liable to this affection; but as information on this point is but rarely
given in the records of these cases, I can only give the results of my own
observations, which go to show that, in a many-flowered inflorescence,
those flowers at the outside, or at the lower portion, seem to be more
excessive growth in some parts of these flowers, as compared with others,
may operate in checking any luxuriant tendency in other directions.
As in the case of median prolification, plants having an indefinite
inflorescence are more liable to be affected with ecblastesis than those
having a definite one. The degree of branching of the inflorescence may be
noticed, as this deformity is far more common in plants whose peduncles
are branched than in those which have either a solitary flower or an
unbranched flower-stalk. More than two thirds of the entire number of
genera cited as the subjects of this malformation have a branched
inflorescence of some form or other; and about two thirds of the cases occur
in genera having some form of indefinite inflorescence. If individual
instances could be accurately computed, the proportion would be even
higher.
Fully three fourths of the entire number of genera recorded as occasionally
the subjects of this irregularity possess in their usual state some peculiarity
of the thalamus; for instance, in about a third of the whole number of genera
the thalamus is more or less prolonged between some or other of the floral
whorl, e.g. Caryophyllaceæ, Potentilla, Anemone, Dictamnus, Umbelliferæ,
&c. About one fourth of the genera have numerous stamens or numerous
carpels, or both, springing naturally from the thalamus. In others (about one
sixth) the thalamus is enlarged into a disc, or else presents one or more
glandular swellings, e.g. Reseda, Nymphæa, Cruciferæ. In the last-named
family, as has been already remarked, prolification is very common. It
would be interesting to ascertain precisely what part of an inflorescence is
most liable to this affection; but as information on this point is but rarely
given in the records of these cases, I can only give the results of my own
observations, which go to show that, in a many-flowered inflorescence,
those flowers at the outside, or at the lower portion, seem to be more
Page 197
frequently the subjects of this change than those situated elsewhere. This
may probably be accounted for by the fact that the malformation is met with
most generally in plants with an indefinite form of inflorescence, and
therefore the lowermost or outermost flowers are most fully nourished; the
upper flowers being in a less advanced condition, the change is more likely
to be overlooked in them; or it may be that from the unusual luxuriance in
the lower flowers, the upper ones may be either present in their ordinary
condition, or may be (as indeed frequently happens) stunted in the size and
proportion of their several parts.
Axillary foliar prolification of the flower.—The formation of an
adventitious leaf-bud in the axil of any of the parts of the flower is not of
such common occurrence as the development of a flower-bud in similar
situations, nor is it so frequent as median foliar prolification. I have seen
leafy shoots proceeding from the axils of the sepals in the flowers of
Brassica, and a similar occurrence has been noticed in Caltha palustris,
Herreria parviflora, and other plants. Dr. Marchand's flowers of Anagallis,
previously referred to at p. 117, showed good illustrations of this
occurrence, as also some specimens described by Kirschleger in A.
phœnicea.[134] Steinheil has figured and described[135] a flower of Scabiosa
in which there was an adventitious formation of leafy shoots in the axil of
the outer calyx. In some flowers, such as Convolvulus, Anemone, &c., the
exact nature of the sub-floral leaves is uncertain, i.e. it is open to doubt
whether the organs in question are bracts or leaves pertaining to the
inflorescence, or whether they are really parts of the flower. When leafy
shoots are formed in the axils of such organs, the adventitious growth may
be referred to extra-floral prolification, prolification of the inflorescence
that is, or to axillary prolification, according to the view taken of the real
nature of the sub-floral leaves. So far as the mere occurrence of
prolification is concerned, it is not very material which view be adopted.
The same remark applies to cases where leaf-buds occur on the outer
may probably be accounted for by the fact that the malformation is met with
most generally in plants with an indefinite form of inflorescence, and
therefore the lowermost or outermost flowers are most fully nourished; the
upper flowers being in a less advanced condition, the change is more likely
to be overlooked in them; or it may be that from the unusual luxuriance in
the lower flowers, the upper ones may be either present in their ordinary
condition, or may be (as indeed frequently happens) stunted in the size and
proportion of their several parts.
Axillary foliar prolification of the flower.—The formation of an
adventitious leaf-bud in the axil of any of the parts of the flower is not of
such common occurrence as the development of a flower-bud in similar
situations, nor is it so frequent as median foliar prolification. I have seen
leafy shoots proceeding from the axils of the sepals in the flowers of
Brassica, and a similar occurrence has been noticed in Caltha palustris,
Herreria parviflora, and other plants. Dr. Marchand's flowers of Anagallis,
previously referred to at p. 117, showed good illustrations of this
occurrence, as also some specimens described by Kirschleger in A.
phœnicea.[134] Steinheil has figured and described[135] a flower of Scabiosa
in which there was an adventitious formation of leafy shoots in the axil of
the outer calyx. In some flowers, such as Convolvulus, Anemone, &c., the
exact nature of the sub-floral leaves is uncertain, i.e. it is open to doubt
whether the organs in question are bracts or leaves pertaining to the
inflorescence, or whether they are really parts of the flower. When leafy
shoots are formed in the axils of such organs, the adventitious growth may
be referred to extra-floral prolification, prolification of the inflorescence
that is, or to axillary prolification, according to the view taken of the real
nature of the sub-floral leaves. So far as the mere occurrence of
prolification is concerned, it is not very material which view be adopted.
The same remark applies to cases where leaf-buds occur on the outer
Page 198
surface of inferior ovaries, as in Rosaceæ, Pomaceæ, Philadelphus, or
Tetragonia expansa, as elsewhere mentioned.
It would seem more consistent with the general arrangements of parts, that
the adventitious buds should be formed more frequently outside than within
the flower proper.
Knight[136] figures and describes the occurrence of small tubers or fleshy
leaf-buds in the axils of the sepals of a potato, a curious illustration of the
real morphological nature of the tuber.
Axillary floral prolification of the flower.—As already stated, this is of
more common occurrence than the formation of a leaf-bud in a similar
situation. Any of the parts of the flower may thus subtend a flower-bud,
though probably the new buds more frequently originate in the axils of the
sepals than in the other whorls. In Cruciferæ the change in question is,
relatively speaking, very common. In cauliflowers and broccoli I have
frequently met with stalked flowers proceeding from the axils of the sepals,
so also in some fuchsias I have seen a ring of stalked flower-buds
alternating with the petals, which, together with the stamens and pistil,
remained unaffected. The number of parts in the supernumerary structures
is generally less than the normal flowers.
In Mr. Herbert Spencer's 'Principles of Biology,' part iv, p. 37, are figured
and described some monstrous inflorescences in Angelica and other
Umbelliferæ, from which, amongst other things, the author draws the
conclusion that there is no absolute distinction between leaf and branch.
Without staying for the moment to discuss this matter, it may here be said
that the Umbellifers in question apparently owe their peculiarities rather to
axillary prolification within the flower, or to prolification of the
Tetragonia expansa, as elsewhere mentioned.
It would seem more consistent with the general arrangements of parts, that
the adventitious buds should be formed more frequently outside than within
the flower proper.
Knight[136] figures and describes the occurrence of small tubers or fleshy
leaf-buds in the axils of the sepals of a potato, a curious illustration of the
real morphological nature of the tuber.
Axillary floral prolification of the flower.—As already stated, this is of
more common occurrence than the formation of a leaf-bud in a similar
situation. Any of the parts of the flower may thus subtend a flower-bud,
though probably the new buds more frequently originate in the axils of the
sepals than in the other whorls. In Cruciferæ the change in question is,
relatively speaking, very common. In cauliflowers and broccoli I have
frequently met with stalked flowers proceeding from the axils of the sepals,
so also in some fuchsias I have seen a ring of stalked flower-buds
alternating with the petals, which, together with the stamens and pistil,
remained unaffected. The number of parts in the supernumerary structures
is generally less than the normal flowers.
In Mr. Herbert Spencer's 'Principles of Biology,' part iv, p. 37, are figured
and described some monstrous inflorescences in Angelica and other
Umbelliferæ, from which, amongst other things, the author draws the
conclusion that there is no absolute distinction between leaf and branch.
Without staying for the moment to discuss this matter, it may here be said
that the Umbellifers in question apparently owe their peculiarities rather to
axillary prolification within the flower, or to prolification of the
Page 199
inflorescence, than to an actual transformation of a flower or any portion of
a flower into an umbellule.[137]
In the 'Gardeners' Chronicle,' 1855, p. 551, an instance is figured of the
production of a supernumerary flower proceeding from the axil of a stamen
in a species of Nymphæa (fig. 65). The ovary in this case was wanting, but
in its place was a tuft of small leaves. It is curious that among Dr. Kirk's
drawings of east tropical African plants now at Kew, there should be one
representing a precisely similar state of things. The species in both instances
was Nymphæa Lotus, or a cultivated variety of it.
M. Wesmael[138] describes a very singular case of what appears to have
been referable to axillary prolification in the flowers of Carex acuta. The
rachillus is described as prolonged through the utricle by the side of the
stigmas, bearing on its side a bract, then a secondary utricle, from the axil
of which sprung a short stem surmounted by an ovary. Wigand, 'Flora,'
1856, mentions a similar change in Carex glauca. In this instance the base
of the female inflorescence bore lateral spikes, which projected from the
utricles; some of these adventitious spikes were female, others female
below and male above, others, again, wholly male.
a flower into an umbellule.[137]
In the 'Gardeners' Chronicle,' 1855, p. 551, an instance is figured of the
production of a supernumerary flower proceeding from the axil of a stamen
in a species of Nymphæa (fig. 65). The ovary in this case was wanting, but
in its place was a tuft of small leaves. It is curious that among Dr. Kirk's
drawings of east tropical African plants now at Kew, there should be one
representing a precisely similar state of things. The species in both instances
was Nymphæa Lotus, or a cultivated variety of it.
M. Wesmael[138] describes a very singular case of what appears to have
been referable to axillary prolification in the flowers of Carex acuta. The
rachillus is described as prolonged through the utricle by the side of the
stigmas, bearing on its side a bract, then a secondary utricle, from the axil
of which sprung a short stem surmounted by an ovary. Wigand, 'Flora,'
1856, mentions a similar change in Carex glauca. In this instance the base
of the female inflorescence bore lateral spikes, which projected from the
utricles; some of these adventitious spikes were female, others female
below and male above, others, again, wholly male.
Page 200
Fig. 65.—Flower of Nymphæa Lotus, var., showing axillary floral prolification. The section also
shows the tuft of leaves that occupied the place of the ovary.
Various changes in the form and arrangement of the several floral whorls
accompany axillary prolification; some of these affect the particular organ
or organs implicated, and these only, while in other cases some other parts
of the flower likewise undergo modification. The changes most commonly
met with are such as may be classed under Goethe's theory of retrograde
metamorphosis; for instance, if a supplementary bud be developed in the
axil of a sepal, that sepal is likely to be more than ordinarily leaf-like in
appearance. The dislocation of the affected sepal from its fellows is a very
frequent occurrence; in cases of this kind the detached sepal is placed below
the others, thus approximating, in position as well as in function, to the
bracts. In some of the instances of proliferous pears, on which I shall have
occasion to comment, the sepals are described as sharing in the succulent
character of the fruit.
shows the tuft of leaves that occupied the place of the ovary.
Various changes in the form and arrangement of the several floral whorls
accompany axillary prolification; some of these affect the particular organ
or organs implicated, and these only, while in other cases some other parts
of the flower likewise undergo modification. The changes most commonly
met with are such as may be classed under Goethe's theory of retrograde
metamorphosis; for instance, if a supplementary bud be developed in the
axil of a sepal, that sepal is likely to be more than ordinarily leaf-like in
appearance. The dislocation of the affected sepal from its fellows is a very
frequent occurrence; in cases of this kind the detached sepal is placed below
the others, thus approximating, in position as well as in function, to the
bracts. In some of the instances of proliferous pears, on which I shall have
occasion to comment, the sepals are described as sharing in the succulent
character of the fruit.
Page 201
The petals, under such circumstances, often exist in the guise of sepals or of
small leaves; and instances are recorded wherein the place of the calyx and
corolla was supplied by a succession of overlapping green scales, from the
axils of which the new buds arose. M. Germain de Saint Pierre records such
a case in Trifolium repens, wherein the calyx and corolla were replaced by
overlapping scales, in the axils of each one of which arose a flower; above
there was a row of stamens, and in the centre a pistil in the guise of a
trifoliate leaf.[139] Such instances seem to afford an extreme degree of a
more common change, viz., the diminished size and contracted appearance
of the sepals and petals when affected with axillary prolification. They have
also a close relationship to such developments as we see in the wheat-ear
carnation, in certain species of the genus Mæsa and others, wherein the
calyx is repeated over and again, to the partial or complete suppression of
the other parts of the flower. All these cases may be in part explained by the
operation of the principle of compensation.
So far as the andrœcium is concerned, the stamens either remain unaltered,
or they are present in a more or less petal-like condition; but it far more
frequently happens that the stamens are entirely suppressed, the
adventitious bud supplying their place; thus was it in the Dianthus
represented in the adjoining woodcut, fig. 66, where the stamens were
entirely absent, and their places supplied by flower-bearing branches. This
Dianthus has the more interest from its similarity to the one described by
Goethe, Metam. der Pflanzen, cap. 16, sect. 105; but in that instance median
prolification also existed. For my specimens I am indebted to Mr. T. Moore.
small leaves; and instances are recorded wherein the place of the calyx and
corolla was supplied by a succession of overlapping green scales, from the
axils of which the new buds arose. M. Germain de Saint Pierre records such
a case in Trifolium repens, wherein the calyx and corolla were replaced by
overlapping scales, in the axils of each one of which arose a flower; above
there was a row of stamens, and in the centre a pistil in the guise of a
trifoliate leaf.[139] Such instances seem to afford an extreme degree of a
more common change, viz., the diminished size and contracted appearance
of the sepals and petals when affected with axillary prolification. They have
also a close relationship to such developments as we see in the wheat-ear
carnation, in certain species of the genus Mæsa and others, wherein the
calyx is repeated over and again, to the partial or complete suppression of
the other parts of the flower. All these cases may be in part explained by the
operation of the principle of compensation.
So far as the andrœcium is concerned, the stamens either remain unaltered,
or they are present in a more or less petal-like condition; but it far more
frequently happens that the stamens are entirely suppressed, the
adventitious bud supplying their place; thus was it in the Dianthus
represented in the adjoining woodcut, fig. 66, where the stamens were
entirely absent, and their places supplied by flower-bearing branches. This
Dianthus has the more interest from its similarity to the one described by
Goethe, Metam. der Pflanzen, cap. 16, sect. 105; but in that instance median
prolification also existed. For my specimens I am indebted to Mr. T. Moore.
Page 202
Fig. 66.—Flower of Dianthus sp., calyx removed; petals turned down so as to show the stalked
flower-buds springing from their axils.
The pistil, too, is necessarily subject to very grave alterations when affected
with this malformation. It is separated into its constituent carpels; and these
assume a leaf-like aspect, and are in the great majority of instances destitute
of ovules. Indeed, virescence or chloranthy is very intimately connected
with this aberration, as might have been anticipated, for if the parts of the
flower assume more or less of the condition of stem-leaves or bracts, it is
quite natural to expect that they will partake likewise of the attributes of
leaves, even at the expense of their own peculiar functions.
It occasionally happens that an adventitious bud arises from the axil of a
monocarpellary pistil. This takes place sometimes in Leguminosæ, and
seems to have been more frequently met with in Trifolium repens than in
other plants. The species named is, as is well known, particularly subject to
a reversion of the outer whorls of the flower to leaves, and even to a leaf-
flower-buds springing from their axils.
The pistil, too, is necessarily subject to very grave alterations when affected
with this malformation. It is separated into its constituent carpels; and these
assume a leaf-like aspect, and are in the great majority of instances destitute
of ovules. Indeed, virescence or chloranthy is very intimately connected
with this aberration, as might have been anticipated, for if the parts of the
flower assume more or less of the condition of stem-leaves or bracts, it is
quite natural to expect that they will partake likewise of the attributes of
leaves, even at the expense of their own peculiar functions.
It occasionally happens that an adventitious bud arises from the axil of a
monocarpellary pistil. This takes place sometimes in Leguminosæ, and
seems to have been more frequently met with in Trifolium repens than in
other plants. The species named is, as is well known, particularly subject to
a reversion of the outer whorls of the flower to leaves, and even to a leaf-
Page 203
like condition of the pistil. There are on record instances wherein a leaf-bud
has been placed in the axil of a more or less leaf-like carpel; while at other
times a second imperfect carpel has been met with in the axil of the first.
[140] I have myself seen numerous imperfectly developed cases of this kind.
It may be asked whether such cases are not more properly referable to
central prolification—whether the axis is not in such flowers terminated by
two, rather than by one carpel? It is, however, generally admitted by
morphologists that the solitary carpel of Leguminosæ is not terminal, but is
the sole existing member of a whorl of carpels, all the other members of
which are suppressed as a general rule, though exceptional instances of the
presence of two and even of five carpels have been described.[141]
Again, the adventitious bud or carpel is placed, not laterally to the primary
one, or opposite to it, on the same level, but slightly higher up—in fact, in
the axil of the primary carpellary leaf. Griffith figures and describes[142] an
instance of the kind in a species of Melilotus. The stalk of the ovary is
mentioned as having a sheathing base, bearing in its axil a prolongation of
the axis of inflorescence, in the form of a short spike with hairy bracts and
imperfect flowers, the latter having a well-formed calyx and rudimentary
petals and stamens. Griffith infers, from this specimen, that the legume is
not to be considered as a terminal leaf.
List of Genera in which Axillary Prolification has been observed.
Leaf-bud or Flower-bud or
Order Genus. From what organ.
Branch Inflorescence
Ranunculaceæ Clematis Flower-bud Sepals.
Caltha Ditto Ditto.
Aconitum Ditto.
Delphinium Ditto Sepals, carpels, &c.
Anemone! Ditto Involucre?
Nymphæaceæ Nymphæa! Fruit?
Nymphæa Flower Petal.
Cruciferæ *Brassica! Leaf-bud Flower-bud Sepals and petals.
has been placed in the axil of a more or less leaf-like carpel; while at other
times a second imperfect carpel has been met with in the axil of the first.
[140] I have myself seen numerous imperfectly developed cases of this kind.
It may be asked whether such cases are not more properly referable to
central prolification—whether the axis is not in such flowers terminated by
two, rather than by one carpel? It is, however, generally admitted by
morphologists that the solitary carpel of Leguminosæ is not terminal, but is
the sole existing member of a whorl of carpels, all the other members of
which are suppressed as a general rule, though exceptional instances of the
presence of two and even of five carpels have been described.[141]
Again, the adventitious bud or carpel is placed, not laterally to the primary
one, or opposite to it, on the same level, but slightly higher up—in fact, in
the axil of the primary carpellary leaf. Griffith figures and describes[142] an
instance of the kind in a species of Melilotus. The stalk of the ovary is
mentioned as having a sheathing base, bearing in its axil a prolongation of
the axis of inflorescence, in the form of a short spike with hairy bracts and
imperfect flowers, the latter having a well-formed calyx and rudimentary
petals and stamens. Griffith infers, from this specimen, that the legume is
not to be considered as a terminal leaf.
List of Genera in which Axillary Prolification has been observed.
Leaf-bud or Flower-bud or
Order Genus. From what organ.
Branch Inflorescence
Ranunculaceæ Clematis Flower-bud Sepals.
Caltha Ditto Ditto.
Aconitum Ditto.
Delphinium Ditto Sepals, carpels, &c.
Anemone! Ditto Involucre?
Nymphæaceæ Nymphæa! Fruit?
Nymphæa Flower Petal.
Cruciferæ *Brassica! Leaf-bud Flower-bud Sepals and petals.
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Brassica! Ditto Stamens.
Brassica! Ditto Ditto Pistil.
Cardamine! Ditto Sepals.
Matthiola! Ditto Sepals and petals.
Cheiranthus! Ditto Sepals.
Erysimum Ditto Sepals and pistils.
Lepidium! Ditto Petals and stamens.
Arabis Ditto Sepals.
Diplotaxis Flower, inflorescence Pistil, calyx and corolla.
Capsella
Capparidaceæ Cleome Flower-bud Sepals.
Resedaceæ *Reseda Ditto Ditto.
Caryophyllaceæ Arenaria Branch Ditto.
Agrostemma Leaf-bud Ditto.
*Lychnis Ditto
Stellaria Ditto
Silene Ditto
*Gypsophila Ditto Ditto Sepals and stamens.
*Dianthus! Ditto Ditto Sepals.
Dianthus! Ditto Inflorescence Petals and stamens.
Cucubalus Sepals
Saponaria! Sepals and petals.
Malvaceæ Alcea Flower-bud Stamen.
Aurantiaceæ Citrus! Ditto Ditto.
Rutaceæ Dictamnus! Ditto Pistil leafy.
Tropæolaceæ Tropæolum! Ditto Petals.
Celastraceæ Celastrus Ditto Sepals.
Leguminosæ *Melilotus! Inflorescence Sepals and petals.
Medicago Flower-bud Sepals.
Coronilla Ditto Ditto.
Second carpel axillary
Trifolium! Ditto Pistil.
to first
Melilotus! Ditto Ditto
Trifolium! Flower-bud Sepals and petals.
Rosaceæ Pyrus! Fruit? Fruit?
Cerasus! Flower-bud Petals and stamens.
Potentilla! Ditto Leafy carpels.
Cratægus! Ditto Petals.
Sepals, petals, stamens and
*Rosa! Ditto Ditto
pistil.
Myrtaceæ Lecythis Ditto Fruit?
Tetragoniaceæ Tetragonia? Ditto Ditto.
Fruit-like
Cactaceæ Opuntia! Tufts of spines.
branch
Pereskia Ditto Sepals?
Echinocactus Ditto Ditto.
Brassica! Ditto Ditto Pistil.
Cardamine! Ditto Sepals.
Matthiola! Ditto Sepals and petals.
Cheiranthus! Ditto Sepals.
Erysimum Ditto Sepals and pistils.
Lepidium! Ditto Petals and stamens.
Arabis Ditto Sepals.
Diplotaxis Flower, inflorescence Pistil, calyx and corolla.
Capsella
Capparidaceæ Cleome Flower-bud Sepals.
Resedaceæ *Reseda Ditto Ditto.
Caryophyllaceæ Arenaria Branch Ditto.
Agrostemma Leaf-bud Ditto.
*Lychnis Ditto
Stellaria Ditto
Silene Ditto
*Gypsophila Ditto Ditto Sepals and stamens.
*Dianthus! Ditto Ditto Sepals.
Dianthus! Ditto Inflorescence Petals and stamens.
Cucubalus Sepals
Saponaria! Sepals and petals.
Malvaceæ Alcea Flower-bud Stamen.
Aurantiaceæ Citrus! Ditto Ditto.
Rutaceæ Dictamnus! Ditto Pistil leafy.
Tropæolaceæ Tropæolum! Ditto Petals.
Celastraceæ Celastrus Ditto Sepals.
Leguminosæ *Melilotus! Inflorescence Sepals and petals.
Medicago Flower-bud Sepals.
Coronilla Ditto Ditto.
Second carpel axillary
Trifolium! Ditto Pistil.
to first
Melilotus! Ditto Ditto
Trifolium! Flower-bud Sepals and petals.
Rosaceæ Pyrus! Fruit? Fruit?
Cerasus! Flower-bud Petals and stamens.
Potentilla! Ditto Leafy carpels.
Cratægus! Ditto Petals.
Sepals, petals, stamens and
*Rosa! Ditto Ditto
pistil.
Myrtaceæ Lecythis Ditto Fruit?
Tetragoniaceæ Tetragonia? Ditto Ditto.
Fruit-like
Cactaceæ Opuntia! Tufts of spines.
branch
Pereskia Ditto Sepals?
Echinocactus Ditto Ditto.
Page 205
Philadelphaceæ Philadelphus Ditto Sepals.
Umbelliferæ *Athamanta Ditto Calyx.
*Daucus! Ditto Calyx and pistil.
Bupleurum Ditto Ditto ditto.
Torilis Ditto Ditto ditto.
Apium Flower-bud Calyx and pistil.
Pastinaca Ditto Ditto ditto.
Heracleum! Ditto Ditto ditto.
Angelica! Umbel Ditto ditto.
Campanulaceæ *Campanula! Branch Sepals.
Prismatocarpus Ditto Fruit Sepals, &c.
Gentianaceæ Gentiana! Flower-bud Sepals.
Convolvulaceæ *Convolvulus! Ditto Outer calyx.
Solanaceæ Solanum! Ditto Sepals.
Solanum Tubers Sepals and petals.
Scrophulariaceæ *Digitalis! Ditto Petals, &c.
Veronica Raceme Calyx.
Primulaceæ Anagallis! Branch Ditto Petals.
Primula Ditto Petals and carpels.
Polygonaceæ Rumex Ditto Sepals.
In place of stamens and
Santalaceæ Thesium Leaf-bud
pistils, both absent.
Euphorbiaceæ? Euphorbia? Ditto ? Outer bracts?
Orchidaceæ Orchis! Flower-bud Perianth.
Amaryllidaceæ Leucoium Ditto Ditto.
Iridaceæ Iris Ditto Pistil.
Liliaceæ Herreria Ditto Sepals.
Hyacinthus Flower and raceme Perianth.
Convallaria Flower-bud Ditto.
Allium Ditto Ditto.
Cyperaceæ Carex Inflorescence Utricle.
Umbelliferæ *Athamanta Ditto Calyx.
*Daucus! Ditto Calyx and pistil.
Bupleurum Ditto Ditto ditto.
Torilis Ditto Ditto ditto.
Apium Flower-bud Calyx and pistil.
Pastinaca Ditto Ditto ditto.
Heracleum! Ditto Ditto ditto.
Angelica! Umbel Ditto ditto.
Campanulaceæ *Campanula! Branch Sepals.
Prismatocarpus Ditto Fruit Sepals, &c.
Gentianaceæ Gentiana! Flower-bud Sepals.
Convolvulaceæ *Convolvulus! Ditto Outer calyx.
Solanaceæ Solanum! Ditto Sepals.
Solanum Tubers Sepals and petals.
Scrophulariaceæ *Digitalis! Ditto Petals, &c.
Veronica Raceme Calyx.
Primulaceæ Anagallis! Branch Ditto Petals.
Primula Ditto Petals and carpels.
Polygonaceæ Rumex Ditto Sepals.
In place of stamens and
Santalaceæ Thesium Leaf-bud
pistils, both absent.
Euphorbiaceæ? Euphorbia? Ditto ? Outer bracts?
Orchidaceæ Orchis! Flower-bud Perianth.
Amaryllidaceæ Leucoium Ditto Ditto.
Iridaceæ Iris Ditto Pistil.
Liliaceæ Herreria Ditto Sepals.
Hyacinthus Flower and raceme Perianth.
Convallaria Flower-bud Ditto.
Allium Ditto Ditto.
Cyperaceæ Carex Inflorescence Utricle.
Page 206
Fig. 67.—Proliferous Rose. Calyx leafy; petals normal, some reflexed; stamens and pistil absent; in
their places a branch with leaves and flowers.
their places a branch with leaves and flowers.
Page 207
Fig. 68.—Rose exhibiting median, axillary, lateral, floral, and leafy prolification in same flower.
Complicated prolification.—From what has been before stated it may be
seen that prolification of two or more kinds may coexist in the same flower.
Mixed leafy and floral prolification is not unfrequent in proliferous roses,
where a shoot is, as it were, prolonged through the centre of the original
flower and terminated by a second flower, or even by a cluster, as is well
shown in the accompanying figure (fig. 67). Median and axillary
prolification, also, not unfrequently coexist in the same flower; thus, in a
proliferous rose forwarded to me by Mr. W. Thomson (fig. 68), the
following changes were observed:—the swollen portion below the calyx,
the "hip," was entirely absent; the sepals were leaf-like in aspect, the petals
unaffected; above the petals the axis was prolonged for a short distance and
then bore a circlet of miniature, sessile roses, destitute, indeed, of calyx, but
provided with numerous petals, stamens, and pistils. Above these lateral
flowers, the prolonged axis bore a number of scales in many rows. The
Complicated prolification.—From what has been before stated it may be
seen that prolification of two or more kinds may coexist in the same flower.
Mixed leafy and floral prolification is not unfrequent in proliferous roses,
where a shoot is, as it were, prolonged through the centre of the original
flower and terminated by a second flower, or even by a cluster, as is well
shown in the accompanying figure (fig. 67). Median and axillary
prolification, also, not unfrequently coexist in the same flower; thus, in a
proliferous rose forwarded to me by Mr. W. Thomson (fig. 68), the
following changes were observed:—the swollen portion below the calyx,
the "hip," was entirely absent; the sepals were leaf-like in aspect, the petals
unaffected; above the petals the axis was prolonged for a short distance and
then bore a circlet of miniature, sessile roses, destitute, indeed, of calyx, but
provided with numerous petals, stamens, and pistils. Above these lateral
flowers, the prolonged axis bore a number of scales in many rows. The
Page 208
scales were in their turn surmounted by a whorl of five perfect leaves,
beyond which, again, the axis was prolonged into a leafy shoot terminated
by a flower bud, the whole constituting a remarkably complicated
admixture of elements belonging to the flower, the bud, the inflorescence,
and the leafshoot.[143]
Proliferous flowers of Orchids also occasionally present great complexity in
the arrangement of their parts. An instance of this kind was described by
myself from specimens furnished by Dr. Moore, of Glasnevin, in the
'Journal of the Linnean Society,' vol. ix, p. 349, tabs. x, xi, and from which
the following summary is extracted:
Fig. 69.—Proliferous Orchis. Diagram showing the arrangement of the several organs in the seven
outer circles of the flower. Each whorl is numbered, and the position of the axillary buds shown by
the small circles.
The primary flowers were composed of five distinct whorls, and of at least
two others less perfectly developed. These primary flowers did not give rise
beyond which, again, the axis was prolonged into a leafy shoot terminated
by a flower bud, the whole constituting a remarkably complicated
admixture of elements belonging to the flower, the bud, the inflorescence,
and the leafshoot.[143]
Proliferous flowers of Orchids also occasionally present great complexity in
the arrangement of their parts. An instance of this kind was described by
myself from specimens furnished by Dr. Moore, of Glasnevin, in the
'Journal of the Linnean Society,' vol. ix, p. 349, tabs. x, xi, and from which
the following summary is extracted:
Fig. 69.—Proliferous Orchis. Diagram showing the arrangement of the several organs in the seven
outer circles of the flower. Each whorl is numbered, and the position of the axillary buds shown by
the small circles.
The primary flowers were composed of five distinct whorls, and of at least
two others less perfectly developed. These primary flowers did not give rise
Page 209
to median formations, but they produced secondary buds in the axils of the
segments of the perianth. These latter buds were themselves the subject of
tertiary prolification of both kinds, median and axillary. The tertiary median
growths, like the primary flower, did not develop median buds, but only
lateral ones—quaternary axillary prolification.
The accompanying diagrams are intended to show the plan of arrangement
in these flowers. Fig. 69 shows the disposition of parts in the primary
flower and the situation of the axillary buds. Fig. 70 shows the primary
flower without any central prolongation, but giving off axillary buds, two of
which are shown in the diagram, 2, 2; these are, each of them, the subject of
both median, 3, 3, and axillary prolification, 4', 4'.
Fig. 70.—Diagram to explain the construction of the double-flowered Orchis.
1. The primary flower, with no median bud, the position of which, had it been present, is
shown by the dotted line.
2. Two axillary buds proceeding from 1, and themselves giving origin to
3, 3. Median buds, and 3', 3', axillary buds.
4' 4'. Axillary buds, proceeding from 3. No median bud is produced from 3; its situation,
had it been present, is indicated by the dotted line.
In Narcissus major a similar combination of both forms of prolification
exists, as described by Morren.[144]
On the general subject of Prolification in flowers, in addition to the
authorities already cited, the reader may refer to the following among many
segments of the perianth. These latter buds were themselves the subject of
tertiary prolification of both kinds, median and axillary. The tertiary median
growths, like the primary flower, did not develop median buds, but only
lateral ones—quaternary axillary prolification.
The accompanying diagrams are intended to show the plan of arrangement
in these flowers. Fig. 69 shows the disposition of parts in the primary
flower and the situation of the axillary buds. Fig. 70 shows the primary
flower without any central prolongation, but giving off axillary buds, two of
which are shown in the diagram, 2, 2; these are, each of them, the subject of
both median, 3, 3, and axillary prolification, 4', 4'.
Fig. 70.—Diagram to explain the construction of the double-flowered Orchis.
1. The primary flower, with no median bud, the position of which, had it been present, is
shown by the dotted line.
2. Two axillary buds proceeding from 1, and themselves giving origin to
3, 3. Median buds, and 3', 3', axillary buds.
4' 4'. Axillary buds, proceeding from 3. No median bud is produced from 3; its situation,
had it been present, is indicated by the dotted line.
In Narcissus major a similar combination of both forms of prolification
exists, as described by Morren.[144]
On the general subject of Prolification in flowers, in addition to the
authorities already cited, the reader may refer to the following among many
Page 210
others:
Linnæus, 'Prolepsis,' §§ vi et vii. Goethe, 'Versuch. Metamorph.,' cap. xv and xvi §§ 103–
106. Moquin-Tandon,' El. Ter. Veg.,' p. 362, &c. Engelmann, 'De Antholys.,' §§ 52–62, &c.
Cramer, 'Bildungsabweichungen,' &c. Orchidaceæ, Umbelliferæ, Compositæ, Leguminosæ,
Primulaceæ, Ranunculaceæ. Fleischer, 'Missbild. Cultur Gewachs.' Schlechtendal,
'Linnæa,' xv, p. 408, Rosa. 'Bot. Zeit.' vol. xx, 1862, p. 382, Cyclamen. 'Bot. Zeit.,' vol. xx,
p. 301, Asphodelus; et Lilium. Seringe, 'Bull. Bot.,' i, t. xi, f. 7, 8, Arabis, Diplotaxis. Clos,
'Mem. Acad. Toulouse,' 5th sér., 1862, Papaver. Wigand, 'Flora,' 1856, p. 716, Hypochæris;
et 'Bot. Untersuch.,' p. 19. Buchenau,' Flora,' 1857, p. 295, Reseda. Roeper, 'Bot. Zeit.,'
1852, p. 427, Orchis. Presl., 'Linnæa,' vi, p. 599, tab. ix, figs. 5–8, Sisymbrium, Vrolik.,
'Flora,' 1846, p. 97, t. i et ii, id. 1844, t. i, Digitalis. See also Schlechtendal, 'Bot. Zeit.,' vol.
ix, 1851, p. 579. Klinsmann, 'Linnæa,' x, p. 604, t. v, Hesperis. Fuckel, 'Flora,' 1848, p.
609. Melilotus. De Candolle, 'Organogr.,' i, 396, t. 33. Turpin, 'Atlas de Goethe,' p. 65, t. 5,
figs. 12, 13. Fenzl. 'Sitzungsbericht d. k. Akad. d. Wissensch. Wien.,' heft, iii, tabs. 3, 4,
Rosa. Kirschleger, 'Flora,' 1845, 613, Dianthus, Rosa. 'Institut.,' 1841, No. 413, p. 421,
Tragopogon. Baron de Melicoq., 'Ann. Sc. Nat.,' 3rd ser., vol. v. 1846, p. 61, Antirrhinum.
Reichenbach, 'Icon. Fl. Germ.,' tab. 100, Reseda—"monstrosa anticipatio Euphorbiacearum
et Capparidearum." Duhamel, 'Phys. Arbres.,' liv. iii, cap. 3, p. 303, pl. xii, f. 306, Rosa.
Caspary, 'Bull. Soc. Bot. Fr.,' vol. vi, 1859, p. 235, Rev. Bibl., Pyrus. Eichler, 'Flora,' 1865,
tab. ix, Cleome. Lindley, 'Elements of Botany,' p. 63, &c., Rosa, Epacris, Anagallis, Pyrus.
Irmish, 'Flora,' 1858, p. 38, Pyrus; and 'Bot. Zeit.,' xix, 1861, p. 342, Hyacinthus.
Duchartre, 'Bull. Soc. Bot. France,' 1861, p. 451, Rosa. Weber, 'Verhandl. Nat. Hist. Verein.
Rhein. Preuss., &c.' 1858 et 1860. Landrin, 'Mem. Soc. Sc. Nat. Seine et Oise,' 1866?[145]
Masters, 'Trans. Linn. Soc.,' vol. xxiii, p. 359, tab. 34 and p. 481, tab. 54.
Prolification of the embryo.—This term was applied by Moquin-Tandon
to a peculiar condition of the almond (Amygdalus), in which, indeed, it is
not of unfrequent occurrence. In these cases one almond encloses within its
cotyledons a second embryo, and this, again, in some instances, a third, the
little plants being thus packed like so many boxes one within the other. The
supplementary embryos are, in the ripe state at least, quite separate and
detached one from another. These cases differ from the ordinary instances
wherein there is an increased number of embryos in one seed in their
position. In the latter case, as often happens in the seeds of the orange, the
new products are placed by the side one of another.[146]
Linnæus, 'Prolepsis,' §§ vi et vii. Goethe, 'Versuch. Metamorph.,' cap. xv and xvi §§ 103–
106. Moquin-Tandon,' El. Ter. Veg.,' p. 362, &c. Engelmann, 'De Antholys.,' §§ 52–62, &c.
Cramer, 'Bildungsabweichungen,' &c. Orchidaceæ, Umbelliferæ, Compositæ, Leguminosæ,
Primulaceæ, Ranunculaceæ. Fleischer, 'Missbild. Cultur Gewachs.' Schlechtendal,
'Linnæa,' xv, p. 408, Rosa. 'Bot. Zeit.' vol. xx, 1862, p. 382, Cyclamen. 'Bot. Zeit.,' vol. xx,
p. 301, Asphodelus; et Lilium. Seringe, 'Bull. Bot.,' i, t. xi, f. 7, 8, Arabis, Diplotaxis. Clos,
'Mem. Acad. Toulouse,' 5th sér., 1862, Papaver. Wigand, 'Flora,' 1856, p. 716, Hypochæris;
et 'Bot. Untersuch.,' p. 19. Buchenau,' Flora,' 1857, p. 295, Reseda. Roeper, 'Bot. Zeit.,'
1852, p. 427, Orchis. Presl., 'Linnæa,' vi, p. 599, tab. ix, figs. 5–8, Sisymbrium, Vrolik.,
'Flora,' 1846, p. 97, t. i et ii, id. 1844, t. i, Digitalis. See also Schlechtendal, 'Bot. Zeit.,' vol.
ix, 1851, p. 579. Klinsmann, 'Linnæa,' x, p. 604, t. v, Hesperis. Fuckel, 'Flora,' 1848, p.
609. Melilotus. De Candolle, 'Organogr.,' i, 396, t. 33. Turpin, 'Atlas de Goethe,' p. 65, t. 5,
figs. 12, 13. Fenzl. 'Sitzungsbericht d. k. Akad. d. Wissensch. Wien.,' heft, iii, tabs. 3, 4,
Rosa. Kirschleger, 'Flora,' 1845, 613, Dianthus, Rosa. 'Institut.,' 1841, No. 413, p. 421,
Tragopogon. Baron de Melicoq., 'Ann. Sc. Nat.,' 3rd ser., vol. v. 1846, p. 61, Antirrhinum.
Reichenbach, 'Icon. Fl. Germ.,' tab. 100, Reseda—"monstrosa anticipatio Euphorbiacearum
et Capparidearum." Duhamel, 'Phys. Arbres.,' liv. iii, cap. 3, p. 303, pl. xii, f. 306, Rosa.
Caspary, 'Bull. Soc. Bot. Fr.,' vol. vi, 1859, p. 235, Rev. Bibl., Pyrus. Eichler, 'Flora,' 1865,
tab. ix, Cleome. Lindley, 'Elements of Botany,' p. 63, &c., Rosa, Epacris, Anagallis, Pyrus.
Irmish, 'Flora,' 1858, p. 38, Pyrus; and 'Bot. Zeit.,' xix, 1861, p. 342, Hyacinthus.
Duchartre, 'Bull. Soc. Bot. France,' 1861, p. 451, Rosa. Weber, 'Verhandl. Nat. Hist. Verein.
Rhein. Preuss., &c.' 1858 et 1860. Landrin, 'Mem. Soc. Sc. Nat. Seine et Oise,' 1866?[145]
Masters, 'Trans. Linn. Soc.,' vol. xxiii, p. 359, tab. 34 and p. 481, tab. 54.
Prolification of the embryo.—This term was applied by Moquin-Tandon
to a peculiar condition of the almond (Amygdalus), in which, indeed, it is
not of unfrequent occurrence. In these cases one almond encloses within its
cotyledons a second embryo, and this, again, in some instances, a third, the
little plants being thus packed like so many boxes one within the other. The
supplementary embryos are, in the ripe state at least, quite separate and
detached one from another. These cases differ from the ordinary instances
wherein there is an increased number of embryos in one seed in their
position. In the latter case, as often happens in the seeds of the orange, the
new products are placed by the side one of another.[146]
Page 211
For other cases of prolification or the adventitious formation of buds on
leaves, roots, &c., see under Heterotaxy.
FOOTNOTES:
[104] Linn., 'Prolepsis,' § vii; Goethe, 'Metamorph.,' §§ 96, 103, 106.
[105] "Diaphysis inflorescentiarum." Engelmann, 'De Anthol.,' § 85.
[106] 'Flora,' 1844, p. 565.
[107] 'Ann. Sc. Nat.,' ser. 3, vol. v, 1846, p. 64.
[108] 'Bot Zeit.,' vol. xx, p. 382.
[109] 'Miscel. Austriac. Bot.,' vol. i, Vindob, 1778, p. 133.
[110] "Umbellati dum prolificantur, augent umbellulam, ut ex umbellula simplici altera
exeat." 'Linn. Phil. Bot.,' § 124.
[111] 'En. Euphorb.,' p. 36.
[112] Meisner. 'Mon. Gen. Polygoni Prodrom.,' p. 20, tab. v, considers the bulbils of this
plant to be modifications of the pedicels of the flower.
[113] See A. Braun. 'Ann. Scienc. Nat.,' 4th series, 1860, vol. xiv, p. 13.
[114] "Prolificatio e latere ex calyci communi proles plurimos pedunculatos emittens, fit in
compositis aggregatis proprie dictis." 'Linn. Phil. Bot.,' § 124.
[115] 'Bot. Zeit.,' 1857, p. 873. See also 'Verhandl. Nat. Hist. Vereins. Preuss. Rheinl. u.
Westphal.,' 1854, t. ix.
[116] "Pannicula spicatim sparsa onusta innumera fœtura herbaceorum flosculorum
racematim cohærentium," 'Lobel. Stirp. Hist.,' p. 163. This is the "Besome Plantain, or
Plantain with spoky tufts," of Ray, 'Synopsis,' p. 314. Gerard's 'Herbal,' Ed. Johnson, p. 420.
Parkinson, 'Theat. Bot.,' p. 494. Baxter, 'Loudon. Mag. Nat. Hist.,' vol. ix. p. 204, and vol. iii,
p. 482. fig. 118.
[117] 'Flora.' 1856. p. 706.
[118] 'Flora of Berwick-on-Tweed,' vol. i. p. 38.
[119] 'Adansonia,' vol. iv. 1864, p. 150, tab. vii. 'Gard. Chron.,' November 19th, 1864.
leaves, roots, &c., see under Heterotaxy.
FOOTNOTES:
[104] Linn., 'Prolepsis,' § vii; Goethe, 'Metamorph.,' §§ 96, 103, 106.
[105] "Diaphysis inflorescentiarum." Engelmann, 'De Anthol.,' § 85.
[106] 'Flora,' 1844, p. 565.
[107] 'Ann. Sc. Nat.,' ser. 3, vol. v, 1846, p. 64.
[108] 'Bot Zeit.,' vol. xx, p. 382.
[109] 'Miscel. Austriac. Bot.,' vol. i, Vindob, 1778, p. 133.
[110] "Umbellati dum prolificantur, augent umbellulam, ut ex umbellula simplici altera
exeat." 'Linn. Phil. Bot.,' § 124.
[111] 'En. Euphorb.,' p. 36.
[112] Meisner. 'Mon. Gen. Polygoni Prodrom.,' p. 20, tab. v, considers the bulbils of this
plant to be modifications of the pedicels of the flower.
[113] See A. Braun. 'Ann. Scienc. Nat.,' 4th series, 1860, vol. xiv, p. 13.
[114] "Prolificatio e latere ex calyci communi proles plurimos pedunculatos emittens, fit in
compositis aggregatis proprie dictis." 'Linn. Phil. Bot.,' § 124.
[115] 'Bot. Zeit.,' 1857, p. 873. See also 'Verhandl. Nat. Hist. Vereins. Preuss. Rheinl. u.
Westphal.,' 1854, t. ix.
[116] "Pannicula spicatim sparsa onusta innumera fœtura herbaceorum flosculorum
racematim cohærentium," 'Lobel. Stirp. Hist.,' p. 163. This is the "Besome Plantain, or
Plantain with spoky tufts," of Ray, 'Synopsis,' p. 314. Gerard's 'Herbal,' Ed. Johnson, p. 420.
Parkinson, 'Theat. Bot.,' p. 494. Baxter, 'Loudon. Mag. Nat. Hist.,' vol. ix. p. 204, and vol. iii,
p. 482. fig. 118.
[117] 'Flora.' 1856. p. 706.
[118] 'Flora of Berwick-on-Tweed,' vol. i. p. 38.
[119] 'Adansonia,' vol. iv. 1864, p. 150, tab. vii. 'Gard. Chron.,' November 19th, 1864.
Page 212
[120] 'Ann. Sc. Nat.,' ser. 3, tom. ii, p. 290; and 'Adansonia,' iii, tab. iv; see also Bureau, in
'Bull. Soc. Bot. France,' x, p. 191.
[121] Baillon, 'Adansonia,' i, 286.
[122] See also figure in 'Hort. Eystett. Ic. Plant. Vern.,' fol. 15, fig. 1. Ranunculus asiaticus.
[123] Duchartre, 'Ann. des sc. nat.,' 3me série, vol. ii, 1844, p. 293.
[124] Reissek, 'Linnæa,' vol. xvii, 1843, p. 641, tab. xix.
[125] The tube of the calyx in these specimens was traversed by ten ribs, apparently
corresponding to the primary ridges of the normal fruit; these ribs were destitute of spines, and
the bristly secondary ridges were entirely absent. Those portions of the carpels which were
detached from the calyx had each three ribs, a central and two lateral ones, which appeared to
be continuous with the ribs of the calyx below,—although in the case of the calyx there were
ten, in the case of the carpels six ribs, three to each. This diversity in number is thus explained:
—A circle of vascular tissue ran round the interior of the calyx-tube, at its junction with the
limb, and at the point of insertion of the petals and stamens. The vascular circle seemed to be
formed from the confluence of the ten ribs from below. Of the five ribs in each half of the
calyx, the three central ones were joined together just at the point of confluence with the
vascular circle, above which they formed but a single rib—that traversing the centre of the
carpellary leaf; the two lateral ribs of each half of the calyx seemed to be continuous, above
the vascular rim, with the lateral ribs of the carpel; these lateral ribs were connected on either
side with the central one by short branches of communication. The disposition of the ten ribs
may be thus represented:—
1 1 1 1 1 1
3 2 3 2 3 3 2 3 2 3
1 1 1 1 1 1 1 1 1 1
The lower line of figures represents the calycine ribs, the middle row shows how each of these
ribs is divided at the vascular rim, and the uppermost row shows their distribution above the
rim. From this it will be seen that six of the calycine ribs divide into three branches, one
prolonged upwards as a lateral or median rib into the carpellary leaf, the other running
horizontally to join with similar branches sent out from the neighbouring rib; the four
intermediate calycine ribs divide into two branches only, which join the side branches of the
first mentioned, but have no direct upward prolongation into the carpel. The ten ridges are
placed opposite to the sepals and petals.
[126] 'Neue Denkschriften der allgemeine Schweizerischen Gesellschaft,' band 5. 1841. tab.
2.
'Bull. Soc. Bot. France,' x, p. 191.
[121] Baillon, 'Adansonia,' i, 286.
[122] See also figure in 'Hort. Eystett. Ic. Plant. Vern.,' fol. 15, fig. 1. Ranunculus asiaticus.
[123] Duchartre, 'Ann. des sc. nat.,' 3me série, vol. ii, 1844, p. 293.
[124] Reissek, 'Linnæa,' vol. xvii, 1843, p. 641, tab. xix.
[125] The tube of the calyx in these specimens was traversed by ten ribs, apparently
corresponding to the primary ridges of the normal fruit; these ribs were destitute of spines, and
the bristly secondary ridges were entirely absent. Those portions of the carpels which were
detached from the calyx had each three ribs, a central and two lateral ones, which appeared to
be continuous with the ribs of the calyx below,—although in the case of the calyx there were
ten, in the case of the carpels six ribs, three to each. This diversity in number is thus explained:
—A circle of vascular tissue ran round the interior of the calyx-tube, at its junction with the
limb, and at the point of insertion of the petals and stamens. The vascular circle seemed to be
formed from the confluence of the ten ribs from below. Of the five ribs in each half of the
calyx, the three central ones were joined together just at the point of confluence with the
vascular circle, above which they formed but a single rib—that traversing the centre of the
carpellary leaf; the two lateral ribs of each half of the calyx seemed to be continuous, above
the vascular rim, with the lateral ribs of the carpel; these lateral ribs were connected on either
side with the central one by short branches of communication. The disposition of the ten ribs
may be thus represented:—
1 1 1 1 1 1
3 2 3 2 3 3 2 3 2 3
1 1 1 1 1 1 1 1 1 1
The lower line of figures represents the calycine ribs, the middle row shows how each of these
ribs is divided at the vascular rim, and the uppermost row shows their distribution above the
rim. From this it will be seen that six of the calycine ribs divide into three branches, one
prolonged upwards as a lateral or median rib into the carpellary leaf, the other running
horizontally to join with similar branches sent out from the neighbouring rib; the four
intermediate calycine ribs divide into two branches only, which join the side branches of the
first mentioned, but have no direct upward prolongation into the carpel. The ten ridges are
placed opposite to the sepals and petals.
[126] 'Neue Denkschriften der allgemeine Schweizerischen Gesellschaft,' band 5. 1841. tab.
2.
Page 213
[127] Bell Salter, 'Gard. Chron.,' March 13th, 1847, and 'Ann. Nat. Hist.,' 1847, vol. xix, p.
471. &c.
[128] 'The Origin and Production of Proliferous Flowers, with the Culture at large for raising
Double Flowers from Single, and Proliferous from the Double.' By J. Hill, M.D. London,
1759.
[129] A. de Candolle, 'Neue Denkschriften,' op. cit., p. 9; also Unger as cited in 'Botanical
Gazette,' May, 1351. p. 70.
[130] Duchartre, op. cit.
[131] 'Ann. Sc. Nat.,' 1844, vol. i, p. 297.
[132] Maout, 'Leçons Elémentaires de Botanique,' vol. ii. p. 488; Ferrari. 'Hesperides.' pls.
271, 315, 405.
[133] Moquin-Tandon, loc. cit., p. 386, &c.; see also Trécul, in the 'Bull. Soc. Bot. France,'
tom. i, p, 307.
[134] 'Bull. Soc. Bot. Fr.,' 1863, vol. x, p. 461.
[135] 'Ann. Sc. Nat.,' 1835, p. 65. See also Le Maout, 'Leçons Element.,' vol. ii, p. 426.
[136] 'Proc. Hort. Soc.,' vol. i, p. 39, fig. 2.
[137] See also 'Nat. Hist. Review,' 1865, p. 377.
[138] 'Acad. Roy. Belg.,' April 11th. 1863.
[139] 'Bull. Soc. Bot. Fr.,' tom. iii, 1856, p. 479.
[140] 'Linnæa,' vol. xv, p. 266, c. ic. Caspary, 'Schriften d. Physik.-Oek. Gesell. zu
Königsberg,' bd. ii, p. 5, tab. iii, fig. 39, &c.
[141] Lindley, 'Veg. King.,' p. 545; also Clarke on the Position of Carpels, Linn. Soc.,'
December, 1850. 'Proc. Linn. Soc.,' ii, p. 105.
[142] 'Notulæ,' vol. i, Dicot. p. 127. 'Atlas,' pl. xliii.
[143] Moquin-Tandon gives the following references to cases of proliferous roses, but some I
have not been able to verify. 'Journ. des Sav.,' 22 Mai 1679. Hottinger, 'Ephem. Nat. Cur.,' dec.
3 ann. 9 et 10, p. 249. Marchant, 'Mem. Acad. Scienc. Paris.' 1707, p. 488. Preussius, 'Ephem.
Nat. Cur.,' cent. 7 et 8. App. p. 83. Schuster, 'Act. Acad. Nat. Cur.,' vol. vi, p. 185. Spadoni,
'Mem. Soc. Ital.,' t. v, p. 488. See also at the end of this section for numerous other references.
471. &c.
[128] 'The Origin and Production of Proliferous Flowers, with the Culture at large for raising
Double Flowers from Single, and Proliferous from the Double.' By J. Hill, M.D. London,
1759.
[129] A. de Candolle, 'Neue Denkschriften,' op. cit., p. 9; also Unger as cited in 'Botanical
Gazette,' May, 1351. p. 70.
[130] Duchartre, op. cit.
[131] 'Ann. Sc. Nat.,' 1844, vol. i, p. 297.
[132] Maout, 'Leçons Elémentaires de Botanique,' vol. ii. p. 488; Ferrari. 'Hesperides.' pls.
271, 315, 405.
[133] Moquin-Tandon, loc. cit., p. 386, &c.; see also Trécul, in the 'Bull. Soc. Bot. France,'
tom. i, p, 307.
[134] 'Bull. Soc. Bot. Fr.,' 1863, vol. x, p. 461.
[135] 'Ann. Sc. Nat.,' 1835, p. 65. See also Le Maout, 'Leçons Element.,' vol. ii, p. 426.
[136] 'Proc. Hort. Soc.,' vol. i, p. 39, fig. 2.
[137] See also 'Nat. Hist. Review,' 1865, p. 377.
[138] 'Acad. Roy. Belg.,' April 11th. 1863.
[139] 'Bull. Soc. Bot. Fr.,' tom. iii, 1856, p. 479.
[140] 'Linnæa,' vol. xv, p. 266, c. ic. Caspary, 'Schriften d. Physik.-Oek. Gesell. zu
Königsberg,' bd. ii, p. 5, tab. iii, fig. 39, &c.
[141] Lindley, 'Veg. King.,' p. 545; also Clarke on the Position of Carpels, Linn. Soc.,'
December, 1850. 'Proc. Linn. Soc.,' ii, p. 105.
[142] 'Notulæ,' vol. i, Dicot. p. 127. 'Atlas,' pl. xliii.
[143] Moquin-Tandon gives the following references to cases of proliferous roses, but some I
have not been able to verify. 'Journ. des Sav.,' 22 Mai 1679. Hottinger, 'Ephem. Nat. Cur.,' dec.
3 ann. 9 et 10, p. 249. Marchant, 'Mem. Acad. Scienc. Paris.' 1707, p. 488. Preussius, 'Ephem.
Nat. Cur.,' cent. 7 et 8. App. p. 83. Schuster, 'Act. Acad. Nat. Cur.,' vol. vi, p. 185. Spadoni,
'Mem. Soc. Ital.,' t. v, p. 488. See also at the end of this section for numerous other references.
Page 214
[144] 'Bull. Acad. Belg.,' t. xx, part ii, p. 271. See also Bellynck, 'Bull. Soc. Bot. Belg.,' t. vi,
ex. 'Bull. Soc. Bot. France,' t. xiv, 1867, Rev. Bibl., p. 241. Orchis ustulata.
[145] I have not been able to meet with this, but it is said to contain a paper on prolification,
with numerous bibliographical references.
[146] 'El. Ter. Veg.,' p. 364, Adnot.
CHAPTER III.
HETEROTAXY.
Under this category are here included a variety of deviations from the
ordinary arrangement and position of parts which cannot conveniently be
classed under the preceding or under other headings. The term heterotaxy is
intended to apply to the production of organs in situations where, under
usual circumstances, they would not be formed. It thus does not include
cases of substitution, where one part is replaced by another, or more or less
metamorphosed, nor cases of multiplication, nor of prolification which are
characterised not only by the production of members in unwonted
situations, but also in unwonted numbers. From the very nature of the
anomalies, and specially from the scanty knowledge we possess concerning
their mode of development, it is not possible to allocate them in all cases
correctly, and moreover many of them might as well be placed in one group
as in another.
Formation of adventitious roots.—This is of exceedingly common
occurrence in a vast number of plants, so much so that in most cases it
cannot be considered as in any way abnormal; there are, however, a few
instances where the formation of these organs may be considered to come
within the scope of teratology, or, at least, where their production is the
ex. 'Bull. Soc. Bot. France,' t. xiv, 1867, Rev. Bibl., p. 241. Orchis ustulata.
[145] I have not been able to meet with this, but it is said to contain a paper on prolification,
with numerous bibliographical references.
[146] 'El. Ter. Veg.,' p. 364, Adnot.
CHAPTER III.
HETEROTAXY.
Under this category are here included a variety of deviations from the
ordinary arrangement and position of parts which cannot conveniently be
classed under the preceding or under other headings. The term heterotaxy is
intended to apply to the production of organs in situations where, under
usual circumstances, they would not be formed. It thus does not include
cases of substitution, where one part is replaced by another, or more or less
metamorphosed, nor cases of multiplication, nor of prolification which are
characterised not only by the production of members in unwonted
situations, but also in unwonted numbers. From the very nature of the
anomalies, and specially from the scanty knowledge we possess concerning
their mode of development, it is not possible to allocate them in all cases
correctly, and moreover many of them might as well be placed in one group
as in another.
Formation of adventitious roots.—This is of exceedingly common
occurrence in a vast number of plants, so much so that in most cases it
cannot be considered as in any way abnormal; there are, however, a few
instances where the formation of these organs may be considered to come
within the scope of teratology, or, at least, where their production is the
Page 215
result of injury or of some unfavorable condition to which the plant is
exposed.
Thus the production of adventitious roots on the stem of the vine is
considered to be due to untoward circumstances impairing the proper action
of the ordinary subterranean roots. So, too, the formation of roots on the
upper portions of stems that are more or less decayed below, as in old
willows, is to be considered as an attempt to obtain fresh supplies through a
more vigorous and healthy channel.
A similar occurrence often arises as a consequence of some injury. Virgil
had this circumstance in view when he wrote
"Quin et, caudicibus sectis, mirabile dictu,
Truditur e sicco radix oleagina ligno."—'Georg.' Bk. ii.
I have seen many specimens of adventitious roots produced on the olive in
the way just mentioned.
In the 'Gardeners' Chronicle,' January 8th, 1853, p. 21, is described a
curious formation of roots in the fissure between two divisions of a
laburnum stem. In the same journal, January 1st, 1853, p. 4, Mr. Booth
mentions the case of a Cornish elm, the trunk of which was divided at the
top into two main divisions, and from the force of the wind or from some
other cause the stem was split down for several feet below the fork. Around
the edges of the fracture, layers of new bark were formed, from which
numerous roots issued, some measuring an inch in diameter and descending
into the cleft portion of the tree: similar instances must be familiar to all
observers.
It may happen that these roots sent down into the cavity of a decaying trunk
may, after a time, become completely concealed within it, by the gradual
exposed.
Thus the production of adventitious roots on the stem of the vine is
considered to be due to untoward circumstances impairing the proper action
of the ordinary subterranean roots. So, too, the formation of roots on the
upper portions of stems that are more or less decayed below, as in old
willows, is to be considered as an attempt to obtain fresh supplies through a
more vigorous and healthy channel.
A similar occurrence often arises as a consequence of some injury. Virgil
had this circumstance in view when he wrote
"Quin et, caudicibus sectis, mirabile dictu,
Truditur e sicco radix oleagina ligno."—'Georg.' Bk. ii.
I have seen many specimens of adventitious roots produced on the olive in
the way just mentioned.
In the 'Gardeners' Chronicle,' January 8th, 1853, p. 21, is described a
curious formation of roots in the fissure between two divisions of a
laburnum stem. In the same journal, January 1st, 1853, p. 4, Mr. Booth
mentions the case of a Cornish elm, the trunk of which was divided at the
top into two main divisions, and from the force of the wind or from some
other cause the stem was split down for several feet below the fork. Around
the edges of the fracture, layers of new bark were formed, from which
numerous roots issued, some measuring an inch in diameter and descending
into the cleft portion of the tree: similar instances must be familiar to all
observers.
It may happen that these roots sent down into the cavity of a decaying trunk
may, after a time, become completely concealed within it, by the gradual
Page 216
formation and extension of new wood over the orifice of the cavity formed
by the death and decay of the old wood. Such is presumed to be the
explanation of a specimen of this kind in the possession of the writer, and
taken from a cavity in an apparently solid block of rosewood; externally
there were no marks to indicate the existence of a central space, but when
the block was sawn up for the use of the cabinet-maker, this root-like
structure was found in the centre and attached to one end of the cavity.
The production of roots which ultimately serve as props to support the
branches, or as buttresses to compensate for the increasing weight of
branches and foliage, is also a familiar occurrence. The huge gnaurs and
burrs met with occasionally on some trees often produce great quantities,
not only of adventitious buds, but of roots also.
Fig. 71.—Production of adventitious roots from leaf stalk of celery.
by the death and decay of the old wood. Such is presumed to be the
explanation of a specimen of this kind in the possession of the writer, and
taken from a cavity in an apparently solid block of rosewood; externally
there were no marks to indicate the existence of a central space, but when
the block was sawn up for the use of the cabinet-maker, this root-like
structure was found in the centre and attached to one end of the cavity.
The production of roots which ultimately serve as props to support the
branches, or as buttresses to compensate for the increasing weight of
branches and foliage, is also a familiar occurrence. The huge gnaurs and
burrs met with occasionally on some trees often produce great quantities,
not only of adventitious buds, but of roots also.
Fig. 71.—Production of adventitious roots from leaf stalk of celery.
Page 217
The leaves, equally with the stems, have the power of emitting roots under
certain conditions, as when the leaves are in close contact with moist soil or
as the result of injury. This happens in some plants more readily than in
others—Bryophyllum calycinum is a well-known instance. Mr. Berkeley has
described the formation of roots from the fractured leaves of celery,[147] and
also in a cabbage where a snail "having gnawed a hole into the middle of a
leaf at its junction with the stem, a fascicle of roots was formed, bursting
through the tissue lining the cavity, and covered with abundant delicate
hairs after the fashion of ordinary radicles."
Fig. 72.—Germinating plant of mango, showing production of roots from one of the cotyledons
(from the Kew Museum).
The production of adventitious roots is not limited to the ordinary leaves of
the plant, but may be manifested on the cotyledons; thus Irmisch describes
cases of this kind in the cotyledons of Bunium creticum and Carum
certain conditions, as when the leaves are in close contact with moist soil or
as the result of injury. This happens in some plants more readily than in
others—Bryophyllum calycinum is a well-known instance. Mr. Berkeley has
described the formation of roots from the fractured leaves of celery,[147] and
also in a cabbage where a snail "having gnawed a hole into the middle of a
leaf at its junction with the stem, a fascicle of roots was formed, bursting
through the tissue lining the cavity, and covered with abundant delicate
hairs after the fashion of ordinary radicles."
Fig. 72.—Germinating plant of mango, showing production of roots from one of the cotyledons
(from the Kew Museum).
The production of adventitious roots is not limited to the ordinary leaves of
the plant, but may be manifested on the cotyledons; thus Irmisch describes
cases of this kind in the cotyledons of Bunium creticum and Carum
Page 218
Bulbocastanum.[148] I have figured and described an analogous case in the
cotyledons of the Mango (fig. 72).[149]
To this formation of adventitious roots the gardener owes the power he has
of propagating plants by cuttings, i.e., small portions of the stem with a bud
or buds attached, or in some cases from portions of the leaves, of the roots
themselves, or even of the fruit, as in the case of the cactus (Baillon). Care
also has to be exercised in grafting certain fruit trees not to allow the
grafted portion to be too close to the ground, else the scion throws out roots
into the soil, and the object of the cultivator is defeated.
Figs. 73 and 74 show formation of roots from leaves induced by the art of the gardener.
Layering is another garden operation dependent on the formation of these
organs, and advantage is also sometimes taken of this tendency of some
plants to produce roots when injured to reduce the dimensions of a plant
when getting too large for the house in which it is growing. By gradually
cotyledons of the Mango (fig. 72).[149]
To this formation of adventitious roots the gardener owes the power he has
of propagating plants by cuttings, i.e., small portions of the stem with a bud
or buds attached, or in some cases from portions of the leaves, of the roots
themselves, or even of the fruit, as in the case of the cactus (Baillon). Care
also has to be exercised in grafting certain fruit trees not to allow the
grafted portion to be too close to the ground, else the scion throws out roots
into the soil, and the object of the cultivator is defeated.
Figs. 73 and 74 show formation of roots from leaves induced by the art of the gardener.
Layering is another garden operation dependent on the formation of these
organs, and advantage is also sometimes taken of this tendency of some
plants to produce roots when injured to reduce the dimensions of a plant
when getting too large for the house in which it is growing. By gradually
Page 219
inducing the production of new roots from the central or upper portions of
the stem, it becomes possible, after a time, to sever the connection between
the original roots and the upper portion of the trunk, and thus secure a
shortened plant.
On the subject of adventitious roots, &c., reference may be made to Trécul, 'Ann. Sc. Nat.,'
1846, t. v, p. 340, et vi, p. 303. Duchartre, 'Elements de Botanique,' p. 219. Lindley,
'Theory and Practice of Horticulture.' Thomson's 'Gardener's Assistant,' pp. 374, et seq.;
and any of the ordinary botanical text-books.
Formation of adventitious buds on roots.—One of the characteristics by
which roots are distinguished from stems in a general way consists in the
absence of buds; but, as is well known, they may be formed on the roots
under certain circumstances, and in certain plants, e.g., Pyrus Japonica,
Anemone Japonica, &c. What are termed suckers, owe their origin to buds
formed in this situation.
If roots be exposed or injured, they will frequently emit buds. The well-
known experiment of Duhamel, in which a willow was placed with the
branches in the soil and the roots in the air, and emitted new buds from the
latter and new roots from the former, depended on this production of
adventitious organs of either kind.
Gardeners often avail themselves of the power that the roots have of
producing buds to propagate plants by cuttings of the roots, but in many of
these cases the organ "parted" or cut is really an underground stem and not
a true root.
M. Claas Mulder has figured and described a case in the turnip-radish of the
unusual formation of a leafy shoot from the root, apparently after injury.[150]
From the figure it appears as if the lower portion of the root had been split
almost to the extremity, while the upper portion seems to have a central
cavity passing through it. From the angle, formed by the split segments
the stem, it becomes possible, after a time, to sever the connection between
the original roots and the upper portion of the trunk, and thus secure a
shortened plant.
On the subject of adventitious roots, &c., reference may be made to Trécul, 'Ann. Sc. Nat.,'
1846, t. v, p. 340, et vi, p. 303. Duchartre, 'Elements de Botanique,' p. 219. Lindley,
'Theory and Practice of Horticulture.' Thomson's 'Gardener's Assistant,' pp. 374, et seq.;
and any of the ordinary botanical text-books.
Formation of adventitious buds on roots.—One of the characteristics by
which roots are distinguished from stems in a general way consists in the
absence of buds; but, as is well known, they may be formed on the roots
under certain circumstances, and in certain plants, e.g., Pyrus Japonica,
Anemone Japonica, &c. What are termed suckers, owe their origin to buds
formed in this situation.
If roots be exposed or injured, they will frequently emit buds. The well-
known experiment of Duhamel, in which a willow was placed with the
branches in the soil and the roots in the air, and emitted new buds from the
latter and new roots from the former, depended on this production of
adventitious organs of either kind.
Gardeners often avail themselves of the power that the roots have of
producing buds to propagate plants by cuttings of the roots, but in many of
these cases the organ "parted" or cut is really an underground stem and not
a true root.
M. Claas Mulder has figured and described a case in the turnip-radish of the
unusual formation of a leafy shoot from the root, apparently after injury.[150]
From the figure it appears as if the lower portion of the root had been split
almost to the extremity, while the upper portion seems to have a central
cavity passing through it. From the angle, formed by the split segments
Page 220
below, proceeds a tuft of leaves, some of which appear to have traversed the
central cavity and to have emerged from the summit, mingling with the
other leaves in that situation. The production of a flower-bud has even been
noticed on the root of a species of Impatiens.
Formation of shoots beneath the cotyledons.—The tigellar or axial
portion of the embryo plant, as contrasted with the radicle proper, is very
variously developed in different cases; sometimes it is a mere "collar"
bearing the cotyledons, while at other times it is of considerable size.
Generally it does not give origin to shoots or leaves other than the seed-
leaves, but occasionally shoots may be seen projecting from it below the
level of the cotyledons. This happens frequently in seedling plants of
Anagallis arvensis, Euphorbia peplus, and other species, Linaria vulgaris,
some Umbelliferæ, &c.[151]
Adventitious formation of leaves.—The term phyllomania has been
vaguely applied both to the production of an unwonted number of leaves
and to their development in unusual situations. Under the present heading
the latter class of cases are alone included. The extraordinary tendency in
some Begonias to develop leaves or leafy excrescences from their surfaces
is elsewhere alluded to, and is, in reality, a species of hypertrophy or over-
luxuriant growth.
In some flowers where the inferior ovary is supposed to be, in part at least,
formed by a dilatation of the top of the flower-stalk, leaves have been met
with proceeding from the surface of the ovary or fruit, as in Cratægus
tanacetifolia, roses, pears, gooseberries, &c. In a specimen of Nymphæa
alba I have met with scale-like leaves projecting from the surface of the
fruit (or torus?), and which did not appear to be metamorphosed stamens or
styles (fig. 76).
central cavity and to have emerged from the summit, mingling with the
other leaves in that situation. The production of a flower-bud has even been
noticed on the root of a species of Impatiens.
Formation of shoots beneath the cotyledons.—The tigellar or axial
portion of the embryo plant, as contrasted with the radicle proper, is very
variously developed in different cases; sometimes it is a mere "collar"
bearing the cotyledons, while at other times it is of considerable size.
Generally it does not give origin to shoots or leaves other than the seed-
leaves, but occasionally shoots may be seen projecting from it below the
level of the cotyledons. This happens frequently in seedling plants of
Anagallis arvensis, Euphorbia peplus, and other species, Linaria vulgaris,
some Umbelliferæ, &c.[151]
Adventitious formation of leaves.—The term phyllomania has been
vaguely applied both to the production of an unwonted number of leaves
and to their development in unusual situations. Under the present heading
the latter class of cases are alone included. The extraordinary tendency in
some Begonias to develop leaves or leafy excrescences from their surfaces
is elsewhere alluded to, and is, in reality, a species of hypertrophy or over-
luxuriant growth.
In some flowers where the inferior ovary is supposed to be, in part at least,
formed by a dilatation of the top of the flower-stalk, leaves have been met
with proceeding from the surface of the ovary or fruit, as in Cratægus
tanacetifolia, roses, pears, gooseberries, &c. In a specimen of Nymphæa
alba I have met with scale-like leaves projecting from the surface of the
fruit (or torus?), and which did not appear to be metamorphosed stamens or
styles (fig. 76).
Page 221
Fig. 75.—Leaf proceeding from hip of the Rose.
Page 222
Fig. 76.—Leaves proceeding from the ovary of Nymphæa.
For other illustrations of increased leaf-formation, see Multiplication of
foliar organs.
Fig. 77.—Leontodon. Scape with two leaves; the bracts of the involucre are also leafy.
Production of leaves on a usually leafless inflorescence.—The
development of the bracts of an inflorescence to such an extent that they
resemble ordinary leaves is elsewhere alluded to as of common occurrence.
It happens far less frequently that leaves are developed on an inflorescence
usually destitute of them, without any metamorphosis or substitution, and
without any formation of adventitious buds, such as happens in
prolification. Such a partial change from a floriferous to a foliiferous branch
may be seen in a specimen of Sambucus nigra in the Smithian herbarium in
the Linnean Society, where the ultimate branches of the cyme bear small
For other illustrations of increased leaf-formation, see Multiplication of
foliar organs.
Fig. 77.—Leontodon. Scape with two leaves; the bracts of the involucre are also leafy.
Production of leaves on a usually leafless inflorescence.—The
development of the bracts of an inflorescence to such an extent that they
resemble ordinary leaves is elsewhere alluded to as of common occurrence.
It happens far less frequently that leaves are developed on an inflorescence
usually destitute of them, without any metamorphosis or substitution, and
without any formation of adventitious buds, such as happens in
prolification. Such a partial change from a floriferous to a foliiferous branch
may be seen in a specimen of Sambucus nigra in the Smithian herbarium in
the Linnean Society, where the ultimate branches of the cyme bear small
Page 223
leaves. My attention was directed to this specimen by the Rev. W.
Newbould.
Jacquin figures an analogous case in Sempervivum sediforme,[152] in which
the branches of the inflorescence were prolonged into leafy shoots.
Sometimes from the side of a flower-stalk or scape, which usually does not
bear leaves, those organs are produced. The common dandelion,
Taraxacum, sometimes offers an illustration of this, and also the daisy
(Bellis).[153] In a specimen of fasciated cowslip given me by Mr. Edgeworth
there was a similar formation of leaves on the flattened stalk.
Production of leaves or scales in place of flower-buds.—The position of
the leaf and of the flower-buds respectively is, in most plants, well defined,
but occasionally it happens that the former is formed where, under ordinary
circumstances, the latter organ should be. This may happen without the
formation of any transitional organs between the two, and without actual
increase in the number of the buds. Where there is evidently a passage from
leaf-bud to flower-bud, or vice versâ, the case would be one of
metamorphy. If the number of buds be augmented, or they be mixed with
the flower-buds, then it would be referable to leafy prolification of the
inflorescence. There remains a class of cases wherein there is a complete
substitution of one structure for the other, it may be without the slightest
indication of transition between the two, and without any admixture of leaf-
buds among flower-buds, or any absolute increase in the number of organs,
as in Prolification. Such a case is represented in fig. 78, which shows a
portion of the stem of a species of Valeriana, bearing at the summit, not an
inflorescence, but a tuft of leaves without the slightest indication of flowers.
Drs. Hooker and Thomson relate that in Northern India the flowers of
Anemone rivularis are very generally absent, and their place supplied by
Newbould.
Jacquin figures an analogous case in Sempervivum sediforme,[152] in which
the branches of the inflorescence were prolonged into leafy shoots.
Sometimes from the side of a flower-stalk or scape, which usually does not
bear leaves, those organs are produced. The common dandelion,
Taraxacum, sometimes offers an illustration of this, and also the daisy
(Bellis).[153] In a specimen of fasciated cowslip given me by Mr. Edgeworth
there was a similar formation of leaves on the flattened stalk.
Production of leaves or scales in place of flower-buds.—The position of
the leaf and of the flower-buds respectively is, in most plants, well defined,
but occasionally it happens that the former is formed where, under ordinary
circumstances, the latter organ should be. This may happen without the
formation of any transitional organs between the two, and without actual
increase in the number of the buds. Where there is evidently a passage from
leaf-bud to flower-bud, or vice versâ, the case would be one of
metamorphy. If the number of buds be augmented, or they be mixed with
the flower-buds, then it would be referable to leafy prolification of the
inflorescence. There remains a class of cases wherein there is a complete
substitution of one structure for the other, it may be without the slightest
indication of transition between the two, and without any admixture of leaf-
buds among flower-buds, or any absolute increase in the number of organs,
as in Prolification. Such a case is represented in fig. 78, which shows a
portion of the stem of a species of Valeriana, bearing at the summit, not an
inflorescence, but a tuft of leaves without the slightest indication of flowers.
Drs. Hooker and Thomson relate that in Northern India the flowers of
Anemone rivularis are very generally absent, and their place supplied by
Page 224
tufts or umbels of leaves.[154] In the collection of the late Mr. N. B. Ward
was a specimen of lupin in which the flowers were all absent, and their
place supplied by tufts of leaves.
Fig. 78.—Tuft of leaves replacing the inflorescence in a species of Valeriana.
A similar appearance has been noticed in Compositæ, and I owe to the
kindness of Professor Oliver the communication of a specimen of a species
of Bidens from Peru, in which the capitula, instead of consisting of florets,
as usual, contained tufts of linear ciliolated bracts within the involucre,
without a trace of flowers. In the eleventh volume of the 'Linnæa,' 1837, p.
301, Von Cesati figures and describes an analogous case in Carduus
crispus. The same author[1] records a similar instance in the umbel of
Seseli coloratum, where the place of the flowers was occupied by stalked
tufts of leaves. In the 'Gardeners' Chronicle,' October 6th, 1860, p. 894, is
mentioned an instance where the blossoms of the pea were entirely absent,
was a specimen of lupin in which the flowers were all absent, and their
place supplied by tufts of leaves.
Fig. 78.—Tuft of leaves replacing the inflorescence in a species of Valeriana.
A similar appearance has been noticed in Compositæ, and I owe to the
kindness of Professor Oliver the communication of a specimen of a species
of Bidens from Peru, in which the capitula, instead of consisting of florets,
as usual, contained tufts of linear ciliolated bracts within the involucre,
without a trace of flowers. In the eleventh volume of the 'Linnæa,' 1837, p.
301, Von Cesati figures and describes an analogous case in Carduus
crispus. The same author[1] records a similar instance in the umbel of
Seseli coloratum, where the place of the flowers was occupied by stalked
tufts of leaves. In the 'Gardeners' Chronicle,' October 6th, 1860, p. 894, is
mentioned an instance where the blossoms of the pea were entirely absent,
Page 225
and their place supplied by accumulations of small, ovate, green scales, thus
presenting an appearance similar to that brought about by the inordinate
multiplication of the sepals in the "wheat-ear carnation," and in the Sweet
William, and not unlike the condition met with in Bryophyllum proliferum.
In Digitalis purpurea a similar anomaly is sometimes met with.
In the apple I have observed leafy shoots bearing terminal tufts of leaves
where the flower should have been, so that what, under ordinary
circumstances would be a corymb of flowers, is here represented by a series
of tufts of leaves. In the cultivated azaleas also, leafy shoots occupying the
position of the flower may occasionally be met with.
In Bouchea hyderabadensis I have seen the inflorescence more than usually
branched and covered with little tufts of bracts, without a trace of true
flower. A similar condition seems not infrequent in Gentiana Amarella, as I
have not only met with the plant myself in this condition, but have been
favoured with specimens by Mr. Pamplin, Mr. Darwin, and others. In
Phyteuma spicatum an analogous appearance has been recorded.
Among Griffith's collections from Affghanistan is a species of willow
(Salix) in which the inflorescence replaced by a much branched panicle,
bearing a quantity of minute bracts, in the axils of which nestle numerous
small buds. In another specimen the inflorescence preserves its usual
catkin-like shape, but the flowers are replaced by little tufts of leaves. M.
Germain de Saint Pierre mentions a case wherein the flowers of Alisma
parnassifolia were completely replaced by leaf-buds.[155]
presenting an appearance similar to that brought about by the inordinate
multiplication of the sepals in the "wheat-ear carnation," and in the Sweet
William, and not unlike the condition met with in Bryophyllum proliferum.
In Digitalis purpurea a similar anomaly is sometimes met with.
In the apple I have observed leafy shoots bearing terminal tufts of leaves
where the flower should have been, so that what, under ordinary
circumstances would be a corymb of flowers, is here represented by a series
of tufts of leaves. In the cultivated azaleas also, leafy shoots occupying the
position of the flower may occasionally be met with.
In Bouchea hyderabadensis I have seen the inflorescence more than usually
branched and covered with little tufts of bracts, without a trace of true
flower. A similar condition seems not infrequent in Gentiana Amarella, as I
have not only met with the plant myself in this condition, but have been
favoured with specimens by Mr. Pamplin, Mr. Darwin, and others. In
Phyteuma spicatum an analogous appearance has been recorded.
Among Griffith's collections from Affghanistan is a species of willow
(Salix) in which the inflorescence replaced by a much branched panicle,
bearing a quantity of minute bracts, in the axils of which nestle numerous
small buds. In another specimen the inflorescence preserves its usual
catkin-like shape, but the flowers are replaced by little tufts of leaves. M.
Germain de Saint Pierre mentions a case wherein the flowers of Alisma
parnassifolia were completely replaced by leaf-buds.[155]
Page 226
Fig. 79.—Spikelets of Willedenovia, composed entirely of scales to the exclusion of flowers.
Page 227
Fig. 80.—Rose Willow, Salix, sp.
Here, also, may be mentioned the curious aggregations of scales which
occur in some grasses, in Restiaceæ, Juncaceæ, and other orders, in which
the inflorescence is made up of collections of scales or bracts with no trace
of floral structure. Fig. 79 shows this in a species of Willdenovia, and a very
good example is figured in a bamboo, Pseudostachyum polymorphum, by
General Munro.[156]
"Rose willows" (fig. 80) owe their peculiar appearance to a similar cause,
the scales of the catkin being here replaced by closely crowded leaves.
These aggregations of scales or leaves are not confined to the inflorescence,
but may be found in other parts of the plant, and may be frequently met
with in the willow, birch, oak, &c., generally as the result of insect
puncture. On the other hand, the production of leaves or leaf-buds in place
of flowers is, as is well known, generally the consequence of an excess of
nutrition, and of the continuance rather than of the arrest of vegetative
development.[157] It has even been asserted that a flower-bud may be
transformed into a leaf-bud by removing the pistil at a very early stage of
development, but this statement requires further confirmation.[158]
Viviparous plants.—The spikelets of certain grasses are frequently found
with some of their constituent parts completely replaced by leaves, like
those of the stem, while the true flowers are usually entirely absent. A
shoot, in fact, is formed in place of a series of flowers. In these cases it
generally happens that the outermost glumes are changed, sometimes,
however, even the outer and inner paleæ are wholly unchanged, while there
is no trace of squamulæ or of stamens and pistils within them, but in their
place is a small shoot with miniature leaves arranged in the ordinary
manner.
Here, also, may be mentioned the curious aggregations of scales which
occur in some grasses, in Restiaceæ, Juncaceæ, and other orders, in which
the inflorescence is made up of collections of scales or bracts with no trace
of floral structure. Fig. 79 shows this in a species of Willdenovia, and a very
good example is figured in a bamboo, Pseudostachyum polymorphum, by
General Munro.[156]
"Rose willows" (fig. 80) owe their peculiar appearance to a similar cause,
the scales of the catkin being here replaced by closely crowded leaves.
These aggregations of scales or leaves are not confined to the inflorescence,
but may be found in other parts of the plant, and may be frequently met
with in the willow, birch, oak, &c., generally as the result of insect
puncture. On the other hand, the production of leaves or leaf-buds in place
of flowers is, as is well known, generally the consequence of an excess of
nutrition, and of the continuance rather than of the arrest of vegetative
development.[157] It has even been asserted that a flower-bud may be
transformed into a leaf-bud by removing the pistil at a very early stage of
development, but this statement requires further confirmation.[158]
Viviparous plants.—The spikelets of certain grasses are frequently found
with some of their constituent parts completely replaced by leaves, like
those of the stem, while the true flowers are usually entirely absent. A
shoot, in fact, is formed in place of a series of flowers. In these cases it
generally happens that the outermost glumes are changed, sometimes,
however, even the outer and inner paleæ are wholly unchanged, while there
is no trace of squamulæ or of stamens and pistils within them, but in their
place is a small shoot with miniature leaves arranged in the ordinary
manner.
Page 228
The grasses most commonly affected in this manner are Dactylis
glomerata!, Poa bulbosa!, Poa annua!, P. trivialis!, pratensis!, alpina!,
angustifolia, and laxa, Cynosurus cristatus, Festuca nemoralis, F. ovina!,
Glyceria fluitans!, Gl. aquatica, Aira alpina!, cæspitosa!, Phleum
phalaroides, Lolium perenne!, Alopecurus pratensis!, Agrostis alba, Holcus
mollis!
Fig. 81.—Portion of panicle of Aira vivipara and separate floret.
From an examination of the structure of viviparous grasses Von Mohl was
led to the conclusion that the lower palea is to be considered as a bract, and
not a perianthial leaf, because the base of the palea surrounds the stem or
axis of the spikelet entirely, and both its margins cohere towards its lower
extremity.[159]
A similar condition occurs not infrequently in Polygonum viviparum, and in
Juncaceæ, Cyperaceæ, &c.
glomerata!, Poa bulbosa!, Poa annua!, P. trivialis!, pratensis!, alpina!,
angustifolia, and laxa, Cynosurus cristatus, Festuca nemoralis, F. ovina!,
Glyceria fluitans!, Gl. aquatica, Aira alpina!, cæspitosa!, Phleum
phalaroides, Lolium perenne!, Alopecurus pratensis!, Agrostis alba, Holcus
mollis!
Fig. 81.—Portion of panicle of Aira vivipara and separate floret.
From an examination of the structure of viviparous grasses Von Mohl was
led to the conclusion that the lower palea is to be considered as a bract, and
not a perianthial leaf, because the base of the palea surrounds the stem or
axis of the spikelet entirely, and both its margins cohere towards its lower
extremity.[159]
A similar condition occurs not infrequently in Polygonum viviparum, and in
Juncaceæ, Cyperaceæ, &c.
Page 229
In the genus Allium an analogous formation of little buds or bulbils takes
place in lieu of flowers; this is specially the case with A. vineale, the
flowers of which are rarely seen.
Other illustrations of a similar character, where the adventitious leaf-buds
are mixed in amongst the flower-buds, are cited under the head of
Prolification of the Inflorescence.
Formation of buds on leaves.—The formation of little bulbs upon the
surfaces or edges of leaves, forming what are called viviparous leaves, has
long been familiar to botanists amongst Alliums. Professor Alexander
Braun,[160] who has paid much attention to this subject, divides cases of this
kind according to the position of the buds; thus, for instance, they are
sometimes formed upon the upper portion of the leaf or petiole, as in many
ferns, in Nymphæa guineensis, some Arads, &c. The same condition has
been met with as a teratological occurrence in the leaves of Cardamine
pratensis, Hyacinthus Pouzolzii, Drosera intermedia,[161] Arabis pumila,
Chelidonium majus, Chirita sinensis,[162] Episcia bicolor,[163] Zamia, &c.
[164] Many species of Begonia possess the power of emitting buds from the
petioles and veins of the leaf; the little ramenta or scales which so
plentifully beset the surface of some of these plants likewise, in some
instances, pass gradually into leaves. B. phyllomaniaca, Mart., is the species
best known as manifesting this tendency, but others have it also.[165]
Buds are also very often formed upon the margins of the leaf, the best
known instance of which occurs in Bryophyllum calycinum; Weinmann[166]
figures an instance of this kind in Alchemilla minima, or they may occur
upon the lower surface of the leaf, as in Ornithogalum scilloides and longe-
bracteatum. M. Duchartre[167] mentions a case in the tomato in which the
leaves gave origin to small leaf-bearing branches, which, of course, must
place in lieu of flowers; this is specially the case with A. vineale, the
flowers of which are rarely seen.
Other illustrations of a similar character, where the adventitious leaf-buds
are mixed in amongst the flower-buds, are cited under the head of
Prolification of the Inflorescence.
Formation of buds on leaves.—The formation of little bulbs upon the
surfaces or edges of leaves, forming what are called viviparous leaves, has
long been familiar to botanists amongst Alliums. Professor Alexander
Braun,[160] who has paid much attention to this subject, divides cases of this
kind according to the position of the buds; thus, for instance, they are
sometimes formed upon the upper portion of the leaf or petiole, as in many
ferns, in Nymphæa guineensis, some Arads, &c. The same condition has
been met with as a teratological occurrence in the leaves of Cardamine
pratensis, Hyacinthus Pouzolzii, Drosera intermedia,[161] Arabis pumila,
Chelidonium majus, Chirita sinensis,[162] Episcia bicolor,[163] Zamia, &c.
[164] Many species of Begonia possess the power of emitting buds from the
petioles and veins of the leaf; the little ramenta or scales which so
plentifully beset the surface of some of these plants likewise, in some
instances, pass gradually into leaves. B. phyllomaniaca, Mart., is the species
best known as manifesting this tendency, but others have it also.[165]
Buds are also very often formed upon the margins of the leaf, the best
known instance of which occurs in Bryophyllum calycinum; Weinmann[166]
figures an instance of this kind in Alchemilla minima, or they may occur
upon the lower surface of the leaf, as in Ornithogalum scilloides and longe-
bracteatum. M. Duchartre[167] mentions a case in the tomato in which the
leaves gave origin to small leaf-bearing branches, which, of course, must
Page 230
have originated from buds, just in the same way as in the Drosera before
mentioned.
Fig. 82.—Formation of shoot on leaf of Episcia bicolor.
Gardeners occasionally avail themselves of this formation of buds from
leaves to propagate plants, e.g. Hoya, Gesnera, Gloxinia, &c.
Formation of buds in the pith.—This is said to be a normal condition in
the curious Stangeria paradoxa,[168] and Mr. Berkeley records an instance
of this in sea-kale[169] (fig. 83) where the crown had been injured, and buds
were seen sprouting from its centre.
mentioned.
Fig. 82.—Formation of shoot on leaf of Episcia bicolor.
Gardeners occasionally avail themselves of this formation of buds from
leaves to propagate plants, e.g. Hoya, Gesnera, Gloxinia, &c.
Formation of buds in the pith.—This is said to be a normal condition in
the curious Stangeria paradoxa,[168] and Mr. Berkeley records an instance
of this in sea-kale[169] (fig. 83) where the crown had been injured, and buds
were seen sprouting from its centre.
Page 231
Fig. 83.—Adventitious buds in sea kale.
Page 232
Fig. 84.—Hyacinth bulb cut across to induce the formation of new bulbs.
Fig. 85.—Showing the formation of new bulbs on the cut edges of an old hyacinth bulb.
It will be remarked that the adventitious production of buds, like that of
roots, is very often consequent on decay or injury. The Dutch bulb-growers
have availed themselves of this latter circumstance in the propagation of
hyacinths. Mr. Fortune, who published some articles on this subject in the
'Gardener's Chronicle,'[170] describes two special modes as adopted by these
skilful horticulturists—the one to make two or three deep cuts at the base of
the bulb, destroying the nascent flower-stalk when, after a time, small bulbs
are formed along the edges of the cut surfaces (figs. 84, 85). The other
method is effected by scooping out the interior of the base of the bulb, thus
leaving exposed the cut ends of the sheathing leaves arranged
concentrically; along these lines the new bulbs are, after some time, formed
in great numbers (fig. 86).
Fig. 85.—Showing the formation of new bulbs on the cut edges of an old hyacinth bulb.
It will be remarked that the adventitious production of buds, like that of
roots, is very often consequent on decay or injury. The Dutch bulb-growers
have availed themselves of this latter circumstance in the propagation of
hyacinths. Mr. Fortune, who published some articles on this subject in the
'Gardener's Chronicle,'[170] describes two special modes as adopted by these
skilful horticulturists—the one to make two or three deep cuts at the base of
the bulb, destroying the nascent flower-stalk when, after a time, small bulbs
are formed along the edges of the cut surfaces (figs. 84, 85). The other
method is effected by scooping out the interior of the base of the bulb, thus
leaving exposed the cut ends of the sheathing leaves arranged
concentrically; along these lines the new bulbs are, after some time, formed
in great numbers (fig. 86).
Page 233
Fig. 86.—Showing the production of small bulbs on the inner surface of the scooped-out bulb of
hyacinth.
For the formation of supernumerary leaves on the surface of the normal
one, see Multiplication and Hypertrophy.
Production of gemmæ in place of spores.—An instance of this is recorded
by Dr. Montagne[171] in the case of a moss, Encamptodon perichætialis, in
which, in the interior of the capsule, in lieu of spores numerous minute
gemmæ of the same nature as those in the cup of Marchantia were seen.
Formation of flowers on leaves.—It is very doubtful whether a flower-bud
has ever been found actually on a leaf. Mere adhesion of the pedicels of the
leaf, such as happens in Ruscus, in Helwingia, Erythrochiton
hypophyllanthus, and a few other plants, is, of course, not really to be
considered in the light of an actual growth from the leaf, and it is very
doubtful in the present state of our knowledge whether the case of the
hyacinth.
For the formation of supernumerary leaves on the surface of the normal
one, see Multiplication and Hypertrophy.
Production of gemmæ in place of spores.—An instance of this is recorded
by Dr. Montagne[171] in the case of a moss, Encamptodon perichætialis, in
which, in the interior of the capsule, in lieu of spores numerous minute
gemmæ of the same nature as those in the cup of Marchantia were seen.
Formation of flowers on leaves.—It is very doubtful whether a flower-bud
has ever been found actually on a leaf. Mere adhesion of the pedicels of the
leaf, such as happens in Ruscus, in Helwingia, Erythrochiton
hypophyllanthus, and a few other plants, is, of course, not really to be
considered in the light of an actual growth from the leaf, and it is very
doubtful in the present state of our knowledge whether the case of the
Page 234
Nepaul barley should find a place here, but for convenience sake it is placed
in this section, as it is uncertain at present where it properly belongs.
Fig. 87.—Three-lobed end of outer palea of Nepaul barley bearing supplementary florets.
in this section, as it is uncertain at present where it properly belongs.
Fig. 87.—Three-lobed end of outer palea of Nepaul barley bearing supplementary florets.
Page 235
Fig. 88.—Three spikelets of Nepaul barley.
Page 236
Fig. 89.—Lip of outer palea of Nepaul barley.
Fig. 90.—Supplementary rachillus or outer palea of Nepaul barley bearing florets.
Fig. 91.—Diagram showing arrangement of supplementary rachillus and florets.
Fig. 90.—Supplementary rachillus or outer palea of Nepaul barley bearing florets.
Fig. 91.—Diagram showing arrangement of supplementary rachillus and florets.
Page 237
Fig. 92.—Supplementary floret of Nepaul barley; palea removed.
This curious plant has been described and figured by Irmisch in the 13th
volume of the 'Linnæa,' p. 124, t. iv; also by Professor Henslow, 'Hooker's
Journal of Botany,' 1849, vol. i, p. 33, tabs. 2, 3. The lower palea of this
plant forms an inverted flower-bud upon its midrib. In some fresh
specimens which I have lately examined I find the structure to be as
follows:—On each notch of the rachis there are three spikelets (fig. 88),
each one-flowered, and each provided with two linear glumes; the outer
palea in all cases is three-lobed at the summit, the central lobe being oblong
and hollow, forming a kind of hood (figs. 87–89), and covered with hairs,
which are directed downwards towards the centre of the plant. The two
lateral lobes are more pointed than the central one; like it they are provided
with hairs, but the hairs, in this case, are turned away from the centre of the
plant. The cavity of the side lobes is generally empty, but that of the central
lobe is occupied by a very slender stalk, which is apparently the termination
of the midrib, but which is bent inwards at an acute angle, so as to occupy
the hollow space (figs. 90–91). On this slender axis are developed two
florets, more or less imperfect in their structure. Only one of the florets that
I have seen contained a perfect ovary. The tips of the lateral lobes of the
This curious plant has been described and figured by Irmisch in the 13th
volume of the 'Linnæa,' p. 124, t. iv; also by Professor Henslow, 'Hooker's
Journal of Botany,' 1849, vol. i, p. 33, tabs. 2, 3. The lower palea of this
plant forms an inverted flower-bud upon its midrib. In some fresh
specimens which I have lately examined I find the structure to be as
follows:—On each notch of the rachis there are three spikelets (fig. 88),
each one-flowered, and each provided with two linear glumes; the outer
palea in all cases is three-lobed at the summit, the central lobe being oblong
and hollow, forming a kind of hood (figs. 87–89), and covered with hairs,
which are directed downwards towards the centre of the plant. The two
lateral lobes are more pointed than the central one; like it they are provided
with hairs, but the hairs, in this case, are turned away from the centre of the
plant. The cavity of the side lobes is generally empty, but that of the central
lobe is occupied by a very slender stalk, which is apparently the termination
of the midrib, but which is bent inwards at an acute angle, so as to occupy
the hollow space (figs. 90–91). On this slender axis are developed two
florets, more or less imperfect in their structure. Only one of the florets that
I have seen contained a perfect ovary. The tips of the lateral lobes of the
Page 238
paleæ in the primary flower are sometimes extended into a long awn. A
similar awn may also be occasionally found on the tips of the paleæ of the
rudimentary florets. The occurrence of an adventitious axial structure with
rudimentary flowers has been adduced in support of the opinion that the
lower paleæ is, at least so far as its midrib is concerned, an axial rather than
a foliar structure, but in the present uncertain state of our knowledge as to
the morphology of grasses it is hazardous to risk any explanation founded
on so exceptional a case as that of the Nepaul barley.[172]
Production of flower-buds in place of leaf-buds.—Under natural
circumstances this does not appear to be of so common occurrence as the
change above alluded to, but by the art of the gardener the change is often
effected. In rhododendrons and in peach trees and roses I have met with this
change occurring without human agency. The means adopted by the
gardener are such as check the luxuriance of the leaf-shoots,[173] and this is
effected in various ways, as by continuous "pinching" or removal of the
leaf-buds, by pruning, ringing the bark, confining the roots, limiting the
supply of nutriment, and other means all based on the same principle. Some
of the Cape bulbs (Cyrtanthus) are known not to produce their flowers till
their leaves have received, in some manner, a check. Fires which often
destroy the herbage thus have the effect of throwing the plant into bloom. A
very remarkable instance is recorded of the production of flower-buds after
an injury to the leaf-buds in the 'Bulletin of the Botanical Society of
France,' vol. ix, p. 146. It appears that during the war of the French against
the Arabs in Algiers, the latter planted several hundreds of Agaves with a
view to obstruct the passage of the French cavalry. The soldiers hacked
these plants with their sabres, and cut out the central tuft of leaves, or the
heart, as gardeners call it. The following season almost every one of these
Agaves sent up their large handsome flower-spikes. It is well known that,
under ordinary circumstances, these plants do not flower except at long
intervals of time.
similar awn may also be occasionally found on the tips of the paleæ of the
rudimentary florets. The occurrence of an adventitious axial structure with
rudimentary flowers has been adduced in support of the opinion that the
lower paleæ is, at least so far as its midrib is concerned, an axial rather than
a foliar structure, but in the present uncertain state of our knowledge as to
the morphology of grasses it is hazardous to risk any explanation founded
on so exceptional a case as that of the Nepaul barley.[172]
Production of flower-buds in place of leaf-buds.—Under natural
circumstances this does not appear to be of so common occurrence as the
change above alluded to, but by the art of the gardener the change is often
effected. In rhododendrons and in peach trees and roses I have met with this
change occurring without human agency. The means adopted by the
gardener are such as check the luxuriance of the leaf-shoots,[173] and this is
effected in various ways, as by continuous "pinching" or removal of the
leaf-buds, by pruning, ringing the bark, confining the roots, limiting the
supply of nutriment, and other means all based on the same principle. Some
of the Cape bulbs (Cyrtanthus) are known not to produce their flowers till
their leaves have received, in some manner, a check. Fires which often
destroy the herbage thus have the effect of throwing the plant into bloom. A
very remarkable instance is recorded of the production of flower-buds after
an injury to the leaf-buds in the 'Bulletin of the Botanical Society of
France,' vol. ix, p. 146. It appears that during the war of the French against
the Arabs in Algiers, the latter planted several hundreds of Agaves with a
view to obstruct the passage of the French cavalry. The soldiers hacked
these plants with their sabres, and cut out the central tuft of leaves, or the
heart, as gardeners call it. The following season almost every one of these
Agaves sent up their large handsome flower-spikes. It is well known that,
under ordinary circumstances, these plants do not flower except at long
intervals of time.
Page 239
Presence of flowers on spines.—That the spine, as a contracted branch,
should occasionally produce flowers is not to be wondered at, though the
occurrence is by no means common. M. Baillon showed at a meeting of the
Botanical Society of France ('Bulletin,' vol. v, 1858, p. 316) a branched
spine of Gleditschia bearing a flower at the end of each of the sub divisions.
This was, therefore, strictly analogous with those cases in which the
peduncle is normally spiney.
Formation of flower-bud on the petals.—An instance of this, it is
believed, the only one on record, is cited in the 'Gardeners' Chronicle' for
1865, p. 760, by the Rev. M. J. Berkeley, who describes the formation of a
flower-bud on the surface of a petal of Clarkia elegans. Reasoning from
analogy there seems no reason why buds should not be formed on the petals
as well as on the leaves.
Formation of buds on fruits.—This is a point of some moment with
reference to the share which the axis takes in the production of "inferior"
fruits. A very frequent malformation in pears is one wherein a second pear
proceeds from the centre of the first, and even a third from the centre of the
second.[174] Pears are occasionally also observed arising either from the
axils of the sepals of the primary pear or from the axil of leaves originating
on the outer surface of the fruits—using the term fruit in its popular sense.
These cases afford strong confirmation of the view that the outer portion of
the so-called fruit in these plants is rather to be considered as an expansion
and hollowing-out of the flower-stalk, than as formed from the calyx-tube.
It is noteworthy that the true carpels and seeds are frequently entirely absent
in these cases.[175] Further reference to these fruits will be made under the
head of Hypertrophy.
M. Trécul has described and figured an instance in a species of
Prismatocarpus, in which a second flower proceeded from the axil of a
should occasionally produce flowers is not to be wondered at, though the
occurrence is by no means common. M. Baillon showed at a meeting of the
Botanical Society of France ('Bulletin,' vol. v, 1858, p. 316) a branched
spine of Gleditschia bearing a flower at the end of each of the sub divisions.
This was, therefore, strictly analogous with those cases in which the
peduncle is normally spiney.
Formation of flower-bud on the petals.—An instance of this, it is
believed, the only one on record, is cited in the 'Gardeners' Chronicle' for
1865, p. 760, by the Rev. M. J. Berkeley, who describes the formation of a
flower-bud on the surface of a petal of Clarkia elegans. Reasoning from
analogy there seems no reason why buds should not be formed on the petals
as well as on the leaves.
Formation of buds on fruits.—This is a point of some moment with
reference to the share which the axis takes in the production of "inferior"
fruits. A very frequent malformation in pears is one wherein a second pear
proceeds from the centre of the first, and even a third from the centre of the
second.[174] Pears are occasionally also observed arising either from the
axils of the sepals of the primary pear or from the axil of leaves originating
on the outer surface of the fruits—using the term fruit in its popular sense.
These cases afford strong confirmation of the view that the outer portion of
the so-called fruit in these plants is rather to be considered as an expansion
and hollowing-out of the flower-stalk, than as formed from the calyx-tube.
It is noteworthy that the true carpels and seeds are frequently entirely absent
in these cases.[175] Further reference to these fruits will be made under the
head of Hypertrophy.
M. Trécul has described and figured an instance in a species of
Prismatocarpus, in which a second flower proceeded from the axil of a
Page 240
bract attached to the side of the fruit of the first flower.[176] A similar growth
was observed in the fruit of Philadelphus speciosus by M. A. Gris, who
observed that the so-called calyx-tube was provided with two small bracts,
from the axil of one of which proceeded a small flower-bud.[177]
Fig. 93.—Small buds projecting from the edges of the fruit in Opuntia.
The fruits of Opuntia Salmiana, O. fragilis,[178] O. monacantha, and of
some species of Echinocactus, have been observed to form small fruit-like
branches around their summits. M. Napoléon Doumet describes the fruit as
ripening as usual, but as being destitute of seeds in the interior; after a little
while the fruit begins to wither, and then a circle of small buds, like those of
the stem, may be seen at the top of the fruit, each bud springing from the
axil of a little tuft of wool and spines found on the fruit. These little buds
elongate into long shoots, produce flowers the following year, which
flowers exhibit the same peculiarity. Gasparini and Tenore are said to have
was observed in the fruit of Philadelphus speciosus by M. A. Gris, who
observed that the so-called calyx-tube was provided with two small bracts,
from the axil of one of which proceeded a small flower-bud.[177]
Fig. 93.—Small buds projecting from the edges of the fruit in Opuntia.
The fruits of Opuntia Salmiana, O. fragilis,[178] O. monacantha, and of
some species of Echinocactus, have been observed to form small fruit-like
branches around their summits. M. Napoléon Doumet describes the fruit as
ripening as usual, but as being destitute of seeds in the interior; after a little
while the fruit begins to wither, and then a circle of small buds, like those of
the stem, may be seen at the top of the fruit, each bud springing from the
axil of a little tuft of wool and spines found on the fruit. These little buds
elongate into long shoots, produce flowers the following year, which
flowers exhibit the same peculiarity. Gasparini and Tenore are said to have
Page 241
recorded the same fact as long since as 1832. The specimen from which the
figure (fig. 93) was taken produced its fruits in the Royal Gardens at Kew,
and is now preserved in the museum of that establishment. The adventitious
growth in these cases appears to arise from the tufts of spines, which, it has
been suggested, are the homologues of the sepals. There can, however, be
little doubt that the outer and lower portion of the fruit of Opuntia and its
allies is a dilatation of the flower-stalk. This is borne out by the fruits of
Pereskia, which bear leaves on their surface arranged spirally; indeed, the
fruits of Pereskia Bleo are mentioned as producing buds from their
summits, in the same way as the Opuntia just cited. P. Bleo is said, by M.
Delavaud,[179] to present this anomaly as a constant occurrence. On the
summit of the primary fruit, arising apparently from the axils of the sepals,
or of small leafy bracts in that situation, are a series of fruit-like branches,
which, in their turn, are surmounted by others, even to the fourth
generation.
The fruits of Tetragonia expansa frequently have attached to their side a
secondary flower or fruit in such a position as to lead to the inference that it
springs from the upper portion of the peduncle which is dilated to invest the
true carpels. In other instances it is due to an adhesion of the pedicel to the
side of the fruit. In either case the production of an adventitious bud might
be considered as an illustration of prolification of the inflorescence, though
not as was supposed by Moquin and others of axillary prolification.[180]
Buds have also been produced artificially on the surface of some of the
fruits in the construction of which the axis is supposed to share; thus, the
unripe fruits of some species of Lecythis were stated by Von Martius, at a
meeting of the German Naturalists at Carlsruhe, to produce buds when
placed in the earth. The fruit of these plants is probably of the same nature
as that of the Pomaceæ, and Baillon[181] succeeded in producing buds on
the surface of the inferior ovary of Jussiæa.
figure (fig. 93) was taken produced its fruits in the Royal Gardens at Kew,
and is now preserved in the museum of that establishment. The adventitious
growth in these cases appears to arise from the tufts of spines, which, it has
been suggested, are the homologues of the sepals. There can, however, be
little doubt that the outer and lower portion of the fruit of Opuntia and its
allies is a dilatation of the flower-stalk. This is borne out by the fruits of
Pereskia, which bear leaves on their surface arranged spirally; indeed, the
fruits of Pereskia Bleo are mentioned as producing buds from their
summits, in the same way as the Opuntia just cited. P. Bleo is said, by M.
Delavaud,[179] to present this anomaly as a constant occurrence. On the
summit of the primary fruit, arising apparently from the axils of the sepals,
or of small leafy bracts in that situation, are a series of fruit-like branches,
which, in their turn, are surmounted by others, even to the fourth
generation.
The fruits of Tetragonia expansa frequently have attached to their side a
secondary flower or fruit in such a position as to lead to the inference that it
springs from the upper portion of the peduncle which is dilated to invest the
true carpels. In other instances it is due to an adhesion of the pedicel to the
side of the fruit. In either case the production of an adventitious bud might
be considered as an illustration of prolification of the inflorescence, though
not as was supposed by Moquin and others of axillary prolification.[180]
Buds have also been produced artificially on the surface of some of the
fruits in the construction of which the axis is supposed to share; thus, the
unripe fruits of some species of Lecythis were stated by Von Martius, at a
meeting of the German Naturalists at Carlsruhe, to produce buds when
placed in the earth. The fruit of these plants is probably of the same nature
as that of the Pomaceæ, and Baillon[181] succeeded in producing buds on
the surface of the inferior ovary of Jussiæa.
Page 242
Some of the cases just mentioned have been considered to be instances of
prolification of the fruit, but the fruit has little to do with the appearances in
question.
Formation of adventitious flowers and fruits within the ovary.—This
generally arises either from substitution of a flower-bud for an ovule or
from prolification; there are certain cases, however, where the new growth
seems not to be either due to metamorphosis or to prolification strictly.
The cut, fig. 94, represents a case where, in the dilated upper portion of the
ovary of Sinapis arvensis, two flower-buds were found projecting from a
raised central line, corresponding, as it would seem, to the midrib, and not
to the margins of the carpel. Similar cases have occurred in Nasturtium
amphibium, Brassica Rapa, and Passiflora quadrangularis.
Fig. 94.—Distended pod of Sinapis arvensis bearing in the interior stalked flower buds.
prolification of the fruit, but the fruit has little to do with the appearances in
question.
Formation of adventitious flowers and fruits within the ovary.—This
generally arises either from substitution of a flower-bud for an ovule or
from prolification; there are certain cases, however, where the new growth
seems not to be either due to metamorphosis or to prolification strictly.
The cut, fig. 94, represents a case where, in the dilated upper portion of the
ovary of Sinapis arvensis, two flower-buds were found projecting from a
raised central line, corresponding, as it would seem, to the midrib, and not
to the margins of the carpel. Similar cases have occurred in Nasturtium
amphibium, Brassica Rapa, and Passiflora quadrangularis.
Fig. 94.—Distended pod of Sinapis arvensis bearing in the interior stalked flower buds.
Page 243
In Bromfield's 'Flora Vectensis,' p. 35, the following account is given of an
abnormal development in Cardamine pratensis: "On the lower part of the
corymb were several seed vessels on pedicels changed from their usual
linear to an ovate elliptical figure, so as to resemble a silicula. These, on
being opened, were found to contain petals of the usual colour, which in the
pods above had burst from their confinement and appeared as semi-double
flowers; the valves of the pod answering to the true calyx. * * * From their
verticillate arrangement it is evident that these petaloid expansions were not
transformed seeds, but simply a development of the common axis within
the ovary into an abortive whorl of floral organs, besides which there were
evident rudiments both of stamens and germens in the centre of the bundle."
Baillon[182] also records a case of the same nature in Sinapis arvensis.
Fig. 94*.—Portion of the interior of the silicle in Cheiranthus Cheiri, showing adventitious pod in
the place of an ovule.
abnormal development in Cardamine pratensis: "On the lower part of the
corymb were several seed vessels on pedicels changed from their usual
linear to an ovate elliptical figure, so as to resemble a silicula. These, on
being opened, were found to contain petals of the usual colour, which in the
pods above had burst from their confinement and appeared as semi-double
flowers; the valves of the pod answering to the true calyx. * * * From their
verticillate arrangement it is evident that these petaloid expansions were not
transformed seeds, but simply a development of the common axis within
the ovary into an abortive whorl of floral organs, besides which there were
evident rudiments both of stamens and germens in the centre of the bundle."
Baillon[182] also records a case of the same nature in Sinapis arvensis.
Fig. 94*.—Portion of the interior of the silicle in Cheiranthus Cheiri, showing adventitious pod in
the place of an ovule.
Page 244
Fig. 95.—Adventitious pod from fig. 94, enlarged.
Here, too, may also be mentioned the presence of an adventitious siliqua
within the ordinary one attached along the same line as the ovules, and
partially divided by a replum into two cavities. In this case there was
nothing to indicate the presence of floral envelopes (figs. 94, 95). A similar
occurrence has been brought under my notice in some grapes which were
observed to be cracking before they were perfectly ripe, and in which
adventitious fruits were found within the parent grape, occupying the
position of seeds (figs. 96, 97).
Similar anomalous growths are noticed under the heads of Substitution and
Prolification.
Formation of stamens within the cavity of the ovary.—The only instance
of this that has come under the author's observation occurred in some
Here, too, may also be mentioned the presence of an adventitious siliqua
within the ordinary one attached along the same line as the ovules, and
partially divided by a replum into two cavities. In this case there was
nothing to indicate the presence of floral envelopes (figs. 94, 95). A similar
occurrence has been brought under my notice in some grapes which were
observed to be cracking before they were perfectly ripe, and in which
adventitious fruits were found within the parent grape, occupying the
position of seeds (figs. 96, 97).
Similar anomalous growths are noticed under the heads of Substitution and
Prolification.
Formation of stamens within the cavity of the ovary.—The only instance
of this that has come under the author's observation occurred in some
Page 245
flowers of Bæckea diosmæfolia, Rudge, for an examination of which he is
indebted to Mr. Bentham.
Fig. 96.—Section of Barbarossa grape showing adventitious grape in the position of a seed.
indebted to Mr. Bentham.
Fig. 96.—Section of Barbarossa grape showing adventitious grape in the position of a seed.
Page 246
Fig. 97.—Grape with supplementary fruit in the interior
In the normal flower there is a turbinate hollow calyx, whose limb is
divided into five serrated lobes; alternating with these latter, and springing
from the throat of the calyx, are the petals. Originating from the same
annular disk as the petals are the stamens, seven or eight in number. The
ovary is partially adherent, is surmounted by a style, and has two or three
loculi with an axile placenta, to which several small curved ovules are
attached. The malformed flowers did not present anything peculiar in their
outer parts, nor did the ovary, partially immersed within the expanded top
of the flower-stalk and the calyx-tube, which is continuous with that organ,
show externally any indication of the change within. On cutting it across,
however, in any direction, numerous perfect stamens (filaments and
anthers) were seen projecting from the walls of the cavity (fig. 98). In most
of the flowers the ovary was one-celled; but in a few there was the usual
axile placenta; yet even in these latter cases the stamens originated from the
walls of the cavity, and not from the placenta. The stamens presented
different degrees of development; in some cases they were fully formed, the
anther-lobes open, and the pollen exposed; while in other instances the
filaments were involute or circinate, just as the ordinary stamens are in the
unexpanded flower-bud. In some cases imperfect stamens were found, mere
barren filaments, with or without rudimentary anthers at the top. In no
instance was there a perfect ovule, or, indeed, any trace of ovules. The
stamens appeared to be arranged irregularly on the walls of the ovarian
cavity; and while they were certainly more numerous at the lower portion
(that now generally considered to be formed by the cup-like end of the
pedicel), they were not wanting in the upper half of the ovary (or that which
is probably formed from the carpellary leaves).
In the normal flower there is a turbinate hollow calyx, whose limb is
divided into five serrated lobes; alternating with these latter, and springing
from the throat of the calyx, are the petals. Originating from the same
annular disk as the petals are the stamens, seven or eight in number. The
ovary is partially adherent, is surmounted by a style, and has two or three
loculi with an axile placenta, to which several small curved ovules are
attached. The malformed flowers did not present anything peculiar in their
outer parts, nor did the ovary, partially immersed within the expanded top
of the flower-stalk and the calyx-tube, which is continuous with that organ,
show externally any indication of the change within. On cutting it across,
however, in any direction, numerous perfect stamens (filaments and
anthers) were seen projecting from the walls of the cavity (fig. 98). In most
of the flowers the ovary was one-celled; but in a few there was the usual
axile placenta; yet even in these latter cases the stamens originated from the
walls of the cavity, and not from the placenta. The stamens presented
different degrees of development; in some cases they were fully formed, the
anther-lobes open, and the pollen exposed; while in other instances the
filaments were involute or circinate, just as the ordinary stamens are in the
unexpanded flower-bud. In some cases imperfect stamens were found, mere
barren filaments, with or without rudimentary anthers at the top. In no
instance was there a perfect ovule, or, indeed, any trace of ovules. The
stamens appeared to be arranged irregularly on the walls of the ovarian
cavity; and while they were certainly more numerous at the lower portion
(that now generally considered to be formed by the cup-like end of the
pedicel), they were not wanting in the upper half of the ovary (or that which
is probably formed from the carpellary leaves).
Page 247
Fig. 98.—1. Vertical section of flower of Bæckea diosmæfolia, showing stamens within the ovary;
magnified ten times. 2. Transverse section of ovary. 3. Stamen. 4. Imperfect stamen.
This case differs from most that have been recorded, and in which there has
been a more or less complete substitution of anther for carpel, or where the
tissues of the carpel have produced pollen, and so taken upon themselves
the appearance and functions of anthers. Instances of this latter kind are not
uncommon; but in the Bæckea there were perfect stamens proceeding from
perfect and completely closed ovaries. Moquin-Tandon[183] cites from
Agardh an instance which seems more closely to resemble the state of
things in the Bæckea, and which occurred in a double hyacinth, wherein
both anthers and ovules were borne on the same placenta. Probably, though
the fact is not stated, the ovary of the hyacinth was open; and we are told
that the flower was double—that it was, in fact, modified and changed in
more organs than one; while in the Bæckea nothing at all unusual was
observed till the ovary was cut open. The style was present even in those
flowers where there was no axile placenta; hence in these cases it could not
magnified ten times. 2. Transverse section of ovary. 3. Stamen. 4. Imperfect stamen.
This case differs from most that have been recorded, and in which there has
been a more or less complete substitution of anther for carpel, or where the
tissues of the carpel have produced pollen, and so taken upon themselves
the appearance and functions of anthers. Instances of this latter kind are not
uncommon; but in the Bæckea there were perfect stamens proceeding from
perfect and completely closed ovaries. Moquin-Tandon[183] cites from
Agardh an instance which seems more closely to resemble the state of
things in the Bæckea, and which occurred in a double hyacinth, wherein
both anthers and ovules were borne on the same placenta. Probably, though
the fact is not stated, the ovary of the hyacinth was open; and we are told
that the flower was double—that it was, in fact, modified and changed in
more organs than one; while in the Bæckea nothing at all unusual was
observed till the ovary was cut open. The style was present even in those
flowers where there was no axile placenta; hence in these cases it could not
Page 248
be, as Lindley stated it to be in the closely allied Babingtonia, a
prolongation of the placenta.[184]
Formation of pollen within the ovules.—This has now been recorded in
two instances by Mr. S. J. A. Salter in Passiflora cærulea and in P. palmata,
[185] and by the author in Rosa arvensis.[186]
Fig. 99.—Pollen within the ovule of Passiflora (after Salter).
In the case of the passion-flower there were various malformations in the
ovaries, which were all more or less split open at the distal end, indicating a
tendency towards dialysis. The pollen-bearing ovules were borne on the
edges of these ovaries, and presented various intermediate conditions
between anthers and ovules, commencing at the distal extremity of the
carpel with a bi-lobed anther, and passing in series to the base of the ovary,
an antheroid body of ovule-like form, a modified ovule containing pollen,
an ovule departing from a perfectly natural condition only in the
prolongation of the placenta.[184]
Formation of pollen within the ovules.—This has now been recorded in
two instances by Mr. S. J. A. Salter in Passiflora cærulea and in P. palmata,
[185] and by the author in Rosa arvensis.[186]
Fig. 99.—Pollen within the ovule of Passiflora (after Salter).
In the case of the passion-flower there were various malformations in the
ovaries, which were all more or less split open at the distal end, indicating a
tendency towards dialysis. The pollen-bearing ovules were borne on the
edges of these ovaries, and presented various intermediate conditions
between anthers and ovules, commencing at the distal extremity of the
carpel with a bi-lobed anther, and passing in series to the base of the ovary,
an antheroid body of ovule-like form, a modified ovule containing pollen,
an ovule departing from a perfectly natural condition only in the
Page 249
development of a few grains of pollen in its nucleus, and, finally, a perfect,
normal ovule.
In the flowers of the Rose the stamens exhibited almost every conceivable
gradation between their ordinary form and that of the carpels, while some of
the ovules contained pollen in greater or less abundance. Speaking
generally, the most common state of things in these flowers was the
occurrence on the throat of the calyx, in the position ordinarily occupied by
the stamens, and sometimes mingled with those organs, of twisted, ribbon-
like filaments, which bore about the centre one or more pendulous,
anatropous ovules on their margins. Immediately above the latter organs
were the anther-lobes, more or less perfectly developed, and surmounting
these a long style, terminating in a fringed, funnel-shaped stigma.
Sometimes the ovules were perfect, at other times the nucleus protruded
through the foramen, while in a third set the nucleus was included within
the tegument, the ovules having in all respects their natural external
conformation, containing, however, not only pollen-grains, but also a layer
of those peculiar spheroidal cells, including a fibrous deposit, which are
among the normal constituents of the anther. In one case, where the coat of
the ovule was imperfect, and allowed the nucleus to protrude, the pollen
was evidently contained within the central mass of the structure. In this
instance the fibrous cells were not detected, these being only found in cases
where the investment of the ovule was perfect; and hence it seems likely
that the fibrous cells were part of the coat of the ovule, while the pollen was
formed within the nucleus. In no case was any trace of embryo sac to be
seen.
The main interest, as Mr. Salter remarks, in these cases is physiological; so
far as structure alone is concerned, there does not appear any reason why
pollen-grains should not be developed in any portion of the plant; the
normal ovule.
In the flowers of the Rose the stamens exhibited almost every conceivable
gradation between their ordinary form and that of the carpels, while some of
the ovules contained pollen in greater or less abundance. Speaking
generally, the most common state of things in these flowers was the
occurrence on the throat of the calyx, in the position ordinarily occupied by
the stamens, and sometimes mingled with those organs, of twisted, ribbon-
like filaments, which bore about the centre one or more pendulous,
anatropous ovules on their margins. Immediately above the latter organs
were the anther-lobes, more or less perfectly developed, and surmounting
these a long style, terminating in a fringed, funnel-shaped stigma.
Sometimes the ovules were perfect, at other times the nucleus protruded
through the foramen, while in a third set the nucleus was included within
the tegument, the ovules having in all respects their natural external
conformation, containing, however, not only pollen-grains, but also a layer
of those peculiar spheroidal cells, including a fibrous deposit, which are
among the normal constituents of the anther. In one case, where the coat of
the ovule was imperfect, and allowed the nucleus to protrude, the pollen
was evidently contained within the central mass of the structure. In this
instance the fibrous cells were not detected, these being only found in cases
where the investment of the ovule was perfect; and hence it seems likely
that the fibrous cells were part of the coat of the ovule, while the pollen was
formed within the nucleus. In no case was any trace of embryo sac to be
seen.
The main interest, as Mr. Salter remarks, in these cases is physiological; so
far as structure alone is concerned, there does not appear any reason why
pollen-grains should not be developed in any portion of the plant; the
Page 250
mother cells in which the pollen is formed not differing, to all outward
appearance, from any other cells, unless it be in size.
The fundamental unity of construction in all the organs of plants could
hardly be better illustrated than by these cases; while, in spite of their
exceptional nature, they must be of great interest physiologically, as
showing the wide limits of possible variation which thus may even involve
the sex, "for an ovule to develop pollen within its interior," says Mr. Salter,
"is equivalent to an ovum in an animal being converted into a capsule of
spermatozoa. It is a conversion of germ into sperm, the most complete
violation of individuality and unity of sex. * * * * The occurrence of an
antheroid ovule and a normal ovule on the same carpellary leaf realises the
simplest and the most absolute form of hermaphroditism."
It must, however, be remarked that the term substitution would be
preferable to conversion. There is, at present, no evidence to show that the
germinal vesicles were present in these cases; on the other hand, it seems
most probable that they were not, so that the presence of the pollen-cells
must be considered as simply adventitious. It can hardly be that they were,
in the first instance, germinal vesicles, which, in course of time, became so
modified as to assume the appearance of pollen-grains. Between the nucleus
of the ovule and the tubercle of cellular tissue constituting the primordial
anther, there is little or no difference, so that it may be said that, for a time,
there is no distinction of sex in the nascent flower, but as development goes
on, the difference becomes perceptible. It cannot at present be stated what
precise circumstances induce the one mass to form mother-cells and pollen-
grains, and the other to develop an embryo sac and germinal vesicles.
Position and external circumstances may have some indirect effect, and it
may, perhaps, be significant that in all the instances of polliniferous ovules,
the ovular structures have been exposed on an open carpel or otherwise, in
place of being confined within the cavity of a closed ovary, as under
appearance, from any other cells, unless it be in size.
The fundamental unity of construction in all the organs of plants could
hardly be better illustrated than by these cases; while, in spite of their
exceptional nature, they must be of great interest physiologically, as
showing the wide limits of possible variation which thus may even involve
the sex, "for an ovule to develop pollen within its interior," says Mr. Salter,
"is equivalent to an ovum in an animal being converted into a capsule of
spermatozoa. It is a conversion of germ into sperm, the most complete
violation of individuality and unity of sex. * * * * The occurrence of an
antheroid ovule and a normal ovule on the same carpellary leaf realises the
simplest and the most absolute form of hermaphroditism."
It must, however, be remarked that the term substitution would be
preferable to conversion. There is, at present, no evidence to show that the
germinal vesicles were present in these cases; on the other hand, it seems
most probable that they were not, so that the presence of the pollen-cells
must be considered as simply adventitious. It can hardly be that they were,
in the first instance, germinal vesicles, which, in course of time, became so
modified as to assume the appearance of pollen-grains. Between the nucleus
of the ovule and the tubercle of cellular tissue constituting the primordial
anther, there is little or no difference, so that it may be said that, for a time,
there is no distinction of sex in the nascent flower, but as development goes
on, the difference becomes perceptible. It cannot at present be stated what
precise circumstances induce the one mass to form mother-cells and pollen-
grains, and the other to develop an embryo sac and germinal vesicles.
Position and external circumstances may have some indirect effect, and it
may, perhaps, be significant that in all the instances of polliniferous ovules,
the ovular structures have been exposed on an open carpel or otherwise, in
place of being confined within the cavity of a closed ovary, as under
Page 251
ordinary circumstances. Even among Conifers the ovuligerous scales are so
closely packed that there is little or no exposure of the ovules. But, apart
from all speculative notions as to the relation between the structure and
functions of the anther and of the ovule respectively, and of the possibility
or the reverse of parthenogenesis, it will clearly be necessary in any future
alleged occurrence of the latter phenomenon to ascertain whether any or all
of the apparent ovules are, or are not, anthers in disguise.
Homomorphic flowers of "Compositæ."—In a large section of the
Compositæ there is, as is well known, a distinction between the florets of
the "disc" and those of the "ray," the latter being ligulate, the former tubular.
In what are erroneously called double flowers in this order, e.g. in the
Chrysanthemum, Dahlia, &c. &c., the florets are all ligulate. This change is
sometimes classed with peloria, but there is no abnormal regularity in these
cases. On the other hand, were the ligulate florets to be all replaced by
tubular ones, the term peloria would be more strictly applicable. It will be
remembered that in the sub-order Ligulifloræ, the florets are naturally all
ligulate, so that the change above mentioned is not in itself a very grave
one.
Heterotaxy affecting the inflorescence.—Under the head of Prolification,
Heterogamy, &c., various deviations from the normal inflorescence are
alluded to. In this place, therefore, it is only necessary to mention certain
rare deviations from the customary arrangement of the inflorescence, such
as the change from a definite centrifugal form of inflorescence to an
indefinite centripetal one. This occurs occasionally in roses, where the
shoot, instead of terminating in a flower-bud, lengthens and bears the
flower-bud on its sides as in a raceme.
In the hyacinth, the inflorescence of which is properly indefinite, the
terminal flower may frequently be found to expand first, though in order of
closely packed that there is little or no exposure of the ovules. But, apart
from all speculative notions as to the relation between the structure and
functions of the anther and of the ovule respectively, and of the possibility
or the reverse of parthenogenesis, it will clearly be necessary in any future
alleged occurrence of the latter phenomenon to ascertain whether any or all
of the apparent ovules are, or are not, anthers in disguise.
Homomorphic flowers of "Compositæ."—In a large section of the
Compositæ there is, as is well known, a distinction between the florets of
the "disc" and those of the "ray," the latter being ligulate, the former tubular.
In what are erroneously called double flowers in this order, e.g. in the
Chrysanthemum, Dahlia, &c. &c., the florets are all ligulate. This change is
sometimes classed with peloria, but there is no abnormal regularity in these
cases. On the other hand, were the ligulate florets to be all replaced by
tubular ones, the term peloria would be more strictly applicable. It will be
remembered that in the sub-order Ligulifloræ, the florets are naturally all
ligulate, so that the change above mentioned is not in itself a very grave
one.
Heterotaxy affecting the inflorescence.—Under the head of Prolification,
Heterogamy, &c., various deviations from the normal inflorescence are
alluded to. In this place, therefore, it is only necessary to mention certain
rare deviations from the customary arrangement of the inflorescence, such
as the change from a definite centrifugal form of inflorescence to an
indefinite centripetal one. This occurs occasionally in roses, where the
shoot, instead of terminating in a flower-bud, lengthens and bears the
flower-bud on its sides as in a raceme.
In the hyacinth, the inflorescence of which is properly indefinite, the
terminal flower may frequently be found to expand first, though in order of
Page 252
development it may have been the last formed.
It occasionally happens that certain plants will, contrary to their usual
custom, bloom twice in the same season; this usually arises from the
premature development of buds which, under ordinary circumstances,
would not unfold till the following spring. In these instances of what the
French term "fleuraison anticipée," the position of inflorescence is not
changed, but there are other cases where the position of the inflorescence is
altered, as in the laburnum, where, in some seasons, racemes may be seen
springing from short lateral "spurs" along the sides of the branches, as well
as from the extremities of long shoots.
Of a similar nature are those cases wherein stems or branches usually sterile
become fertile; this happens in Equisetaceæ,[187] in Restiaceæ, and other
orders. In the equisetums, the condition in question has been specially
noticed to occur after prolonged drought.
Equisetaceæ are likewise subject to an anomaly called by Duval Jouve
interruption of the spike, and wherein the scales bearing the spore cases are
separated by whorls of branches instead of forming one compact unbroken
spike as usual.
This alternation of the organs of vegetation and reproduction may also be
seen occasionally in Typha, and other plants.
Kirschleger describes a case in which the male catkins of Salix cinerea were
placed at the ends of the branches instead of being lateral productions;
moreover the usual articulation was not formed, so that the catkin was
persistent instead of deciduous.[188]
Supra-soriferous ferns.—In the great majority of ferns the sori or clusters
of spore cases are placed on the under surface of the fronds; nevertheless, a
It occasionally happens that certain plants will, contrary to their usual
custom, bloom twice in the same season; this usually arises from the
premature development of buds which, under ordinary circumstances,
would not unfold till the following spring. In these instances of what the
French term "fleuraison anticipée," the position of inflorescence is not
changed, but there are other cases where the position of the inflorescence is
altered, as in the laburnum, where, in some seasons, racemes may be seen
springing from short lateral "spurs" along the sides of the branches, as well
as from the extremities of long shoots.
Of a similar nature are those cases wherein stems or branches usually sterile
become fertile; this happens in Equisetaceæ,[187] in Restiaceæ, and other
orders. In the equisetums, the condition in question has been specially
noticed to occur after prolonged drought.
Equisetaceæ are likewise subject to an anomaly called by Duval Jouve
interruption of the spike, and wherein the scales bearing the spore cases are
separated by whorls of branches instead of forming one compact unbroken
spike as usual.
This alternation of the organs of vegetation and reproduction may also be
seen occasionally in Typha, and other plants.
Kirschleger describes a case in which the male catkins of Salix cinerea were
placed at the ends of the branches instead of being lateral productions;
moreover the usual articulation was not formed, so that the catkin was
persistent instead of deciduous.[188]
Supra-soriferous ferns.—In the great majority of ferns the sori or clusters
of spore cases are placed on the under surface of the fronds; nevertheless, a
Page 253
few cases are on record where the fructification is produced on the upper as
well as on the lower surface, and sometimes abundantly so. This
occasionally happens from the elongation of the normally placed sorus,
which thus extends to the margin, and returns on the upper side, when the
sori chance to be placed opposite to the marginal crenatures. But it is also
frequently the case that the sori are produced on the upper side, distinctly
within the margin, and where there are no corresponding sori beneath.
Those varieties which have the margin crenated or lobed seem most liable
to assume this abnormal supra-soriferous condition. Among the ferns in
which this condition has been observed are the following: Scolopendrium
vulgare, Polypodium anomalum, Hook., Asplenium Trichomanes,
Cionidium Moorei.[189]
FOOTNOTES:
[147] 'Gard. Chron.' 1852, p. 51.
[148] 'Flora.' 1858, pp. 32–42.
[149] 'Journ. Linn. Soc.,' vol. vi; "Botany," 1862, p. 24.
[150] 'Tijdschrift voor Natuur. Geschied,' 1836, vol. iii, tab. vii, p. 171.
[151] Rœper, 'Enum. Euphorb.,' p. 19. Bernhardi, 'Linnæa,' vii, p. 561, tab. xiv, f. 1. Wydler,
"Subcotyled. sprossbildung," 'Flora,' 1850, p. 337. Hooker, 'Trans. Linn. Soc.,' vol. xxiv, p. 20
(Welwitschia).
[152] 'Misc. Austriac. ad Bot.,' vol. i, p. 133, t. 5.
[153] See also Carrière, 'Revue Horticole,' 1866, p. 442; and as to pears, Radlkofer in
'Bericht über die Thätigkert der Baierischen Gartenbau Gesellschaft,' 1862, p. 74, t. i.
[154] 'Flora Indica,' p. 23.
[155] 'Bull. Soc. Bot. Fr.,' 1856, p. 53.
[156] 'Trans. Linn. Soc.' xxvi, p. 142, tab. iv, B.
[157] "Si arbusculam, quæ in ollâ antea posita, quotannis floruit et fructus protulit, deinde
deponamus in uberiori terra calidi caldarii, proferet illa per plures annos multos ac frondosos
well as on the lower surface, and sometimes abundantly so. This
occasionally happens from the elongation of the normally placed sorus,
which thus extends to the margin, and returns on the upper side, when the
sori chance to be placed opposite to the marginal crenatures. But it is also
frequently the case that the sori are produced on the upper side, distinctly
within the margin, and where there are no corresponding sori beneath.
Those varieties which have the margin crenated or lobed seem most liable
to assume this abnormal supra-soriferous condition. Among the ferns in
which this condition has been observed are the following: Scolopendrium
vulgare, Polypodium anomalum, Hook., Asplenium Trichomanes,
Cionidium Moorei.[189]
FOOTNOTES:
[147] 'Gard. Chron.' 1852, p. 51.
[148] 'Flora.' 1858, pp. 32–42.
[149] 'Journ. Linn. Soc.,' vol. vi; "Botany," 1862, p. 24.
[150] 'Tijdschrift voor Natuur. Geschied,' 1836, vol. iii, tab. vii, p. 171.
[151] Rœper, 'Enum. Euphorb.,' p. 19. Bernhardi, 'Linnæa,' vii, p. 561, tab. xiv, f. 1. Wydler,
"Subcotyled. sprossbildung," 'Flora,' 1850, p. 337. Hooker, 'Trans. Linn. Soc.,' vol. xxiv, p. 20
(Welwitschia).
[152] 'Misc. Austriac. ad Bot.,' vol. i, p. 133, t. 5.
[153] See also Carrière, 'Revue Horticole,' 1866, p. 442; and as to pears, Radlkofer in
'Bericht über die Thätigkert der Baierischen Gartenbau Gesellschaft,' 1862, p. 74, t. i.
[154] 'Flora Indica,' p. 23.
[155] 'Bull. Soc. Bot. Fr.,' 1856, p. 53.
[156] 'Trans. Linn. Soc.' xxvi, p. 142, tab. iv, B.
[157] "Si arbusculam, quæ in ollâ antea posita, quotannis floruit et fructus protulit, deinde
deponamus in uberiori terra calidi caldarii, proferet illa per plures annos multos ac frondosos
Page 254
ramos, sine ullo fructu. Id quod argumento est, folia inde crescere, unde prius enati sunt flores;
quemadmodum vicissim, quod in folia nunc succrescit, id, naturâ ita moderante, in flores
mutatur, si eadem arbor iterum in ollâ seritur."—Linnæus, 'Prolepsis,' § iii.
[158] 'Rev. Hortic.' May, 1868, 'Gardeners' Chronicle,' 1868, pp. 572, 737.
[159] Cited in 'Annals Nat. Hist.,' 1845, vol. xv, p. 177.
[160] 'Ann. Scienc. Nat.,' vol. xiv, 1860, p. 13.
[161] Naudin, 'Ann. Sc. Nat.,' 2nd ser., 1840, vol. xiv, p. 14, fig. 6, pl. i (Drosera). St.
Hilaire, 'Comptes Rendus,' ix, p. 437.
[162] Hance, 'Hook. Journ. Botany,' 1849, vol. i, p. 141, pl. v.
[163] Booth, 'Gard. Chron.,' Jan. 1st, 1853, p. 4.
[164] Lindley, 'Theory of Horticulture,' ed. 2, p. 273.
[165] 'Hook. Journ. of Botany,' 1852, iv, p. 206. See also the curious Begonia gemmipara,
'Hook. fil. Illust. Himal. Plant.,' t. xiv.
[166] 'Phytanth.,' n. 36, d.
[167] 'Ann. Scienc. Nat.,' 3rd series. 1853. vol. xix, p. 251, tab. 14.
[168] Carrière, 'Revue Horticole.' 1868, p. 184.
[169] 'Gard. Chron.,' 1858, p. 556.
[170] 1863, p. 556, &c.
[171] 'Ann. Nat. Hist.,' 1845, vol. xvi, p. 355.
[172] See also Lindley, 'Veg. Kingd.,' p. 109 et 116a, where the views of Raspail, R. Brown,
Mohl, Henslow, and others, are discussed.
[173] It has been observed that if a plant is supplied with copious nourishment the flowering-
period is delayed; but that moderate or even scanty nourishment accelerates it. Goethe,
'Metam.,' § 30. See also Wolff, 'Theoria Generationis,' 1759; Linn. 'Prolepsis,' §§ 3 and 10.
[174] Moquin-Tandon, p. 384; also Lindl., 'Elements of Botany,' p. 65, fig. 130; "Theory of
Horticulture," p. 86. 'Gard. Chron.,' 1851, p. 723; Irmish, 'Flora,' 1858, p. 38, &c.
[175] Caspary, 'Bull. Soc. Bot. Fr.,' vol. vi, 1859, p. 235; also Payer, ibid., vol. i, 1854. p.
283.
quemadmodum vicissim, quod in folia nunc succrescit, id, naturâ ita moderante, in flores
mutatur, si eadem arbor iterum in ollâ seritur."—Linnæus, 'Prolepsis,' § iii.
[158] 'Rev. Hortic.' May, 1868, 'Gardeners' Chronicle,' 1868, pp. 572, 737.
[159] Cited in 'Annals Nat. Hist.,' 1845, vol. xv, p. 177.
[160] 'Ann. Scienc. Nat.,' vol. xiv, 1860, p. 13.
[161] Naudin, 'Ann. Sc. Nat.,' 2nd ser., 1840, vol. xiv, p. 14, fig. 6, pl. i (Drosera). St.
Hilaire, 'Comptes Rendus,' ix, p. 437.
[162] Hance, 'Hook. Journ. Botany,' 1849, vol. i, p. 141, pl. v.
[163] Booth, 'Gard. Chron.,' Jan. 1st, 1853, p. 4.
[164] Lindley, 'Theory of Horticulture,' ed. 2, p. 273.
[165] 'Hook. Journ. of Botany,' 1852, iv, p. 206. See also the curious Begonia gemmipara,
'Hook. fil. Illust. Himal. Plant.,' t. xiv.
[166] 'Phytanth.,' n. 36, d.
[167] 'Ann. Scienc. Nat.,' 3rd series. 1853. vol. xix, p. 251, tab. 14.
[168] Carrière, 'Revue Horticole.' 1868, p. 184.
[169] 'Gard. Chron.,' 1858, p. 556.
[170] 1863, p. 556, &c.
[171] 'Ann. Nat. Hist.,' 1845, vol. xvi, p. 355.
[172] See also Lindley, 'Veg. Kingd.,' p. 109 et 116a, where the views of Raspail, R. Brown,
Mohl, Henslow, and others, are discussed.
[173] It has been observed that if a plant is supplied with copious nourishment the flowering-
period is delayed; but that moderate or even scanty nourishment accelerates it. Goethe,
'Metam.,' § 30. See also Wolff, 'Theoria Generationis,' 1759; Linn. 'Prolepsis,' §§ 3 and 10.
[174] Moquin-Tandon, p. 384; also Lindl., 'Elements of Botany,' p. 65, fig. 130; "Theory of
Horticulture," p. 86. 'Gard. Chron.,' 1851, p. 723; Irmish, 'Flora,' 1858, p. 38, &c.
[175] Caspary, 'Bull. Soc. Bot. Fr.,' vol. vi, 1859, p. 235; also Payer, ibid., vol. i, 1854. p.
283.
Page 255
[176] Trécul, 'Ann. Sc. Nat.,' 2nd ser., vol. xx, p. 339.
[177] 'Bull. Soc. Bot. Fr.,' vol. vii, 1858, p. 331.
[178] 'Bull. Soc. Bot. Fr.,' vol. i. p. 306, vol. v, p. 115. 'Illustr. Hortic.,' xii, 1865, Misc. 79.
'Rev. Horticole,' 1860 p. 204, et 1867 p. 43.
[179] 'Bull. Soc. Bot. Fr.,' 1858, p. 685.
[180] The structure of this flower is discussed at some length in a paper by the author on
axillary prolification. 'Trans. Linn. Soc.,' vol. xxiii, p. 486, t. liv. fig. 3. See also 'Clos. Bull.
Soc. Bot. Fr.,' vol. v, 1855, p. 672. Seringe et Heyland, 'Bull. Bot.,' i, p. 8. 'Pallas Enum. Plant.
Hort. Demidoff,' append, c, ic.
[181] 'Adansonia,' i, 181.
[182] 'Adansonia.' vol. iii, p. 351, tab. xii.
[183] 'Elém. Térat. Végét.,' p, 218.
[184] Masters, 'Journ. Linn. Soc.,' vol. ix, 1866, p. 334.
[185] 'Trans. Linn. Soc.,' vol. xxiv, p. 143. tab. xxiv.
[186] 'Brit. Assoc. Report,' Dundee, 1867; and Seemann's 'Journal of Botany,' 1867, p. 319,
tab. lxxii, figs. B 1–9.
[187] Duval Jouve, 'Hist. Equiset. France.' 1864, p. 154.
[188] 'Flora,' t. xxiv, 1841, p. 340.
[189] Moore, 'Nature-Printed British Ferns,' 8vo edition, vol. ii. p. 135. tab. lxxxv, B, &c.
CHAPTER IV.
HETEROGAMY.
This term is here intended to apply to all those cases in which the
arrangement of the sexual organs is different from what it is habitually. It is
evident that in many instances there is no malformation, no monstrosity, but
[177] 'Bull. Soc. Bot. Fr.,' vol. vii, 1858, p. 331.
[178] 'Bull. Soc. Bot. Fr.,' vol. i. p. 306, vol. v, p. 115. 'Illustr. Hortic.,' xii, 1865, Misc. 79.
'Rev. Horticole,' 1860 p. 204, et 1867 p. 43.
[179] 'Bull. Soc. Bot. Fr.,' 1858, p. 685.
[180] The structure of this flower is discussed at some length in a paper by the author on
axillary prolification. 'Trans. Linn. Soc.,' vol. xxiii, p. 486, t. liv. fig. 3. See also 'Clos. Bull.
Soc. Bot. Fr.,' vol. v, 1855, p. 672. Seringe et Heyland, 'Bull. Bot.,' i, p. 8. 'Pallas Enum. Plant.
Hort. Demidoff,' append, c, ic.
[181] 'Adansonia,' i, 181.
[182] 'Adansonia.' vol. iii, p. 351, tab. xii.
[183] 'Elém. Térat. Végét.,' p, 218.
[184] Masters, 'Journ. Linn. Soc.,' vol. ix, 1866, p. 334.
[185] 'Trans. Linn. Soc.,' vol. xxiv, p. 143. tab. xxiv.
[186] 'Brit. Assoc. Report,' Dundee, 1867; and Seemann's 'Journal of Botany,' 1867, p. 319,
tab. lxxii, figs. B 1–9.
[187] Duval Jouve, 'Hist. Equiset. France.' 1864, p. 154.
[188] 'Flora,' t. xxiv, 1841, p. 340.
[189] Moore, 'Nature-Printed British Ferns,' 8vo edition, vol. ii. p. 135. tab. lxxxv, B, &c.
CHAPTER IV.
HETEROGAMY.
This term is here intended to apply to all those cases in which the
arrangement of the sexual organs is different from what it is habitually. It is
evident that in many instances there is no malformation, no monstrosity, but
Page 256
rather a restoration of organs habitually suppressed, a tendency towards
structural completeness rather than the reverse. It must be also understood
that the following remarks apply to structural points only, and are not
intended to include the question of function. The occurrence of
heteromorphic unions renders it necessary to keep in mind that plants
hermaphrodite as to structure are by no means necessarily so as to function.
The simplest case of this alteration in the relative position of the sexes is
that which occurs in monœcious plants, where the male and female flowers
have a definite position, but which in exceptional instances is altered.
Change in the relative position of male and female flowers may thus
occur in any monœcious plant. Cultivated maize, Zea Mays, frequently
exhibits alterations of this kind; under ordinary circumstances, the male
inflorescence is a compound spike, occupying the extremity of the stem,
while the female flowers are borne in simple spikes at a lower level, but
specimens may now and then be found where the sexes are mixed in the
same inflorescence; the upper branching panicle usually containing male
flowers only, under these circumstances, bears female flowers also.[190] In
like manner, but less frequently, the female inflorescence occasionally
produces male flowers as well.
Among the species of Carex it is a common thing for the terminal spike to
consist of male flowers at the top, and female flowers at the base; the
converse of this, where the female flowers are at the summit of the spike, is
much more uncommon. An illustration of this occurrence is given in the
figure (fig. 100). Among the Coniferæ numerous instances have been
recorded of the presence of male and female flowers on the same spike,
thus Mr. now Professor Alexander Dickson exhibited at the Botanical
Society of Edinburgh in July, 1860, some malformed cones of Abies
excelsa, in which the inferior part of the axis was covered with stamens,
structural completeness rather than the reverse. It must be also understood
that the following remarks apply to structural points only, and are not
intended to include the question of function. The occurrence of
heteromorphic unions renders it necessary to keep in mind that plants
hermaphrodite as to structure are by no means necessarily so as to function.
The simplest case of this alteration in the relative position of the sexes is
that which occurs in monœcious plants, where the male and female flowers
have a definite position, but which in exceptional instances is altered.
Change in the relative position of male and female flowers may thus
occur in any monœcious plant. Cultivated maize, Zea Mays, frequently
exhibits alterations of this kind; under ordinary circumstances, the male
inflorescence is a compound spike, occupying the extremity of the stem,
while the female flowers are borne in simple spikes at a lower level, but
specimens may now and then be found where the sexes are mixed in the
same inflorescence; the upper branching panicle usually containing male
flowers only, under these circumstances, bears female flowers also.[190] In
like manner, but less frequently, the female inflorescence occasionally
produces male flowers as well.
Among the species of Carex it is a common thing for the terminal spike to
consist of male flowers at the top, and female flowers at the base; the
converse of this, where the female flowers are at the summit of the spike, is
much more uncommon. An illustration of this occurrence is given in the
figure (fig. 100). Among the Coniferæ numerous instances have been
recorded of the presence of male and female flowers on the same spike,
thus Mr. now Professor Alexander Dickson exhibited at the Botanical
Society of Edinburgh in July, 1860, some malformed cones of Abies
excelsa, in which the inferior part of the axis was covered with stamens,
Page 257
whilst the terminal portion produced bracts and scales like an ordinary
female cone. The stamens of the lower division were serially continuous
with the bracts above. Some of the lower scales of the female portion were
in the axils of the uppermost stamens, which last were somewhat modified,
the anther cells being diminished, whilst the scale-like crest had become
more elongated and pointed, in fact, more or less resembling the ordinary
bracts.[191] Mohl, Schleiden, and A. Braun have observed similar cones in
Pinus alba, and Cramer figures and describes androgynous cones in Larix
microcarpa. C. A. Meyer ('Bull. Phys. Math.,' t. x, 1850) also describes
some catkins of Alnus fruticosa which bore male flowers at the top, and
female flowers at the base.
Fig. 100.—Spike of Carex acuta, with female flowers at the summit.
On the subject of this section the reader may consult A. Braun,. 'Das Individ.,' 1853, p. 65.
Caspary, 'De Abietin. flor. fem. struct. morphol.' Schleiden. 'Principles,' English edition, p.
299. Mohl, 'Verm. Schrift.,' p. 45. Meyen in 'Wiegm. Archiv.,' 1838, p. 155. Cramer,
female cone. The stamens of the lower division were serially continuous
with the bracts above. Some of the lower scales of the female portion were
in the axils of the uppermost stamens, which last were somewhat modified,
the anther cells being diminished, whilst the scale-like crest had become
more elongated and pointed, in fact, more or less resembling the ordinary
bracts.[191] Mohl, Schleiden, and A. Braun have observed similar cones in
Pinus alba, and Cramer figures and describes androgynous cones in Larix
microcarpa. C. A. Meyer ('Bull. Phys. Math.,' t. x, 1850) also describes
some catkins of Alnus fruticosa which bore male flowers at the top, and
female flowers at the base.
Fig. 100.—Spike of Carex acuta, with female flowers at the summit.
On the subject of this section the reader may consult A. Braun,. 'Das Individ.,' 1853, p. 65.
Caspary, 'De Abietin. flor. fem. struct. morphol.' Schleiden. 'Principles,' English edition, p.
299. Mohl, 'Verm. Schrift.,' p. 45. Meyen in 'Wiegm. Archiv.,' 1838, p. 155. Cramer,
Page 258
'Bildungsabweich,' p. 4, tab. v, figs. 13–17. Parlatore, 'Ann. Sc. Nat.,' ser. iv, vol. xvi, p.
215, tab. 13a. See also under the head of Prolification, Substitutions, &c.
Change from the monœcious to the diœcious condition.—This is of less
frequent occurrence than might have been anticipated. In the 'Gardeners'
Chronicle,' 1847, pp. 541 and 558, several instances are noted of walnut
trees bearing female flowers to the exclusion of males. The mulberry tree
has also been noticed to produce female blossoms only, while in other
plants male flowers only are developed.
It seems probable that the age of the plant may have something to do with
this production of flowers of one sex to the exclusion of the other.
Change from the diœcious to the monœcious condition.—Androgynism.
—This is of far more common occurrence than the preceding.
Fig. 101.—Monœcious inflorescence of Hop.
215, tab. 13a. See also under the head of Prolification, Substitutions, &c.
Change from the monœcious to the diœcious condition.—This is of less
frequent occurrence than might have been anticipated. In the 'Gardeners'
Chronicle,' 1847, pp. 541 and 558, several instances are noted of walnut
trees bearing female flowers to the exclusion of males. The mulberry tree
has also been noticed to produce female blossoms only, while in other
plants male flowers only are developed.
It seems probable that the age of the plant may have something to do with
this production of flowers of one sex to the exclusion of the other.
Change from the diœcious to the monœcious condition.—Androgynism.
—This is of far more common occurrence than the preceding.
Fig. 101.—Monœcious inflorescence of Hop.
Page 259
In the hop (Humulus Lupulus), when monœcious, the female catkins are
usually borne on the ends of the branches as shown in the cut (fig. 101), and
a similar thing has been noticed in Urtica dioica by Clos, 'Bull. Soc. Bot.
France,' vol. 9, p. 7.
Baillon ('Etudes du groupe des Euphorbiacées,' p. 205) mentions the
following species of that order as having been seen by him with monœcious
inflorescence: Schismatopera distichophylla, Mozinna peltata, Hermesia
castaneifolia. Oliver mentions ('Hook. Icon. Plant.,' t. 1044) that in
Leitneria floridana the upper scales of the male catkin occasionally subtend
an ovary.
It would seem that external conditions have some effect in determining the
formation of one sex, as in some species of Carex, while in the case of Salix
repens, Hampe[192] says that when grown partially or for a time under
water, those twigs which are thrust up above the surface bear female
flowers, while those twigs that blossom after the water is dried up, produce
male flowers only.
Carrière[193] says that a plant of Stauntonia latifolia which for some years
produced stamens only, now produces flowers of both sexes; it was
diœcious, but is now monœcious. The same author alludes to a similar
occurrence in Juniperus Virginiana. The hops is also said to vary in sexual
characteristics from time to time.[194] In addition to the genera, already
named, in which this production of flowers of both sexes has been observed
may be mentioned Taxus! Gunnera! Urtica! Mercurialis! Restio! Cannabis!
Salix! Humulus! as well as others in which the change is less frequent.
Among cryptogams a similar change occurs. As an illustration may be cited
Leucobryum giganteum, as quoted from Müller in Henfrey's 'Botanical
Gazette,' i, p. 100.
usually borne on the ends of the branches as shown in the cut (fig. 101), and
a similar thing has been noticed in Urtica dioica by Clos, 'Bull. Soc. Bot.
France,' vol. 9, p. 7.
Baillon ('Etudes du groupe des Euphorbiacées,' p. 205) mentions the
following species of that order as having been seen by him with monœcious
inflorescence: Schismatopera distichophylla, Mozinna peltata, Hermesia
castaneifolia. Oliver mentions ('Hook. Icon. Plant.,' t. 1044) that in
Leitneria floridana the upper scales of the male catkin occasionally subtend
an ovary.
It would seem that external conditions have some effect in determining the
formation of one sex, as in some species of Carex, while in the case of Salix
repens, Hampe[192] says that when grown partially or for a time under
water, those twigs which are thrust up above the surface bear female
flowers, while those twigs that blossom after the water is dried up, produce
male flowers only.
Carrière[193] says that a plant of Stauntonia latifolia which for some years
produced stamens only, now produces flowers of both sexes; it was
diœcious, but is now monœcious. The same author alludes to a similar
occurrence in Juniperus Virginiana. The hops is also said to vary in sexual
characteristics from time to time.[194] In addition to the genera, already
named, in which this production of flowers of both sexes has been observed
may be mentioned Taxus! Gunnera! Urtica! Mercurialis! Restio! Cannabis!
Salix! Humulus! as well as others in which the change is less frequent.
Among cryptogams a similar change occurs. As an illustration may be cited
Leucobryum giganteum, as quoted from Müller in Henfrey's 'Botanical
Gazette,' i, p. 100.
Page 260
As to androgynous willows, in addition to the references given under the head of
Substitution of stamens for pistils, see Schlechtendal, 'Flora Berol.,' ii. p. 259. Tausch, 'Bot.
Zeit.,' 1833, i. p. 229. Koch, 'Synops. Flor. Germ.,' 740. Host, 'Flor. Aust,.' ii, p. 641 (S.
mirabilis). See also Hegelmaier, 'Württemberg Naturwissenshaft Jahreshefte,' 1866, p. 30.
Other references to less accessible works are given in 'Linnæa,' xiv, p. 372.
Change from hermaphroditism to unisexuality.—Many flowers
ordinarily hermaphrodite as to structure, become unisexual by the abortion
or suppression of their stamens, or of their carpels, as the case may be. This
phenomenon is lessened in interest since the demonstration of the fact by
Darwin and others, that many plants, structurally hermaphrodite, require for
the full and perfect performance of their functions the cooperation of the
stamens and pistils, belonging to different individuals of the same species.
Some of the Ranunculaceæ constantly exhibit a tendency towards the
diœcious condition, and the rarity with which perfect seeds of Ranunculus
Ficaria are formed is to be attributed, in great measure, to the deficiency of
pollen in the anthers of these flowers. Ranunculus auricomus also is
frequently sterile. Specimens of Ranunculus bulbosus may be met with in
which every flower is furnished with carpels, most of which have evidently
been fertilised, although there are no perfect stamens in the flowers.
Knight and other vegetable physiologists have been of opinion that a high
temperature favours the production of stamens, while a lower degree of heat
is considered more favorable to the production of pistils, and in this way the
occurrence of "blind" strawberries has been accounted for. Mr. R.
Thompson, writing on this subject, speaks of a plantation of Hautbois
strawberries which in one season were wholly sterile, and accounts for the
circumstance as follows: the plants were taken from the bearing beds the
year previous, and were planted in a rich well-manured border, in which
they started rapidly into too great luxuriance, the growth being to leaves
rather than to fruit. The following season these same plants bore a most
Substitution of stamens for pistils, see Schlechtendal, 'Flora Berol.,' ii. p. 259. Tausch, 'Bot.
Zeit.,' 1833, i. p. 229. Koch, 'Synops. Flor. Germ.,' 740. Host, 'Flor. Aust,.' ii, p. 641 (S.
mirabilis). See also Hegelmaier, 'Württemberg Naturwissenshaft Jahreshefte,' 1866, p. 30.
Other references to less accessible works are given in 'Linnæa,' xiv, p. 372.
Change from hermaphroditism to unisexuality.—Many flowers
ordinarily hermaphrodite as to structure, become unisexual by the abortion
or suppression of their stamens, or of their carpels, as the case may be. This
phenomenon is lessened in interest since the demonstration of the fact by
Darwin and others, that many plants, structurally hermaphrodite, require for
the full and perfect performance of their functions the cooperation of the
stamens and pistils, belonging to different individuals of the same species.
Some of the Ranunculaceæ constantly exhibit a tendency towards the
diœcious condition, and the rarity with which perfect seeds of Ranunculus
Ficaria are formed is to be attributed, in great measure, to the deficiency of
pollen in the anthers of these flowers. Ranunculus auricomus also is
frequently sterile. Specimens of Ranunculus bulbosus may be met with in
which every flower is furnished with carpels, most of which have evidently
been fertilised, although there are no perfect stamens in the flowers.
Knight and other vegetable physiologists have been of opinion that a high
temperature favours the production of stamens, while a lower degree of heat
is considered more favorable to the production of pistils, and in this way the
occurrence of "blind" strawberries has been accounted for. Mr. R.
Thompson, writing on this subject, speaks of a plantation of Hautbois
strawberries which in one season were wholly sterile, and accounts for the
circumstance as follows: the plants were taken from the bearing beds the
year previous, and were planted in a rich well-manured border, in which
they started rapidly into too great luxuriance, the growth being to leaves
rather than to fruit. The following season these same plants bore a most
Page 261
abundant crop, hence these plants were accidentally prevented from
perfecting their female organs.[195]
Mr. Darwin[196] cites from various sources the following details relating to
strawberries which it may be useful to insert in this place, as throwing some
light upon the production of unisexual flowers. "Several English varieties,
which in this country are free from any such tendency, when cultivated in
rich soils under the climate of North America commonly produce plants
with separate sexes. Thus, a whole acre of Keen's seedlings in the United
States has been observed to be almost sterile in the absence of male flowers;
but the more general rule is, that the male plants over-run the females....
The most successful cultivators in Ohio plant, for every seven rows of
pistillate flowers, one row of hermaphrodites, which afford pollen for both
kinds; but the hermaphrodites, owing to their expenditure in the production
of pollen, bear less fruit than the female plants."
Stratiotes aloides has been said to produce its carpels with greater
abundance towards the northern limits of its geographical distribution, and
its stamens, on the other hand, are stated to be more frequently developed in
more southern districts.
Honckenya peploides affords another illustration of the sexual arrangements
in the flower being altered as it would seem by climatal conditions. Thus, in
the United States, according to Professor Asa Gray, the flowers are
frequently hermaphrodite, while in this country they are usually sub-
diœcious.[197]
Treviranus[198] says that the flowers of Hippuris and Callitriche are apt to
be hermaphrodite in summer, but female only at a later period.
For further remarks on this subject, see sections relating to suppression of
stamens and pistils.
perfecting their female organs.[195]
Mr. Darwin[196] cites from various sources the following details relating to
strawberries which it may be useful to insert in this place, as throwing some
light upon the production of unisexual flowers. "Several English varieties,
which in this country are free from any such tendency, when cultivated in
rich soils under the climate of North America commonly produce plants
with separate sexes. Thus, a whole acre of Keen's seedlings in the United
States has been observed to be almost sterile in the absence of male flowers;
but the more general rule is, that the male plants over-run the females....
The most successful cultivators in Ohio plant, for every seven rows of
pistillate flowers, one row of hermaphrodites, which afford pollen for both
kinds; but the hermaphrodites, owing to their expenditure in the production
of pollen, bear less fruit than the female plants."
Stratiotes aloides has been said to produce its carpels with greater
abundance towards the northern limits of its geographical distribution, and
its stamens, on the other hand, are stated to be more frequently developed in
more southern districts.
Honckenya peploides affords another illustration of the sexual arrangements
in the flower being altered as it would seem by climatal conditions. Thus, in
the United States, according to Professor Asa Gray, the flowers are
frequently hermaphrodite, while in this country they are usually sub-
diœcious.[197]
Treviranus[198] says that the flowers of Hippuris and Callitriche are apt to
be hermaphrodite in summer, but female only at a later period.
For further remarks on this subject, see sections relating to suppression of
stamens and pistils.
Page 262
Change from unisexuality to hermaphroditism.—This occurrence
depends on one of two causes, either organs are developed (stamens or
pistils as the case may be), which are habitually absent in the particular
flower; or some of the stamens may be more or less completely converted
into or replaced by pistils, or vice versâ.
The first condition is the opposite of suppression; it is, as it were, a
restoration of symmetry, and might be included under the head of regular
peloria, inasmuch as certain organs which habitually undergo suppression at
a certain stage in their development, by exception, go on growing, and
produce a perfect, instead of an imperfect flower. In teratological records it
is not always stated clearly to which of the two above-named causes the
unusual hermaphroditism belongs, though it is generally easy to ascertain
this point. Very many, perhaps all, diclinous flowers may, under certain
conditions, become perfect, at least structurally. I have myself seen
hermaphrodite flowers in Cucurbita,[199] Mercurialis, Cannabis, Zea Mays,
and Aucuba japonica, as well as in many Restiaceæ, notably Cannamois
virgata and Lepyrodia hermaphrodita. Spinacia oleracea, Rhodiola rosea,
Cachrys taurica, and Empetrum nigrum are also occasionally
hermaphrodite.
Gubler[200] alludes to a similar occurrence in Pistacia Lentiscus, wherein,
however, he adds that there was a deficiency of pollen in the flowers.
Schnizlein[201] observed hermaphrodite flowers in the beech, Fagus
sylvatica, the ovaries being smaller than usual, and the stamens epigynous.
Baillon[202] enumerates the following Euphorbiaceæ as having
exceptionally produced hermaphrodite flowers, Crozophora tinctoria,
Suregada sp., Phyllanthus longifolius, Breynia sp., Philyra brasiliensis,
depends on one of two causes, either organs are developed (stamens or
pistils as the case may be), which are habitually absent in the particular
flower; or some of the stamens may be more or less completely converted
into or replaced by pistils, or vice versâ.
The first condition is the opposite of suppression; it is, as it were, a
restoration of symmetry, and might be included under the head of regular
peloria, inasmuch as certain organs which habitually undergo suppression at
a certain stage in their development, by exception, go on growing, and
produce a perfect, instead of an imperfect flower. In teratological records it
is not always stated clearly to which of the two above-named causes the
unusual hermaphroditism belongs, though it is generally easy to ascertain
this point. Very many, perhaps all, diclinous flowers may, under certain
conditions, become perfect, at least structurally. I have myself seen
hermaphrodite flowers in Cucurbita,[199] Mercurialis, Cannabis, Zea Mays,
and Aucuba japonica, as well as in many Restiaceæ, notably Cannamois
virgata and Lepyrodia hermaphrodita. Spinacia oleracea, Rhodiola rosea,
Cachrys taurica, and Empetrum nigrum are also occasionally
hermaphrodite.
Gubler[200] alludes to a similar occurrence in Pistacia Lentiscus, wherein,
however, he adds that there was a deficiency of pollen in the flowers.
Schnizlein[201] observed hermaphrodite flowers in the beech, Fagus
sylvatica, the ovaries being smaller than usual, and the stamens epigynous.
Baillon[202] enumerates the following Euphorbiaceæ as having
exceptionally produced hermaphrodite flowers, Crozophora tinctoria,
Suregada sp., Phyllanthus longifolius, Breynia sp., Philyra brasiliensis,
Page 263
Ricinus communis, Conceveiba macrophylla, Cluytia semperflorens, Wall,
non Roxb. Mercurialis annua and Cleistanthus polystachyus.
In some of these cases the hermaphroditism is due to the development of
anthers on the usually barren staminodes, though, in other cases, the
stamens would seem to be separate, independent formations, as they do not
occupy the same relative position that the ordinary stamens would do if
developed.[203]
Fig. 102.—Flower of Fuchsia in which the calyx was leafy, the petals normal (reflexed in the figure),
the stamens partially converted into ovaries, the ordinary inferior ovary being absent. See
Substitution.
Robert Brown[204] observed stamens within the utricle of Carex acuta, and
Gay is stated by Moquin ('El. Ter. Veg.,' p. 343) to have observed a similar
occurrence in Carex glauca.
non Roxb. Mercurialis annua and Cleistanthus polystachyus.
In some of these cases the hermaphroditism is due to the development of
anthers on the usually barren staminodes, though, in other cases, the
stamens would seem to be separate, independent formations, as they do not
occupy the same relative position that the ordinary stamens would do if
developed.[203]
Fig. 102.—Flower of Fuchsia in which the calyx was leafy, the petals normal (reflexed in the figure),
the stamens partially converted into ovaries, the ordinary inferior ovary being absent. See
Substitution.
Robert Brown[204] observed stamens within the utricle of Carex acuta, and
Gay is stated by Moquin ('El. Ter. Veg.,' p. 343) to have observed a similar
occurrence in Carex glauca.
Page 264
Paasch[205] observed a similar occurrence in C. cæspitosa, and Schauer, in
C. paludosa,[206] though in the latter instance the case seems to have been
one of transformation or substitution rather than one of hermaphroditism.
The second cause of this pseudo-hermaphroditism is due either to the more
or less perfect mutation of male and female organs, or it may be to the
complete absence of one and its replacement by another, as when out of
many stamens, one or more are deficient, and their places occupied by
carpels. This happens very frequently in willows and poplars, and has been
seen in the beech.[207]
Fig. 103.—Hermaphrodite flower of Carica Papaya.
In Begonia frigida[208] the anomaly is increased by the position of the
ovaries above, the perianth, a position due, not to any solution or
detachment of the latter from the former, but simply to the presence of
ovaries where, under ordinary circumstances, stamens only are formed, as
C. paludosa,[206] though in the latter instance the case seems to have been
one of transformation or substitution rather than one of hermaphroditism.
The second cause of this pseudo-hermaphroditism is due either to the more
or less perfect mutation of male and female organs, or it may be to the
complete absence of one and its replacement by another, as when out of
many stamens, one or more are deficient, and their places occupied by
carpels. This happens very frequently in willows and poplars, and has been
seen in the beech.[207]
Fig. 103.—Hermaphrodite flower of Carica Papaya.
In Begonia frigida[208] the anomaly is increased by the position of the
ovaries above, the perianth, a position due, not to any solution or
detachment of the latter from the former, but simply to the presence of
ovaries where, under ordinary circumstances, stamens only are formed, as
Page 265
happened also in a garden variety of a Fuchsia, wherein, however, the
change was less perfect than in the Begonia, and in which, as the flower is
naturally hermaphrodite, the alteration is of the less importance.
Fig. 104.—Ovuliferous anthers—Cucurbita.
In hermaphrodite flowers of Carica Papaya (fig. 103) there is a single row
of five stamens instead of two rows of five each as in the normal male
flowers, the position of the second or inner row of stamens being occupied
by five carpels, which, however, are not adherent to the corolla as the
stamens are, thus, supposing the arrangement of parts in the normal male
flowers to be as follows:
---------------------------
S S S S S
---------------------------
| p p p p p
|
change was less perfect than in the Begonia, and in which, as the flower is
naturally hermaphrodite, the alteration is of the less importance.
Fig. 104.—Ovuliferous anthers—Cucurbita.
In hermaphrodite flowers of Carica Papaya (fig. 103) there is a single row
of five stamens instead of two rows of five each as in the normal male
flowers, the position of the second or inner row of stamens being occupied
by five carpels, which, however, are not adherent to the corolla as the
stamens are, thus, supposing the arrangement of parts in the normal male
flowers to be as follows:
---------------------------
S S S S S
---------------------------
| p p p p p
|
Page 266
| st st st st st
|
| st st st st st
|
That of the hermaphrodite blossoms would be, in brief, as follows:
| 5 S
|------------
| 5 p
|
| 5 st
|
| 5 c
|
One of the most curious cases of this kind recorded is one mentioned by Mr.
Berkeley,[209] wherein a large white-seeded gourd presented a majority of
flowers in which the pollen was replaced by ovules. It would seem probable
from the appearances presented by the figure that these ovules were, some
of them, polliniferous, like those of the Passiflora, &c., described at p. 185,
but nothing is stated on the subject.
See also section on Regular Peloria, Substitution, Pistillody of the stamens,
&c.
FOOTNOTES:
[190] See also Clos., 'Mem. Acad. Toulouse,' sixth ser., t. iii, pp. 294–305. Scott, 'Trans. Bot.
Soc. Edinburgh,' t. viii, p. 60. Wigand, 'Flora,' 1856, p. 707.
[191] Professor Dickson concludes from the examination of these structures that the male
cone, consisting of simple stamens developed on one common axis, must be regarded as a
simple male flower, while the axillary scales of the female cone are by him compared with the
flattened shoots of Ruscus.
[192] 'Linnæa,' xiv, 367.
[193] Rev. Hortic.,' January, 1867.
|
| st st st st st
|
That of the hermaphrodite blossoms would be, in brief, as follows:
| 5 S
|------------
| 5 p
|
| 5 st
|
| 5 c
|
One of the most curious cases of this kind recorded is one mentioned by Mr.
Berkeley,[209] wherein a large white-seeded gourd presented a majority of
flowers in which the pollen was replaced by ovules. It would seem probable
from the appearances presented by the figure that these ovules were, some
of them, polliniferous, like those of the Passiflora, &c., described at p. 185,
but nothing is stated on the subject.
See also section on Regular Peloria, Substitution, Pistillody of the stamens,
&c.
FOOTNOTES:
[190] See also Clos., 'Mem. Acad. Toulouse,' sixth ser., t. iii, pp. 294–305. Scott, 'Trans. Bot.
Soc. Edinburgh,' t. viii, p. 60. Wigand, 'Flora,' 1856, p. 707.
[191] Professor Dickson concludes from the examination of these structures that the male
cone, consisting of simple stamens developed on one common axis, must be regarded as a
simple male flower, while the axillary scales of the female cone are by him compared with the
flattened shoots of Ruscus.
[192] 'Linnæa,' xiv, 367.
[193] Rev. Hortic.,' January, 1867.
Page 267
[194] See Royle, 'Man. Materia Medica,' ed. 1, p. 567.
[195] Thomson, 'Gardener's Assistant,' p. 577.
[196] 'Variation of Animals and Plants,' i, 353.
[197] Babington, 'Ann. Nat. Hist.,' vol. ix, 1852, p. 156.
[198] 'Phys. der Gewächse,' ii, p. 323.
[199] See also Schlechtendal, 'Linnæa,' viii, p. 623, and Lindley, 'Veg. Kingd.,' p. 315.
[200] 'Bull. Soc. Bot. France,' vol. ix, p. 81.
[201] Cited in Henfrey, 'Bot. Gazette.' 3, p. 11.
[202] Baillon. 'Etudes du Groupe des Euphorbiacées,' p. 205, tab. xv, fig. 19, tab. xix, fig. 31.
[203] See also Guillemin, 'Mém. Soc. Nat. Hist. Paris,' I, p. 16; hermaphrodite flowers in
Euphorbia esula.
[204] 'Prod. Flor. N. Holl.,' p. 242.
[205] 'Bot. Zeit.,' 1837, p. 335.
[206] 'Pflanz, Terat.,' von Moquin-Tandon, p. 208.
[207] Schnizlein, loc. cit.
[208] 'Bot. Mag.,' tab. 5160, fig. 4. See also 'Gard. Chron.,' 1860, pp. 146, 170; 1861, p.
1092.
[209] 'Gard. Chron.,' 1851, p. 499.
CHAPTER V.
ALTERATIONS IN THE DIRECTION OF ORGANS.
The deviations from the ordinary direction of organs partake for the most
part more of the nature of variations than of absolute malposition or
displacement. It must also be borne in mind how frequently the direction of
[195] Thomson, 'Gardener's Assistant,' p. 577.
[196] 'Variation of Animals and Plants,' i, 353.
[197] Babington, 'Ann. Nat. Hist.,' vol. ix, 1852, p. 156.
[198] 'Phys. der Gewächse,' ii, p. 323.
[199] See also Schlechtendal, 'Linnæa,' viii, p. 623, and Lindley, 'Veg. Kingd.,' p. 315.
[200] 'Bull. Soc. Bot. France,' vol. ix, p. 81.
[201] Cited in Henfrey, 'Bot. Gazette.' 3, p. 11.
[202] Baillon. 'Etudes du Groupe des Euphorbiacées,' p. 205, tab. xv, fig. 19, tab. xix, fig. 31.
[203] See also Guillemin, 'Mém. Soc. Nat. Hist. Paris,' I, p. 16; hermaphrodite flowers in
Euphorbia esula.
[204] 'Prod. Flor. N. Holl.,' p. 242.
[205] 'Bot. Zeit.,' 1837, p. 335.
[206] 'Pflanz, Terat.,' von Moquin-Tandon, p. 208.
[207] Schnizlein, loc. cit.
[208] 'Bot. Mag.,' tab. 5160, fig. 4. See also 'Gard. Chron.,' 1860, pp. 146, 170; 1861, p.
1092.
[209] 'Gard. Chron.,' 1851, p. 499.
CHAPTER V.
ALTERATIONS IN THE DIRECTION OF ORGANS.
The deviations from the ordinary direction of organs partake for the most
part more of the nature of variations than of absolute malposition or
displacement. It must also be borne in mind how frequently the direction of
Page 268
the leaves, or of the flower, varies according to the stage of development
which it has arrived at, to unequal or disproportionate growth of some parts,
or to the presence of some impediment either accidental or resulting from
the natural growth of the plant. These and other causes tend to alter the
direction of parts very materially.
Change in the direction of axile organs, roots, stems, &c.—The roots
frequently exhibit good illustrations of the effect of the causes above
mentioned in altering the natural direction. The roots are put out of their
course by meeting with any obstacle in their way. Almost the only
exception to the rule in accordance with which roots descend under natural
circumstances, is that furnished by Trapa natans, the roots of which in
germination are directed upwards towards the surface of the water. So in
Sechium edule, the seed of which germinates while still in the fruit, the
roots are necessarily, owing to the inverted position of the embryo, directed
upwards in the first instance.
A downward direction of the stem or branches occurs in many weak-
stemmed plants growing upon rocks or walls, or in trees with very long
slender branches as in Salix Babylonica, and the condition may often be
produced artificially as in the weeping ash.
The opposite change occurs in what are termed fastigiate varieties, where
the branches, in place of assuming more or less of a horizontal direction,
become erect and nearly parallel with the main stem as in the Lombardy
poplar, which is supposed to be merely a form of the black Italian poplar.
M. de Selys-Longchamps has described a similar occurrence in another
species of Poplar (P. virginiana Desf.), and amongst a number of seedling
plants fastigiate varieties may frequently be found, which may be
perpetuated by cuttings or grafts, or sometimes even by seed; hence the
which it has arrived at, to unequal or disproportionate growth of some parts,
or to the presence of some impediment either accidental or resulting from
the natural growth of the plant. These and other causes tend to alter the
direction of parts very materially.
Change in the direction of axile organs, roots, stems, &c.—The roots
frequently exhibit good illustrations of the effect of the causes above
mentioned in altering the natural direction. The roots are put out of their
course by meeting with any obstacle in their way. Almost the only
exception to the rule in accordance with which roots descend under natural
circumstances, is that furnished by Trapa natans, the roots of which in
germination are directed upwards towards the surface of the water. So in
Sechium edule, the seed of which germinates while still in the fruit, the
roots are necessarily, owing to the inverted position of the embryo, directed
upwards in the first instance.
A downward direction of the stem or branches occurs in many weak-
stemmed plants growing upon rocks or walls, or in trees with very long
slender branches as in Salix Babylonica, and the condition may often be
produced artificially as in the weeping ash.
The opposite change occurs in what are termed fastigiate varieties, where
the branches, in place of assuming more or less of a horizontal direction,
become erect and nearly parallel with the main stem as in the Lombardy
poplar, which is supposed to be merely a form of the black Italian poplar.
M. de Selys-Longchamps has described a similar occurrence in another
species of Poplar (P. virginiana Desf.), and amongst a number of seedling
plants fastigiate varieties may frequently be found, which may be
perpetuated by cuttings or grafts, or sometimes even by seed; hence the
Page 269
origin of fastigiate varieties of elms, oaks, thorns, chestnuts, and other
plants which may be met with in the nurseries.
Sometimes when the top of the main stem is destroyed by disease or
accident, one of the heretofore lateral shoots takes its place, and continues
the development of the tree in the original direction. It is often an object
with the gardener to restore the symmetry of an injured tree so that its
beauty may ultimately not be impaired.[210]
Climate appears sometimes to have some influence on the direction of
branches, thus Dr. Falconer, as quoted by Darwin,[211] relates that in the
hotter parts of India "the English Ribston-pippin apple, a Himalayan oak, a
Prunus and a Pyrus all assume a fastigiate or pyramidal habit, and this fact
is the more interesting as a Chinese tropical species of Pyrus naturally has
this habit of growth. Nevertheless many of the fastigiate varieties seen in
gardens have originated in this country by variation of seeds or buds."
M. Carrière has also recorded a curious circumstance with reference to the
fastigiate variety of the false acacia Robinia pseudacacia; he states that if a
cutting or a graft be taken from the upper portion of the tree, the fastigiate
habit will be reproduced, and the branches will be furrowed and covered
with short prickles; but if the plant be multiplied by detaching portions of
the root-stock, then instead of getting a pyramidal tree with erect branches,
a spreading bushy shrub is produced, with more or less horizontal,
cylindrical branches, destitute of prickles.[212]
Eversion of the axis.—In the case of the fig, the peculiar inflorescence is
usually explained on the supposition that the termination of the axis
becomes concave, during growth, bearing the true flowers in the hollow
thus formed. The cavity in this case would probably be due not to any real
process of excavation, but to a disproportionate growth of the outer as
plants which may be met with in the nurseries.
Sometimes when the top of the main stem is destroyed by disease or
accident, one of the heretofore lateral shoots takes its place, and continues
the development of the tree in the original direction. It is often an object
with the gardener to restore the symmetry of an injured tree so that its
beauty may ultimately not be impaired.[210]
Climate appears sometimes to have some influence on the direction of
branches, thus Dr. Falconer, as quoted by Darwin,[211] relates that in the
hotter parts of India "the English Ribston-pippin apple, a Himalayan oak, a
Prunus and a Pyrus all assume a fastigiate or pyramidal habit, and this fact
is the more interesting as a Chinese tropical species of Pyrus naturally has
this habit of growth. Nevertheless many of the fastigiate varieties seen in
gardens have originated in this country by variation of seeds or buds."
M. Carrière has also recorded a curious circumstance with reference to the
fastigiate variety of the false acacia Robinia pseudacacia; he states that if a
cutting or a graft be taken from the upper portion of the tree, the fastigiate
habit will be reproduced, and the branches will be furrowed and covered
with short prickles; but if the plant be multiplied by detaching portions of
the root-stock, then instead of getting a pyramidal tree with erect branches,
a spreading bushy shrub is produced, with more or less horizontal,
cylindrical branches, destitute of prickles.[212]
Eversion of the axis.—In the case of the fig, the peculiar inflorescence is
usually explained on the supposition that the termination of the axis
becomes concave, during growth, bearing the true flowers in the hollow
thus formed. The cavity in this case would probably be due not to any real
process of excavation, but to a disproportionate growth of the outer as
Page 270
contrasted with the central parts of the fig. Some species of Sempervivum
have a similar mode of growth, so that ultimately a kind of tube is formed,
lined by the leaves, the central and innermost being the youngest. The hip
of the Rose may be explained in a similar manner by the greater
proportionate growth of the outer as contrasted with the central portions of
the apex of the flower-stalk. In cases of median prolification, already
referred to, the process is reversed, the central portions then elongate into a
shoot and no cavity is formed. A fig observed by Zuccarini (figs. 105, 106)
appears to have been formed in a similar manner, the flower-bearing
summit of the stalk not being contracted as usual, the flowers projected
beyond the orifice of the fig. If this view be correct the case would be one
rather of lengthening of the axis than of absolute eversion since it was never
inverted.
Fig. 105.—Fig showing prolonged inflorescence and projecting flowers.
have a similar mode of growth, so that ultimately a kind of tube is formed,
lined by the leaves, the central and innermost being the youngest. The hip
of the Rose may be explained in a similar manner by the greater
proportionate growth of the outer as contrasted with the central portions of
the apex of the flower-stalk. In cases of median prolification, already
referred to, the process is reversed, the central portions then elongate into a
shoot and no cavity is formed. A fig observed by Zuccarini (figs. 105, 106)
appears to have been formed in a similar manner, the flower-bearing
summit of the stalk not being contracted as usual, the flowers projected
beyond the orifice of the fig. If this view be correct the case would be one
rather of lengthening of the axis than of absolute eversion since it was never
inverted.
Fig. 105.—Fig showing prolonged inflorescence and projecting flowers.
Page 271
Fig. 106.—Section of the same.
Altered direction of leaves.—The leaves partake more or less of the
altered direction of the axis, as in fastigiate elms, but this is not universally
the case, for though the stem is bent downwards the leaves may be placed in
the opposite direction; thus in some specimens of Galium Aparine growing
on the side of a cliff from which there had been a fall of chalk, the stems,
owing apparently to the landslip, were pendent, but the leaves were abruptly
bent upwards.
One of the most singular instances of an inverted direction of the leaves is
that presented by a turnip (fig. 107) presented to the Museum of King's
College, London, by the late Professor Edward Forbes. The turnip is hollow
in the interior and the majority of the leaves springing from its apex instead
of ascending into the light and air become bent downwards so as to occupy
the cavity, and in such a manner as to bring to mind the position of an
inverted embryo in a seed.
Altered direction of leaves.—The leaves partake more or less of the
altered direction of the axis, as in fastigiate elms, but this is not universally
the case, for though the stem is bent downwards the leaves may be placed in
the opposite direction; thus in some specimens of Galium Aparine growing
on the side of a cliff from which there had been a fall of chalk, the stems,
owing apparently to the landslip, were pendent, but the leaves were abruptly
bent upwards.
One of the most singular instances of an inverted direction of the leaves is
that presented by a turnip (fig. 107) presented to the Museum of King's
College, London, by the late Professor Edward Forbes. The turnip is hollow
in the interior and the majority of the leaves springing from its apex instead
of ascending into the light and air become bent downwards so as to occupy
the cavity, and in such a manner as to bring to mind the position of an
inverted embryo in a seed.
Page 272
Fig. 107.—Hollow turnip, showing some of the leaves inverted and occupying the cavity.
Altered direction of the flower and its parts.—The changes which take
place in the relative position either of the flower as a whole or of its several
parts during growth are well known, as also are the relations which some of
these movements bear to the process of fertilisation, so that but little space
need here be given to the subject beyond what is necessary to point out the
frequent changes of direction which necessarily accompany various
deviations from the ordinary form and arrangement of parts.
In cases where an habitually irregular flower becomes regular, the change in
form is frequently associated with an alteration in direction both of the
flower as a whole and, to a greater or less extent, of its individual members,
for instance of Gloxinia, the normal flowers of which are irregular and
pendent, there is now in common cultivation a peloriate race in which the
flowers are regular in form and erect in position.
Altered direction of the flower and its parts.—The changes which take
place in the relative position either of the flower as a whole or of its several
parts during growth are well known, as also are the relations which some of
these movements bear to the process of fertilisation, so that but little space
need here be given to the subject beyond what is necessary to point out the
frequent changes of direction which necessarily accompany various
deviations from the ordinary form and arrangement of parts.
In cases where an habitually irregular flower becomes regular, the change in
form is frequently associated with an alteration in direction both of the
flower as a whole and, to a greater or less extent, of its individual members,
for instance of Gloxinia, the normal flowers of which are irregular and
pendent, there is now in common cultivation a peloriate race in which the
flowers are regular in form and erect in position.
Page 273
Fig. 108.—Flower of normal Gloxinia.
Fig. 109.—Flower of Gloxinia, erect and regular (regular Peloria).
Fig. 108 shows the usual irregular form of Gloxinia, with which may be
contrasted figs. 109, 110 and 111.
Fig. 109.—Flower of Gloxinia, erect and regular (regular Peloria).
Fig. 108 shows the usual irregular form of Gloxinia, with which may be
contrasted figs. 109, 110 and 111.
Page 274
Fig. 109 shows the regular erect form; fig. 110 the calyx of the same flower;
while in fig. 111 are shown the stamens and style of the two plants
respectively. In the upper figure the style of the peloriate variety is shown
as nearly straight, and the stamens undergo a corresponding change. No
doubt the relative fertility and capacity for impregnation of the two varieties
is affected in proportion to the change of form. The Gloxinia affords an
instance of regular congenital peloria in which the regularity of form and
the erect direction are due to an arrest, not of growth, but of development,
in consequence of which the changes that ordinarily ensue during the
progress of the flower from its juvenile to its fully formed condition do not
take place.
Fig. 110.—Calyx of erect Gloxinia.
while in fig. 111 are shown the stamens and style of the two plants
respectively. In the upper figure the style of the peloriate variety is shown
as nearly straight, and the stamens undergo a corresponding change. No
doubt the relative fertility and capacity for impregnation of the two varieties
is affected in proportion to the change of form. The Gloxinia affords an
instance of regular congenital peloria in which the regularity of form and
the erect direction are due to an arrest, not of growth, but of development,
in consequence of which the changes that ordinarily ensue during the
progress of the flower from its juvenile to its fully formed condition do not
take place.
Fig. 110.—Calyx of erect Gloxinia.
Page 275
Fig. 111.—Stamens of erect regular, and of pendent irregular-flowered Gloxinia.
A similar alteration accompanies this form of peloria in other flowers (see
Peloria). A change in direction may result also from other circumstances
than those just alluded to. Abortion or suppression of organs will induce
such an alteration; thus in a flower of Pelargonium now before me three of
the five carpels, from some cause or other, are abortive and much smaller
than usual, and the style and the beak-like torus are bent downwards
towards the stunted carpels instead of being, as they usually are, straight.
Amongst orchids, where the pedicel of the flower or the ovary is normally
twisted, so that the labellum occupies the anterior or inferior part of the
flower, it frequently happens, in cases of peloria and other changes, that the
primitive position is retained, the twist does not take place, and so with
other resupinate flowers. In Azaleas a curious deflexion of the parts of the
flower may occasionally be met with. Fig. 112 shows an instance of this in
which the corolla, the stamens and the style were abruptly bent downwards:
A similar alteration accompanies this form of peloria in other flowers (see
Peloria). A change in direction may result also from other circumstances
than those just alluded to. Abortion or suppression of organs will induce
such an alteration; thus in a flower of Pelargonium now before me three of
the five carpels, from some cause or other, are abortive and much smaller
than usual, and the style and the beak-like torus are bent downwards
towards the stunted carpels instead of being, as they usually are, straight.
Amongst orchids, where the pedicel of the flower or the ovary is normally
twisted, so that the labellum occupies the anterior or inferior part of the
flower, it frequently happens, in cases of peloria and other changes, that the
primitive position is retained, the twist does not take place, and so with
other resupinate flowers. In Azaleas a curious deflexion of the parts of the
flower may occasionally be met with. Fig. 112 shows an instance of this in
which the corolla, the stamens and the style were abruptly bent downwards:
Page 276
as young flowers of this singular variety have not been examined it is
difficult to form an opinion as to the cause of this variation. In one plant the
change occurred in connection with the suppression of all the flowers but
one in the cluster, or rather the place of the flowers was occupied by an
equal number of leafy shoots.
Fig. 112.—Flower of Azalea, showing the corolla reflected.
Moquin[213] mentions a flower of Rosa alpina in which two of the petals
were erect, while the remaining ones were much larger and expanded
horizontally. The same author quotes from M. Desmoulins the case of a
species of Orobanche, in which a disjunction of the petals constituting the
upper lip took place, thus liberating the style and allowing it to assume a
vertical direction.
difficult to form an opinion as to the cause of this variation. In one plant the
change occurred in connection with the suppression of all the flowers but
one in the cluster, or rather the place of the flowers was occupied by an
equal number of leafy shoots.
Fig. 112.—Flower of Azalea, showing the corolla reflected.
Moquin[213] mentions a flower of Rosa alpina in which two of the petals
were erect, while the remaining ones were much larger and expanded
horizontally. The same author quotes from M. Desmoulins the case of a
species of Orobanche, in which a disjunction of the petals constituting the
upper lip took place, thus liberating the style and allowing it to assume a
vertical direction.
Page 277
Fig. 113.—Flower of Cuphea miniata enlarged, showing protrusion and hypertrophy of an erect
placenta, after Morren.
Fig. 114.—Placenta from the flower shown at fig. 113; the ovary is membranous and torn, the
placenta, erect and ovuliferous, after Morren.
M. Carrière[214] has described an instance wherein two apples were joined
together, a larger and a smaller one; the former was directed away from the
centre of the tree as usual, while the smaller one was pointed in exactly the
placenta, after Morren.
Fig. 114.—Placenta from the flower shown at fig. 113; the ovary is membranous and torn, the
placenta, erect and ovuliferous, after Morren.
M. Carrière[214] has described an instance wherein two apples were joined
together, a larger and a smaller one; the former was directed away from the
centre of the tree as usual, while the smaller one was pointed in exactly the
Page 278
opposite direction. The larger fruit had the customary parchment-like
carpels, the smaller was destitute of them.
Sometimes the direction assumed by one flower as an abnormal occurrence
is the same as that which is proper to an allied species or genus under
natural circumstances; thus flowers of the vine (Vitis) have been met with in
which the petals were spreading like a star (fleurs avalidouires), as in the
genus Cissus.[215]
Morren describes a curious condition in some flowers of Cuphea miniata,
in which the placenta protruded through an orifice in the ovary, and losing
the horizontal direction became erect (figs. 113, 114). A similar occurrence
happened in Lobelia erinus. To this condition the Belgian savant gave the
name of gymnaxony.[216]
FOOTNOTES:
[210] The following details as to the method pursued by Mr. McNab, of the Edinburgh
Botanic Garden, may not be uninteresting in this place. They are from the pen of Mr.
Anderson, and originally appeared in the 'Gardeners' Chronicle.'
"The mode of inducing leaders to proceed from laterals is a matter of comparatively little
concern among the generality of deciduous trees, for they are often provided with subsidiary
branches around the leader, at an angle of elevation scarcely less perpendicular, but the laterals
of all Conifers stand, as nearly as possible, at right angles. Imagine the consternation of most
people when the leader of, say, Picea nobilis, P. Nordmanniana, or P. Lowii is destroyed."
In a specimen of the latter plant the leader had been mischievously destroyed, to remedy which
Mr. McNab adopted means which Mr. Anderson goes on to describe. "Looking from the leader
downward to the first tier of laterals, there appeared to have been a number of adventitious
leaf-buds created, owing to the coronal bud being destroyed. These were allowed to plump up
unmolested until the return of spring, when every one was scarified or rubbed off but the one
nearest the extremity. To assist its development and restrain the action of the numerous
laterals, every one was cut back in autumn, and this restraint upon the sap acted so favorably
upon the incipient leader as to give it the strength and stamina of the original leader, so that
nothing detrimental was evident twelve months after the accident had happened, and only a
carpels, the smaller was destitute of them.
Sometimes the direction assumed by one flower as an abnormal occurrence
is the same as that which is proper to an allied species or genus under
natural circumstances; thus flowers of the vine (Vitis) have been met with in
which the petals were spreading like a star (fleurs avalidouires), as in the
genus Cissus.[215]
Morren describes a curious condition in some flowers of Cuphea miniata,
in which the placenta protruded through an orifice in the ovary, and losing
the horizontal direction became erect (figs. 113, 114). A similar occurrence
happened in Lobelia erinus. To this condition the Belgian savant gave the
name of gymnaxony.[216]
FOOTNOTES:
[210] The following details as to the method pursued by Mr. McNab, of the Edinburgh
Botanic Garden, may not be uninteresting in this place. They are from the pen of Mr.
Anderson, and originally appeared in the 'Gardeners' Chronicle.'
"The mode of inducing leaders to proceed from laterals is a matter of comparatively little
concern among the generality of deciduous trees, for they are often provided with subsidiary
branches around the leader, at an angle of elevation scarcely less perpendicular, but the laterals
of all Conifers stand, as nearly as possible, at right angles. Imagine the consternation of most
people when the leader of, say, Picea nobilis, P. Nordmanniana, or P. Lowii is destroyed."
In a specimen of the latter plant the leader had been mischievously destroyed, to remedy which
Mr. McNab adopted means which Mr. Anderson goes on to describe. "Looking from the leader
downward to the first tier of laterals, there appeared to have been a number of adventitious
leaf-buds created, owing to the coronal bud being destroyed. These were allowed to plump up
unmolested until the return of spring, when every one was scarified or rubbed off but the one
nearest the extremity. To assist its development and restrain the action of the numerous
laterals, every one was cut back in autumn, and this restraint upon the sap acted so favorably
upon the incipient leader as to give it the strength and stamina of the original leader, so that
nothing detrimental was evident twelve months after the accident had happened, and only a
Page 279
practical eye could detect that there had been any mishap at all. This beautifully simple
process saved the baby tree.
"Another example of retrieving lost leaders may be quoted as illustrative of many in similar
circumstances. Pícea Webbiana had its leader completely destroyed down to the first tier of
laterals. There was no such provision left for inducing leaf-buds as was the case with P. Lowii
above referred to. Resort must, therefore, be had to one of the best favoured laterals, but how
is it to be coaxed from the horizontal position of a lateral to the perpendicular position of a
leader? The uninitiated in these matters, and, in fact, practical gardeners generally, would at
once reply, by supporting to a stake with the all-powerful Cuba or bast-matting. But no. A far
simpler method than that, namely, by fore-shortening all the laterals of the upper tier but the
one selected for a leader. Nature becomes the handmaid of art here; for without the slightest
prop the lateral gradually raises itself erect, and takes the place of the lost leader. All that the
operator requires to attend to is the amputation of the laterals until this adventitious fellow has
gained a supremacy. Singular provision in nature this, which, thanks to the undivided attention
of a careful observer, has been fully appreciated and utilized."
[211] 'Variation of Animals and Plants,' ii, p. 277.
[212] Quoted in 'Gard. Chron.,' 1867, p. 654.
[213] Loc. cit., p. 315.
[214] 'Rev. Hortic.,' 1868, p. 110.
[215] Planchon and Marès, 'Ann. Sc. Nat.,' 5 ser., tom. vi, 1866, p. 228, tab. xii.
[216] 'Bull. Acad. Belg.,' xviii, part ii, p. 293.
process saved the baby tree.
"Another example of retrieving lost leaders may be quoted as illustrative of many in similar
circumstances. Pícea Webbiana had its leader completely destroyed down to the first tier of
laterals. There was no such provision left for inducing leaf-buds as was the case with P. Lowii
above referred to. Resort must, therefore, be had to one of the best favoured laterals, but how
is it to be coaxed from the horizontal position of a lateral to the perpendicular position of a
leader? The uninitiated in these matters, and, in fact, practical gardeners generally, would at
once reply, by supporting to a stake with the all-powerful Cuba or bast-matting. But no. A far
simpler method than that, namely, by fore-shortening all the laterals of the upper tier but the
one selected for a leader. Nature becomes the handmaid of art here; for without the slightest
prop the lateral gradually raises itself erect, and takes the place of the lost leader. All that the
operator requires to attend to is the amputation of the laterals until this adventitious fellow has
gained a supremacy. Singular provision in nature this, which, thanks to the undivided attention
of a careful observer, has been fully appreciated and utilized."
[211] 'Variation of Animals and Plants,' ii, p. 277.
[212] Quoted in 'Gard. Chron.,' 1867, p. 654.
[213] Loc. cit., p. 315.
[214] 'Rev. Hortic.,' 1868, p. 110.
[215] Planchon and Marès, 'Ann. Sc. Nat.,' 5 ser., tom. vi, 1866, p. 228, tab. xii.
[216] 'Bull. Acad. Belg.,' xviii, part ii, p. 293.
Page 280
BOOK II.
DEVIATIONS FROM THE ORDINARY FORM
OF ORGANS.
In a morphological point of view the form of the various parts or organs of
plants and the changes to which they are subjected during their
development are only second in importance to the diversities of
arrangement and, indeed, in some cases, do not in any degree hold a second
place.
Taken together, the arrangement, form, and number of the several parts of
the flower, make up what has been termed the symmetry of the flower.[217]
Referring to the assumed standard of comparison, see p. 4, it will be seen
that in the typically regular flower all the various organs are supposed to be
regular in their dimensions and form. At one time it was even supposed that
all flowers, no matter how irregular they subsequently became, began by
being strictly symmetrical or regular, and that subsequent alterations were
produced by inequality of growth or development. The researches of
organogenists have, however, dispelled this idea of unvarying primordial
regularity, by showing that in many cases flowers are irregular from the
very first, that some begin by being irregular, and subsequently become
regular, and even in some cases resume their original condition during the
course of their development.[218] Under these circumstances an artificial
DEVIATIONS FROM THE ORDINARY FORM
OF ORGANS.
In a morphological point of view the form of the various parts or organs of
plants and the changes to which they are subjected during their
development are only second in importance to the diversities of
arrangement and, indeed, in some cases, do not in any degree hold a second
place.
Taken together, the arrangement, form, and number of the several parts of
the flower, make up what has been termed the symmetry of the flower.[217]
Referring to the assumed standard of comparison, see p. 4, it will be seen
that in the typically regular flower all the various organs are supposed to be
regular in their dimensions and form. At one time it was even supposed that
all flowers, no matter how irregular they subsequently became, began by
being strictly symmetrical or regular, and that subsequent alterations were
produced by inequality of growth or development. The researches of
organogenists have, however, dispelled this idea of unvarying primordial
regularity, by showing that in many cases flowers are irregular from the
very first, that some begin by being irregular, and subsequently become
regular, and even in some cases resume their original condition during the
course of their development.[218] Under these circumstances an artificial
Page 281
standard of comparison becomes almost an absolute necessity for the time
being.
Changes of form very generally, but not always, are accompanied with a
change in regularity: thus a flower habitually bi-lateral may assume the
characters of radiating symmetry and vice versâ. Increase or decrease of
size very frequently also are co-existent with an alteration in the usual form.
In the case of the arrangement of organs it is often difficult or impossible, in
the present state of our knowledge, to determine whether a given
arrangement is congenital or acquired subsequently to the first
development, whether for instance an isolation of parts be due to primordial
separation or to a subsequent disunion of originally combined organs, see p.
58. With reference to the changes in the form of organs, however, it is in
general more easy to ascertain the proximate cause of the appearance, and
thus teratological changes of form may be grouped according as they are
due to, 1, arrest of development; 2, undue or excessive development; 3,
perverted development; and 4, irregular development; hence the use of the
following terms—Stasimorphy, Pleiomorphy, Metamorphy, and
Heteromorphy—to include teratological changes really or apparently due to
one or other of the causes above mentioned. The classification here adopted
is of course to a considerable extent an arbitrary one and subject to
correction or modification, as the knowledge of the development of the
flowers in the various genera of plants advances.
FOOTNOTES:
[217] The word symmetry has been used in very different senses by different botanists,
sometimes as synonymous with "regularity," at other times to express the assumed typical
form of a flower. Payer understands it to be that arrangement of parts which permits of the
whole flower being divided vertically into two symmetrical halves (bi-lateral symmetry).
Others, again, have applied the term symmetry to the number of the parts of the flower,
reserving the terms "regularity" or "irregularity" for the form. It is here used in a general sense
being.
Changes of form very generally, but not always, are accompanied with a
change in regularity: thus a flower habitually bi-lateral may assume the
characters of radiating symmetry and vice versâ. Increase or decrease of
size very frequently also are co-existent with an alteration in the usual form.
In the case of the arrangement of organs it is often difficult or impossible, in
the present state of our knowledge, to determine whether a given
arrangement is congenital or acquired subsequently to the first
development, whether for instance an isolation of parts be due to primordial
separation or to a subsequent disunion of originally combined organs, see p.
58. With reference to the changes in the form of organs, however, it is in
general more easy to ascertain the proximate cause of the appearance, and
thus teratological changes of form may be grouped according as they are
due to, 1, arrest of development; 2, undue or excessive development; 3,
perverted development; and 4, irregular development; hence the use of the
following terms—Stasimorphy, Pleiomorphy, Metamorphy, and
Heteromorphy—to include teratological changes really or apparently due to
one or other of the causes above mentioned. The classification here adopted
is of course to a considerable extent an arbitrary one and subject to
correction or modification, as the knowledge of the development of the
flowers in the various genera of plants advances.
FOOTNOTES:
[217] The word symmetry has been used in very different senses by different botanists,
sometimes as synonymous with "regularity," at other times to express the assumed typical
form of a flower. Payer understands it to be that arrangement of parts which permits of the
whole flower being divided vertically into two symmetrical halves (bi-lateral symmetry).
Others, again, have applied the term symmetry to the number of the parts of the flower,
reserving the terms "regularity" or "irregularity" for the form. It is here used in a general sense
Page 282
to express the plan of the flower, and thus includes the arrangement, form, and number of its
component elements.
[218] See Baillon, 'Adansonia,' v, 176.
PART I.
STASIMORPHY.[219]
Deviations from the ordinary form of organs arising from stasis or arrest of
development are included under this heading.
There are many cases in which the forms proper to a juvenile condition of
the plant are retained for a much longer period than ordinary, or even
throughout the life of the individual growth goes on, but "development" is
checked. Such conditions may even be propagated by seed or bud. It is a
very general thing for botanists to consider these cases as reversions to a
simpler, primitive type, and this may be so; but on the other hand, they may
be degenerations from a complex type, or they may have no direct relation
to any antecedent condition. Stasimorphic changes affecting principally the
relative size of organs—such, for instance, as the non-development of
internodes, or the atrophy or suppression of parts will be found mentioned
in the sections relating to those subjects. In the present part those alterations
which affect the form of organs principally are treated of.
FOOTNOTES:
[219] Στασις-μορφωσις.
component elements.
[218] See Baillon, 'Adansonia,' v, 176.
PART I.
STASIMORPHY.[219]
Deviations from the ordinary form of organs arising from stasis or arrest of
development are included under this heading.
There are many cases in which the forms proper to a juvenile condition of
the plant are retained for a much longer period than ordinary, or even
throughout the life of the individual growth goes on, but "development" is
checked. Such conditions may even be propagated by seed or bud. It is a
very general thing for botanists to consider these cases as reversions to a
simpler, primitive type, and this may be so; but on the other hand, they may
be degenerations from a complex type, or they may have no direct relation
to any antecedent condition. Stasimorphic changes affecting principally the
relative size of organs—such, for instance, as the non-development of
internodes, or the atrophy or suppression of parts will be found mentioned
in the sections relating to those subjects. In the present part those alterations
which affect the form of organs principally are treated of.
FOOTNOTES:
[219] Στασις-μορφωσις.
Page 283
CHAPTER I.
PERSISTENCE OF JUVENILE FORMS.
The retention in adult life of a form characteristic of an early stage of
development, and therefore usually transient, may be manifested in any of
the organs of the plant. As these cases are for the most part treated under
separate headings, it is here only necessary to allude to a few, which it is
difficult to allocate satisfactorily, while the reader may be referred for other
instances of like nature to the sections on Peloria, Atrophy, Suppression,
Dimorphy, Substitutions, &c.
Fig. 115.—Juniperus sinensis. Two forms of leaves on branches of the same shrub.
Stasimorphy in the leaves of conifers.—In many conifers the leaves
produced in the young state of the plant are different, both in arrangement
and form, from those subsequently developed (see pp. 89, 90). But it
occasionally happens that the plant continues to form throughout its
existence leaves such as are usually produced only in a young state; thus M.
Gubler ('Bull. Soc. Bot., Fr.,' vol. viii, 1861, p. 527) describes a plant of
Pinus pinea in which the primordial, usually transitory, foliage was
PERSISTENCE OF JUVENILE FORMS.
The retention in adult life of a form characteristic of an early stage of
development, and therefore usually transient, may be manifested in any of
the organs of the plant. As these cases are for the most part treated under
separate headings, it is here only necessary to allude to a few, which it is
difficult to allocate satisfactorily, while the reader may be referred for other
instances of like nature to the sections on Peloria, Atrophy, Suppression,
Dimorphy, Substitutions, &c.
Fig. 115.—Juniperus sinensis. Two forms of leaves on branches of the same shrub.
Stasimorphy in the leaves of conifers.—In many conifers the leaves
produced in the young state of the plant are different, both in arrangement
and form, from those subsequently developed (see pp. 89, 90). But it
occasionally happens that the plant continues to form throughout its
existence leaves such as are usually produced only in a young state; thus M.
Gubler ('Bull. Soc. Bot., Fr.,' vol. viii, 1861, p. 527) describes a plant of
Pinus pinea in which the primordial, usually transitory, foliage was
Page 284
permanent, leaves of the ordinary shape not being developed at all. It more
often happens that some only of the leaves retain their young form while
others assume other shapes, see fig. 115. This happens frequently in the
larch and constantly in the Chinese juniper when it has arrived at a
considerable age. In Cupressus funebris two forms of leaves may often be
found on the same plant, the one representing the juvenile state, the other
the more developed condition. What is very singular, is that a cutting taken
from the branch with leaves of the young form grows up into a shrub
bearing leaves of no other shape, so that an ordinary observer unacquainted
with the history of the plant would imagine that he had to deal with two
distinct species. This fact is the more interesting when compared with the
alternation of generations which takes place among the lower animals.
The regular development of all the parts of the flower in a plant habitually
producing irregular flowers is referred to under the head of Peloria, but it
still remains to consider those examples in which some only of the parts of
the flower are affected in this manner.[220] Most of these cases are
elsewhere referred to in this volume under the particular form of
malformation assumed; but the following case may here be noticed as not
coming under any of the previous heads. It is an instance recorded by
Professor Babington ('Phytologist,' August, 1853), and in which the pod of
Medicago maculata, which is usually rolled up like a snail shell and
provided with spines, was sickle-shaped and unarmed.
FOOTNOTES:
[220] See a paper of Professor C. Morren's on "Floral Stesomy" in 'Bull. Acad. Belg.,' t. xix,
part ii, p. 519.
often happens that some only of the leaves retain their young form while
others assume other shapes, see fig. 115. This happens frequently in the
larch and constantly in the Chinese juniper when it has arrived at a
considerable age. In Cupressus funebris two forms of leaves may often be
found on the same plant, the one representing the juvenile state, the other
the more developed condition. What is very singular, is that a cutting taken
from the branch with leaves of the young form grows up into a shrub
bearing leaves of no other shape, so that an ordinary observer unacquainted
with the history of the plant would imagine that he had to deal with two
distinct species. This fact is the more interesting when compared with the
alternation of generations which takes place among the lower animals.
The regular development of all the parts of the flower in a plant habitually
producing irregular flowers is referred to under the head of Peloria, but it
still remains to consider those examples in which some only of the parts of
the flower are affected in this manner.[220] Most of these cases are
elsewhere referred to in this volume under the particular form of
malformation assumed; but the following case may here be noticed as not
coming under any of the previous heads. It is an instance recorded by
Professor Babington ('Phytologist,' August, 1853), and in which the pod of
Medicago maculata, which is usually rolled up like a snail shell and
provided with spines, was sickle-shaped and unarmed.
FOOTNOTES:
[220] See a paper of Professor C. Morren's on "Floral Stesomy" in 'Bull. Acad. Belg.,' t. xix,
part ii, p. 519.
Page 285
CHAPTER II.
REGULAR PELORIA.
Fig. 116.—Regular Peloria, Delphinium.
REGULAR PELORIA.
Fig. 116.—Regular Peloria, Delphinium.
Page 286
Fig. 117.—Sepal, petal, &c., of regular-flowered Delphinium.
Page 287
Fig. 118—Regular peloria, Viola.
Fig. 119—Double Violet, flower regular, petals multiplied, stamens and pistils petaloid.
When an habitually irregular flower becomes regular, it does so in one of
two ways; either by the non-development of the irregular portions, or by the
formation of irregular parts in increased number, so that the symmetry of
the flower is rendered perfect, as in the original peloria of Linnæus, and
which may be called irregular peloria, while the former case may be called
regular peloria. This latter appearance is therefore congenital, and due to an
arrest of development.[221] As the true nature of these cases has not been in
all cases recognised (even Moquin places them under the head of
deformities—they being less entitled to rank in that class than are the usual
flowers), it may be well to cite a few instances taken from various families.
In Delphinium peregrinum I have met with perfectly regular flowers having
five sepals and five oblong stalked petals, and a similar occurrence has been
noted in other species of this genus. Baillon,[222] in referring to these
Fig. 119—Double Violet, flower regular, petals multiplied, stamens and pistils petaloid.
When an habitually irregular flower becomes regular, it does so in one of
two ways; either by the non-development of the irregular portions, or by the
formation of irregular parts in increased number, so that the symmetry of
the flower is rendered perfect, as in the original peloria of Linnæus, and
which may be called irregular peloria, while the former case may be called
regular peloria. This latter appearance is therefore congenital, and due to an
arrest of development.[221] As the true nature of these cases has not been in
all cases recognised (even Moquin places them under the head of
deformities—they being less entitled to rank in that class than are the usual
flowers), it may be well to cite a few instances taken from various families.
In Delphinium peregrinum I have met with perfectly regular flowers having
five sepals and five oblong stalked petals, and a similar occurrence has been
noted in other species of this genus. Baillon,[222] in referring to these
Page 288
flowers, points out the resemblance that they bear to the double varieties of
Nigella. In the stellate columbines (Aquilegia) of gardens the tubular petals
are replaced by flat ones often in increased numbers. In violets both forms
of peloria occur, that in which there is an unusual number of spurs, and that
in which there are no spurs (var. anectaria). In the more perfect forms of
regular peloria occurring in the last-named genus the following changes
may be noticed: 1, an alteration in the direction of the flower so that it
remains in an erect position, and is not bent downwards as usual; 2, equality
of proportion in the sepals and petals; 3, absence of spurs, as also of hairs
on the lateral petals; 4, equal stamens whose anthers are sometimes entirely
destitute of the prolonged crest which forms so prominent a feature under
ordinary circumstances; 5, erect, not curved styles, and the stigmas not
prolonged into a beak, but having a more or less capitate form; ovary with
three or five cells, ovules normal.
These are cases where the change in question is most strongly marked, the
bi-lateral is completely replaced by the radiating symmetry. The absence of
the usual nectary, and of hairs on the side petals, the alterations in the form
of the style, etc., all show how much the process of fertilisation must be
altered from that which occurs under ordinary circumstances. In some of
the double violets now cultivated in gardens, a similar regularity of
proportion in the parts of the flower may be seen combined with the
substitution of petals for stamens and pistils, and with the development of
an increased number of petal-like organs.[223] Between these cases and the
ordinary spurred forms as well as those with an increased number of spurs,
many intermediate forms may be met with. That such regularity should
occur in this family is not to be wondered at seeing that there is a whole
subdivision of the order (Alsodeiæ) in which regular flowers are the rule.
In cultivated Pelargoniums the central flower of the umbel or "truss"
frequently retains its regularity of proportion, so as closely to approximate
Nigella. In the stellate columbines (Aquilegia) of gardens the tubular petals
are replaced by flat ones often in increased numbers. In violets both forms
of peloria occur, that in which there is an unusual number of spurs, and that
in which there are no spurs (var. anectaria). In the more perfect forms of
regular peloria occurring in the last-named genus the following changes
may be noticed: 1, an alteration in the direction of the flower so that it
remains in an erect position, and is not bent downwards as usual; 2, equality
of proportion in the sepals and petals; 3, absence of spurs, as also of hairs
on the lateral petals; 4, equal stamens whose anthers are sometimes entirely
destitute of the prolonged crest which forms so prominent a feature under
ordinary circumstances; 5, erect, not curved styles, and the stigmas not
prolonged into a beak, but having a more or less capitate form; ovary with
three or five cells, ovules normal.
These are cases where the change in question is most strongly marked, the
bi-lateral is completely replaced by the radiating symmetry. The absence of
the usual nectary, and of hairs on the side petals, the alterations in the form
of the style, etc., all show how much the process of fertilisation must be
altered from that which occurs under ordinary circumstances. In some of
the double violets now cultivated in gardens, a similar regularity of
proportion in the parts of the flower may be seen combined with the
substitution of petals for stamens and pistils, and with the development of
an increased number of petal-like organs.[223] Between these cases and the
ordinary spurred forms as well as those with an increased number of spurs,
many intermediate forms may be met with. That such regularity should
occur in this family is not to be wondered at seeing that there is a whole
subdivision of the order (Alsodeiæ) in which regular flowers are the rule.
In cultivated Pelargoniums the central flower of the umbel or "truss"
frequently retains its regularity of proportion, so as closely to approximate
Page 289
to the normal condition in the allied genus Geranium; this resemblance is
rendered greater by the fact that, under such circumstances, the patches of
darker colour characteristic of the ordinary flower are completely wanting;
the flower is as uniform in colour as in shape. Even the nectary which is
adherent to the upper surface of the pedicel in the normal flower disappears
—sometimes completely, at other tunes partially. The direction of the
stamens and style, and even that of the whole flower, becomes altered from
the inclined to the vertical position. In addition to these changes, which are
those most commonly met with, the number of the parts of the flower is
sometimes augmented, and a tendency to pass from the verticillate to the
spiral arrangement manifested. Schlechtendal mentions some flowers of
Tropæolum majus in which the flowers were perfectly regular and devoid of
spurs[224], while in the double varieties, now commonly grown in
greenhouses, the condition of parts is precisely the same as in the double
violet before alluded to. Among the Papilionaceæ the Laburnum and others
have been noticed to produce occasionally a perfectly regular flower in the
centre, or at the extremity of the inflorescence, though the peloria in this
flower is usually irregular. In the Gentianaceous genus Halenia, H.
heterantha is remarkable for the absence of spurs. Amongst Gesneraceæ,
Bignoniaceæ, Scrophulariaceæ, and other families of like structure, regular
peloria is not uncommon. Fig. 120 represents a case of this kind in
Eccremocarpus scaber, conjoined, as is frequently the case, with dialysis or
separation of the petals.[225] Many of the cultivated Gloxinias also show
erect, regular, five stamened flowers, but these are probably cases of
irregular peloria.
rendered greater by the fact that, under such circumstances, the patches of
darker colour characteristic of the ordinary flower are completely wanting;
the flower is as uniform in colour as in shape. Even the nectary which is
adherent to the upper surface of the pedicel in the normal flower disappears
—sometimes completely, at other tunes partially. The direction of the
stamens and style, and even that of the whole flower, becomes altered from
the inclined to the vertical position. In addition to these changes, which are
those most commonly met with, the number of the parts of the flower is
sometimes augmented, and a tendency to pass from the verticillate to the
spiral arrangement manifested. Schlechtendal mentions some flowers of
Tropæolum majus in which the flowers were perfectly regular and devoid of
spurs[224], while in the double varieties, now commonly grown in
greenhouses, the condition of parts is precisely the same as in the double
violet before alluded to. Among the Papilionaceæ the Laburnum and others
have been noticed to produce occasionally a perfectly regular flower in the
centre, or at the extremity of the inflorescence, though the peloria in this
flower is usually irregular. In the Gentianaceous genus Halenia, H.
heterantha is remarkable for the absence of spurs. Amongst Gesneraceæ,
Bignoniaceæ, Scrophulariaceæ, and other families of like structure, regular
peloria is not uncommon. Fig. 120 represents a case of this kind in
Eccremocarpus scaber, conjoined, as is frequently the case, with dialysis or
separation of the petals.[225] Many of the cultivated Gloxinias also show
erect, regular, five stamened flowers, but these are probably cases of
irregular peloria.
Page 290
Fig. 120.—Regular peloria, Eccremocarpus scaber.
A solitary flower of Pedicularis sylvatica was found by the Marquis of
Stafford near Dunrobin Castle in Sutherlandshire, in which the usual ringent
form of the corolla was replaced by the form called salver-shaped. There
were six stamens, four long and two short. Sir W. Hooker and Mr. Borrer
are stated to have found a similar flower in the same locality in 1809.[226]
The passage of ligulate to tubular corollas among Compositæ is not of such
common occurrence as is the converse change. I owe to Mr. Berkeley the
communication of a capitulum of a species of Bidens, in which there was a
transition from the form of ligulate corollas to those that were deeply
divided into three, four, or five oblong lobes. These then were instances of
regular peloria.
A solitary flower of Pedicularis sylvatica was found by the Marquis of
Stafford near Dunrobin Castle in Sutherlandshire, in which the usual ringent
form of the corolla was replaced by the form called salver-shaped. There
were six stamens, four long and two short. Sir W. Hooker and Mr. Borrer
are stated to have found a similar flower in the same locality in 1809.[226]
The passage of ligulate to tubular corollas among Compositæ is not of such
common occurrence as is the converse change. I owe to Mr. Berkeley the
communication of a capitulum of a species of Bidens, in which there was a
transition from the form of ligulate corollas to those that were deeply
divided into three, four, or five oblong lobes. These then were instances of
regular peloria.
Page 291
Fig. 121.—Flower of Cattleya marginata. Lip replaced by a flat petal.
In Orchidaceæ a similar change is not by any means infrequent; in a few,
indeed, a regular flower is the normal character, as in Dendrobium normale,
Oncidium heteranthum, Thelymitra, etc. Fig. 121, reduced from a cut in the
'Gardeners' Chronicle,' 1854, p. 804, represents an instance of this kind in
Cattleya marginata.
From the same journal the following account of a case of peloria in
Phalænopsis Schilleriana is also cited as a good illustration of this peculiar
change. The terminal flower differed entirely from all the others; instead of
the peculiar labellum there were three petals all exactly alike, and three
sepals also exactly alike; the petals resembled those of the other flowers of
the spike, and the upper sepal also; but the two lower sepals had no spots,
and were not reflexed as in the ordinary way: thus, these six parts of the
flower were all in one plane, and being close together at their edges, made
almost a full round flower; the column and pollen-glands were unaffected.
In Orchidaceæ a similar change is not by any means infrequent; in a few,
indeed, a regular flower is the normal character, as in Dendrobium normale,
Oncidium heteranthum, Thelymitra, etc. Fig. 121, reduced from a cut in the
'Gardeners' Chronicle,' 1854, p. 804, represents an instance of this kind in
Cattleya marginata.
From the same journal the following account of a case of peloria in
Phalænopsis Schilleriana is also cited as a good illustration of this peculiar
change. The terminal flower differed entirely from all the others; instead of
the peculiar labellum there were three petals all exactly alike, and three
sepals also exactly alike; the petals resembled those of the other flowers of
the spike, and the upper sepal also; but the two lower sepals had no spots,
and were not reflexed as in the ordinary way: thus, these six parts of the
flower were all in one plane, and being close together at their edges, made
almost a full round flower; the column and pollen-glands were unaffected.
Page 292
Professor Reichenbach also exhibited at the Amsterdam Botanical
Congress, of 1865, a flower of Selenipedium caudatum with a flat lip.
M. Gris[227] has placed on record some interesting cases of peloria of this
kind in Zingiber zerumbet; in the more complete forms the andrœcium or
staminal series was composed of six distinct pieces, the three inner of which
were fertile, while in the ordinary flower the andrœcium is composed of
two pieces, "a lip" and a fertile stamen. "Is it not a matter of regret," says
M. Gris, "to be obliged to call the latter the normal flower?"
Under this head may likewise be mentioned those cases in which the
normal, or at least the typical symmetry of the flower is restored by the
formation of parts usually suppressed; thus Moquin cites an abnormal
flower of Atriplex[228] hortensis described by M. Fenzl as having a true
calyx within the two bracts that usually alone encircle the stamens.
Adanson, also cited by Moquin, found a specimen of Bocconia with a
corolla. Arum maculatum has likewise been met with provided with a
genuine perianth as in Acorus and other Orontiads. The unusual
development of the sexual organs in diclinous flowers has been alluded to
under the head of heterogamy, and other cases where the symmetry of the
flower is rendered regular, by the development of parts ordinarily
suppressed, will be found in the chapters relating to deviations from the
usual number of organs.
This change, or rather this persistence of a form that is usually transient, is
generally accompanied by some other alterations. Change of direction, as
has been already mentioned, is one of the most common of these; separation
of the petals (Antirrhinum, Verbascum, &c.), and even their appearance in
leaf-like guise, are not infrequent (Delphinium, Antirrhinum, Verbascum,
&c.) At other times multiplication or increased number of the whorls of
petals takes place, often, but not always, at the expense of the sexual organs
Congress, of 1865, a flower of Selenipedium caudatum with a flat lip.
M. Gris[227] has placed on record some interesting cases of peloria of this
kind in Zingiber zerumbet; in the more complete forms the andrœcium or
staminal series was composed of six distinct pieces, the three inner of which
were fertile, while in the ordinary flower the andrœcium is composed of
two pieces, "a lip" and a fertile stamen. "Is it not a matter of regret," says
M. Gris, "to be obliged to call the latter the normal flower?"
Under this head may likewise be mentioned those cases in which the
normal, or at least the typical symmetry of the flower is restored by the
formation of parts usually suppressed; thus Moquin cites an abnormal
flower of Atriplex[228] hortensis described by M. Fenzl as having a true
calyx within the two bracts that usually alone encircle the stamens.
Adanson, also cited by Moquin, found a specimen of Bocconia with a
corolla. Arum maculatum has likewise been met with provided with a
genuine perianth as in Acorus and other Orontiads. The unusual
development of the sexual organs in diclinous flowers has been alluded to
under the head of heterogamy, and other cases where the symmetry of the
flower is rendered regular, by the development of parts ordinarily
suppressed, will be found in the chapters relating to deviations from the
usual number of organs.
This change, or rather this persistence of a form that is usually transient, is
generally accompanied by some other alterations. Change of direction, as
has been already mentioned, is one of the most common of these; separation
of the petals (Antirrhinum, Verbascum, &c.), and even their appearance in
leaf-like guise, are not infrequent (Delphinium, Antirrhinum, Verbascum,
&c.) At other times multiplication or increased number of the whorls of
petals takes place, often, but not always, at the expense of the sexual organs
Page 293
of the flower. Perhaps even more frequent is the increased number of parts
in the same whorl in cases of regular peloria; thus, in the Pelargoniums
before alluded to, the parts of the flower are frequently regulated by the
number six instead of five.
This form of peloria is most generally met with in flowers that are placed at
the end or in the centre of the inflorescence, or in such flowers as occur
singly at the end of the flower-stalk, as in Tropæolum, Viola, &c. It would
hence seem as if the freedom from pressure or restriction on one side
allowed the flower to develop equally in all directions, and thus to produce
regularity of form.
It is obvious, from what has been before said, that the process of
fertilisation is in many cases interfered with and altered by the change in the
conformation or the flower.
From overlooking the occasional existence of this form of peloria, new
genera have sometimes been formed on insufficient grounds. The genus
Aceranthus, for instance, consists of species of Epimedium in which the
customary spurs are not formed.[229]
The occurrence both of regular and irregular peloria on the same plant has
frequently been observed in Linaria. It has also been remarked that the
seedlings raised from these forms are not always constant; thus, the late Mr.
Crocker, formerly foreman in the Royal Gardens, Kew, informed me that he
fertilised some flowers of a drooping Gloxinia with their own pollen, and
that when the seedlings blossomed a large number of them produced the
erect regular flowers.
From what has been already said it will be seen that regular peloria is
closely allied to what Morren called epanody, or a return to the normal
condition. The reversion of a monstrous form to the normal one, as, for
in the same whorl in cases of regular peloria; thus, in the Pelargoniums
before alluded to, the parts of the flower are frequently regulated by the
number six instead of five.
This form of peloria is most generally met with in flowers that are placed at
the end or in the centre of the inflorescence, or in such flowers as occur
singly at the end of the flower-stalk, as in Tropæolum, Viola, &c. It would
hence seem as if the freedom from pressure or restriction on one side
allowed the flower to develop equally in all directions, and thus to produce
regularity of form.
It is obvious, from what has been before said, that the process of
fertilisation is in many cases interfered with and altered by the change in the
conformation or the flower.
From overlooking the occasional existence of this form of peloria, new
genera have sometimes been formed on insufficient grounds. The genus
Aceranthus, for instance, consists of species of Epimedium in which the
customary spurs are not formed.[229]
The occurrence both of regular and irregular peloria on the same plant has
frequently been observed in Linaria. It has also been remarked that the
seedlings raised from these forms are not always constant; thus, the late Mr.
Crocker, formerly foreman in the Royal Gardens, Kew, informed me that he
fertilised some flowers of a drooping Gloxinia with their own pollen, and
that when the seedlings blossomed a large number of them produced the
erect regular flowers.
From what has been already said it will be seen that regular peloria is
closely allied to what Morren called epanody, or a return to the normal
condition. The reversion of a monstrous form to the normal one, as, for
Page 294
instance, when the fern-leaved beech reverts to the normal type, was called
by the same author epistrophy.[230]
The following are the genera in which regular peloria has been most often
observed. It must, however, be remarked that in some of the flowers
recorded as peloric there is no indication as to which form of peloria the
case should be referred to. For other illustrations refer to chapters on
Heterogamy, Number, Irregular Peloria, &c.
*Delphinium peregrinum!
*Nigella damascena!
*Aquilegia vulgaris!
*Viola odorata!
hirta.
Epimedium, sp.
*Pelargonium zonale!
*inquinans!
Tropæolum majus!
*Wistaria sinensis.
Lupinus.
*Cytisus Laburnum!
Trifolium repens!
*Compositæ, gen. pl.!
Lonicera Periclymenum!
Streptocarpus Rexii.
*Digitalis purpurea.
*Scrophularia aquatica.
*Pentstemon.
*Linaria vulgaris!
*Antirrhinum majus!
Verbascum nigrum!
Columnea Schiedeana.
Halenia heterantha.
Galeobdolon luteum.
Prunella vulgaris!
Salvia, sp.!
Teucrium campanulatum.
by the same author epistrophy.[230]
The following are the genera in which regular peloria has been most often
observed. It must, however, be remarked that in some of the flowers
recorded as peloric there is no indication as to which form of peloria the
case should be referred to. For other illustrations refer to chapters on
Heterogamy, Number, Irregular Peloria, &c.
*Delphinium peregrinum!
*Nigella damascena!
*Aquilegia vulgaris!
*Viola odorata!
hirta.
Epimedium, sp.
*Pelargonium zonale!
*inquinans!
Tropæolum majus!
*Wistaria sinensis.
Lupinus.
*Cytisus Laburnum!
Trifolium repens!
*Compositæ, gen. pl.!
Lonicera Periclymenum!
Streptocarpus Rexii.
*Digitalis purpurea.
*Scrophularia aquatica.
*Pentstemon.
*Linaria vulgaris!
*Antirrhinum majus!
Verbascum nigrum!
Columnea Schiedeana.
Halenia heterantha.
Galeobdolon luteum.
Prunella vulgaris!
Salvia, sp.!
Teucrium campanulatum.
Page 295
Betonica alopecuros.
Eccremocarpus scaber.
Pedicularis sylvatica.
Zingiber Zerumbet.
Phalænopsis amabilis!
Phalænopsis Schilleriana.
Habenaria.
*Orchis morio.
mascula.
*Dendrobium, sp.
Atriplex, sp.
Cattleya Mossiæ!
marginata.
Calanthe vestita!
Oncidium, sp.!
Selenipedium caudatum.
Arum maculatum.
In addition to the references already given, further information on this
subject may be gained from consulting the following publications. See also
Irregular Peloria.
Giraud, 'Bot. Soc. Edinb.,' Dec. 12, 1839, Antirrhinum. Dareste, 'Ann. Sc. Nat.,' ser. 2,
1842, xviii, p. 220, Delphinium. C. Morren, 'Fuchsia,' p. 90, Calceolaria, 'Bull. Acad.
Belg.,' xx, part ii, p. 57; and E. Morren, 'Bull. Acad. Belg.,' 2nd ser., xix. p. 224, Gloxinia.
Richard, 'Mém. Soc. d'hist. nat.,' ii, p. 212, tab. 3. Lindley, 'Journ. Linn. Soc.,' iii, p. 9,
Dendrobium. Michalet, 'Bull. Soc, Bot. France,' vii, p. 625, Betonica. Gubler, 'Bull. Soc.
Bot. Fr.,' ix, 81, 'Des anomalies aberrantes et regularisantes.' Reichenbach fil. 'De pollinis
orchid. genesi ac structura,' 1852, Oncidium. Clos, 'Mém. Acad. Toulouse,' vi, 1862, Salvia.
Caspary, 'Verhandl. Phys. Œkon. Gesell. Königsberg,' 1860, i, 59, Columnea. Bureau,
'Bull. Soc. Bot. Fr.,' 1861, vol. viii, p. 710, Streptocarpus. Darwin, 'Variation of Animals
and Plants,' ii, pp. 59 and 396. Godron, 'Ex. Bull. Bot. Soc. Fr.,' xiv, p. 165, 'Rev. Bibl.,'
Wistaria. Marchand, 'Adansonia,' iv, p. 172, Lonicera. Baillon, 'Adansonia,' v, p. 177, 'Sur
la regularité transitoire de quelques fleurs irreg.,' shows that during the development of
some flowers which begin and end by being irregular, there is an intermediate state when
all the parts are regular. Helye, 'Revue Horticole,' Sept., 1868, p. 327. In this last paper,
published as this sheet is going through the press, the author states that he has raised from
seed three generations of plants of Antirrhinum with regular spur-less flowers. The original
Eccremocarpus scaber.
Pedicularis sylvatica.
Zingiber Zerumbet.
Phalænopsis amabilis!
Phalænopsis Schilleriana.
Habenaria.
*Orchis morio.
mascula.
*Dendrobium, sp.
Atriplex, sp.
Cattleya Mossiæ!
marginata.
Calanthe vestita!
Oncidium, sp.!
Selenipedium caudatum.
Arum maculatum.
In addition to the references already given, further information on this
subject may be gained from consulting the following publications. See also
Irregular Peloria.
Giraud, 'Bot. Soc. Edinb.,' Dec. 12, 1839, Antirrhinum. Dareste, 'Ann. Sc. Nat.,' ser. 2,
1842, xviii, p. 220, Delphinium. C. Morren, 'Fuchsia,' p. 90, Calceolaria, 'Bull. Acad.
Belg.,' xx, part ii, p. 57; and E. Morren, 'Bull. Acad. Belg.,' 2nd ser., xix. p. 224, Gloxinia.
Richard, 'Mém. Soc. d'hist. nat.,' ii, p. 212, tab. 3. Lindley, 'Journ. Linn. Soc.,' iii, p. 9,
Dendrobium. Michalet, 'Bull. Soc, Bot. France,' vii, p. 625, Betonica. Gubler, 'Bull. Soc.
Bot. Fr.,' ix, 81, 'Des anomalies aberrantes et regularisantes.' Reichenbach fil. 'De pollinis
orchid. genesi ac structura,' 1852, Oncidium. Clos, 'Mém. Acad. Toulouse,' vi, 1862, Salvia.
Caspary, 'Verhandl. Phys. Œkon. Gesell. Königsberg,' 1860, i, 59, Columnea. Bureau,
'Bull. Soc. Bot. Fr.,' 1861, vol. viii, p. 710, Streptocarpus. Darwin, 'Variation of Animals
and Plants,' ii, pp. 59 and 396. Godron, 'Ex. Bull. Bot. Soc. Fr.,' xiv, p. 165, 'Rev. Bibl.,'
Wistaria. Marchand, 'Adansonia,' iv, p. 172, Lonicera. Baillon, 'Adansonia,' v, p. 177, 'Sur
la regularité transitoire de quelques fleurs irreg.,' shows that during the development of
some flowers which begin and end by being irregular, there is an intermediate state when
all the parts are regular. Helye, 'Revue Horticole,' Sept., 1868, p. 327. In this last paper,
published as this sheet is going through the press, the author states that he has raised from
seed three generations of plants of Antirrhinum with regular spur-less flowers. The original
Page 296
wild plant was only partially peloric, but all the flowers produced on its descendants were
regular.
FOOTNOTES:
[221] "On the existence of two forms of Peloria," by M. T. Masters. 'Nat. Hist. Review,'
April, 1863.
[222] Baillon, 'Adansonia,' iv. p. 149.
[223] Similar cases are figured in 'Hort. Eystettens. Ic. Pl. Vern.' fol. 4, f. 1, 2. Viola martia
multiplici flore.
[224] 'Linnæa,' 1837, p. 128.
[225] M. Bureau, 'Bull. Soc. Bot. Fr.,' ix, p. 91, describes two genera of Bignoniaceæ in
which the flowers are normally regular and six parted.
[226] See 'Trans. Linn. Soc.,' vol. x. p. 227.
[227] 'Ann. Sc. Nat.,' ser. 4, 1859. tom. xi, p. 264, tab. 3.
[228] 'El. Ter. Veg.,' p. 342.
[229] Marchand, 'Adansonia,' vol. iv, p. 127.
[230] 'Bull. Acad. Belg.,' xvii. p. 17. "Fuchsia," p. 169.
PART II.
PLEIOMORPHY.[231]
Most irregular flowers owe their irregularity to an unequal development of
some of their organs as compared with that of others. When such flowers
become exceptionally regular they do so either because development does
not keep pace with growth, and a regular flower is thus the result of an
arrest of the former process (regular peloria), or because the comparatively
excessive development, which usually occurs in a few parts is, in
regular.
FOOTNOTES:
[221] "On the existence of two forms of Peloria," by M. T. Masters. 'Nat. Hist. Review,'
April, 1863.
[222] Baillon, 'Adansonia,' iv. p. 149.
[223] Similar cases are figured in 'Hort. Eystettens. Ic. Pl. Vern.' fol. 4, f. 1, 2. Viola martia
multiplici flore.
[224] 'Linnæa,' 1837, p. 128.
[225] M. Bureau, 'Bull. Soc. Bot. Fr.,' ix, p. 91, describes two genera of Bignoniaceæ in
which the flowers are normally regular and six parted.
[226] See 'Trans. Linn. Soc.,' vol. x. p. 227.
[227] 'Ann. Sc. Nat.,' ser. 4, 1859. tom. xi, p. 264, tab. 3.
[228] 'El. Ter. Veg.,' p. 342.
[229] Marchand, 'Adansonia,' vol. iv, p. 127.
[230] 'Bull. Acad. Belg.,' xvii. p. 17. "Fuchsia," p. 169.
PART II.
PLEIOMORPHY.[231]
Most irregular flowers owe their irregularity to an unequal development of
some of their organs as compared with that of others. When such flowers
become exceptionally regular they do so either because development does
not keep pace with growth, and a regular flower is thus the result of an
arrest of the former process (regular peloria), or because the comparatively
excessive development, which usually occurs in a few parts is, in
Page 297
exceptional cases manifested by all, hence the flower becomes regular from
the increase in number of its irregular elements. These latter cases, then, are
due to an excess of development, hence the application of the term
pleiomorphy. It must be understood that mere increase in the number of the
organs of a flower is not included under this head, but under that of
deviations from the ordinary number of parts.
FOOTNOTES:
[231] Ρλειος-μορφωσις.
CHAPTER I.
IRREGULAR PELORIA.
The term peloria was originally given by Linné to a malformation of
Linaria vulgaris, with five spurs and five stamens, which was first found in
1742 near Upsal. This was considered so marvellous a circumstance that the
term peloria, from the Greek πελωρ, a prodigy, was applied to it.[232] After a
time other irregular flowers were found in like condition, and so the term
peloria became applied to all cases wherein, on a plant habitually producing
irregular flowers, regular ones were formed. The fact that this regularity
might arise from two totally different causes was overlooked, or at least not
fully recognised, even by Moquin-Tandon himself. Where a flower retains
throughout life the same relative size in its parts that it had when those parts
first originated the result is, of course, a regular flower, as happens in
violets and other plants. This kind of peloria may for distinction sake be
called regular or congenital peloria (see chapter on that subject); but where
a flower becomes regular by the increase in number of its irregular portions,
as in the Linaria already alluded to, where not only one petal is spurred, but
the increase in number of its irregular elements. These latter cases, then, are
due to an excess of development, hence the application of the term
pleiomorphy. It must be understood that mere increase in the number of the
organs of a flower is not included under this head, but under that of
deviations from the ordinary number of parts.
FOOTNOTES:
[231] Ρλειος-μορφωσις.
CHAPTER I.
IRREGULAR PELORIA.
The term peloria was originally given by Linné to a malformation of
Linaria vulgaris, with five spurs and five stamens, which was first found in
1742 near Upsal. This was considered so marvellous a circumstance that the
term peloria, from the Greek πελωρ, a prodigy, was applied to it.[232] After a
time other irregular flowers were found in like condition, and so the term
peloria became applied to all cases wherein, on a plant habitually producing
irregular flowers, regular ones were formed. The fact that this regularity
might arise from two totally different causes was overlooked, or at least not
fully recognised, even by Moquin-Tandon himself. Where a flower retains
throughout life the same relative size in its parts that it had when those parts
first originated the result is, of course, a regular flower, as happens in
violets and other plants. This kind of peloria may for distinction sake be
called regular or congenital peloria (see chapter on that subject); but where
a flower becomes regular by the increase in number of its irregular portions,
as in the Linaria already alluded to, where not only one petal is spurred, but
Page 298
all five of them are furnished with such appendages, and which are the
result of an irregular development of those organs, the peloria is evidently
not congenital, but occurs at a more or less advanced stage of development.
To this latter form of peloria it is proposed to give the distinctive epithet of
irregular.
Peloria is either complete or incomplete; it is complete when the flower
appears perfectly symmetrical, it is incomplete when only a portion of the
flower is thus rendered regular. It is very common, for instance, to find
violets or Linarias with two or three spurs, and these intermediate stages are
very interesting, as they serve to show in what way the irregularity is
brought about. In Antirrhinum, Linaria, &c., intermediate forms show very
clearly that it is to the repetition of the form usually assumed by the petals
of the lower lip that the condition is due. This is also obvious in peloric
flowers of the Calceolaria. The perfect peloria of this flower is in general
erect, with five regular sepals, a regular corolla contracted at the base and at
the apex, but distended in the centre so as to resemble a lady's sleeve, tight
at the shoulder and wrist, and puffed in the centre!
result of an irregular development of those organs, the peloria is evidently
not congenital, but occurs at a more or less advanced stage of development.
To this latter form of peloria it is proposed to give the distinctive epithet of
irregular.
Peloria is either complete or incomplete; it is complete when the flower
appears perfectly symmetrical, it is incomplete when only a portion of the
flower is thus rendered regular. It is very common, for instance, to find
violets or Linarias with two or three spurs, and these intermediate stages are
very interesting, as they serve to show in what way the irregularity is
brought about. In Antirrhinum, Linaria, &c., intermediate forms show very
clearly that it is to the repetition of the form usually assumed by the petals
of the lower lip that the condition is due. This is also obvious in peloric
flowers of the Calceolaria. The perfect peloria of this flower is in general
erect, with five regular sepals, a regular corolla contracted at the base and at
the apex, but distended in the centre so as to resemble a lady's sleeve, tight
at the shoulder and wrist, and puffed in the centre!
Page 299
Fig. 122.—Peloric flower of Calceolaria.
Morren[233] describes a form intermediate between the ordinary slipper-
shaped corolla and the perfect peloria just described, and which he calls
sigmoid peloria. This flower is intermediate in direction between the erect
peloria and the ordinary reflected flower. The tube is curved like a swan's
neck and is dilated in front into two hollow bosses, such as we see in the
lower lip of an ordinary flower; beyond these it is contracted and is
prolonged into a slender beak terminating in two hollow teeth, between
which is the narrow orifice of the corolla. The colour at the base of the tube
inside is as in the perfect peloria; while round the summit of the tube, in
both cases, the intensity of colour is greatest on the outside. Now, in a
normal flower the deepest colour is within just opposite the orifice of the
corolla; this deep colour is also seen outside of the central and most
elevated portions of the lower lip. In the peloria the deep colour at the base
of the tube represents that which is near the orifice under ordinary
Morren[233] describes a form intermediate between the ordinary slipper-
shaped corolla and the perfect peloria just described, and which he calls
sigmoid peloria. This flower is intermediate in direction between the erect
peloria and the ordinary reflected flower. The tube is curved like a swan's
neck and is dilated in front into two hollow bosses, such as we see in the
lower lip of an ordinary flower; beyond these it is contracted and is
prolonged into a slender beak terminating in two hollow teeth, between
which is the narrow orifice of the corolla. The colour at the base of the tube
inside is as in the perfect peloria; while round the summit of the tube, in
both cases, the intensity of colour is greatest on the outside. Now, in a
normal flower the deepest colour is within just opposite the orifice of the
corolla; this deep colour is also seen outside of the central and most
elevated portions of the lower lip. In the peloria the deep colour at the base
of the tube represents that which is near the orifice under ordinary
Page 300
circumstances, while the outer patch of colour at the apex corresponds to
that formed on the upper surface of the lower lip. On the other hand, in
peloric flowers of Cytisus Laburnum, Clitoria Ternatea, Trifolium repens,
and other Papilionaceæ, it is the "standard," the form of which is repeated.
In the case of peloric aconites[234] the lateral and sometimes the inferior
coloured sepals assume the hooded form usually peculiar to the upper sepal
only, the number of the petals or nectaries being correspondingly increased.
Balsams become peloric by the augmentation in the number of spurs.[235]
So when orchids are affected with irregular peloria it is the form of the
labellum that is repeated, the accessory lips being sometimes the
representatives of stamens, which are usually suppressed in these flowers,
[236] but at other times the appearance is due simply to the fact that all three
petals assume the form usually confined to the lip, the staminal column
being unaffected, except that its direction and relative position with
reference to the other parts of the flower is different from ordinary. This
was the case in some flowers of Phalænopsis equestris sent to me by Mr.
Wentworth Buller. Fig. 123 represents a flower of Aristolochia caudata
with two lips, for which I am indebted to Mr. W. H. Baxter.
From these cases it is evident that the flowers in question become regular
by the repetition of the irregular parts.
that formed on the upper surface of the lower lip. On the other hand, in
peloric flowers of Cytisus Laburnum, Clitoria Ternatea, Trifolium repens,
and other Papilionaceæ, it is the "standard," the form of which is repeated.
In the case of peloric aconites[234] the lateral and sometimes the inferior
coloured sepals assume the hooded form usually peculiar to the upper sepal
only, the number of the petals or nectaries being correspondingly increased.
Balsams become peloric by the augmentation in the number of spurs.[235]
So when orchids are affected with irregular peloria it is the form of the
labellum that is repeated, the accessory lips being sometimes the
representatives of stamens, which are usually suppressed in these flowers,
[236] but at other times the appearance is due simply to the fact that all three
petals assume the form usually confined to the lip, the staminal column
being unaffected, except that its direction and relative position with
reference to the other parts of the flower is different from ordinary. This
was the case in some flowers of Phalænopsis equestris sent to me by Mr.
Wentworth Buller. Fig. 123 represents a flower of Aristolochia caudata
with two lips, for which I am indebted to Mr. W. H. Baxter.
From these cases it is evident that the flowers in question become regular
by the repetition of the irregular parts.
Page 301
Fig. 123.—Two-lipped flower of Aristolochia caudata.
It is probable that peloria may occur in any habitually irregular flower, and
that, if more attention were directed to the subject, illustrations might be
obtained from a larger number of natural families than can be done at
present. It is, however, necessary to exercise discrimination, and not to
attribute to peloria all the cases that at first sight appear to be so referable.
Thus, Professor Dickson exhibited at the Botanical Society of Edinburgh,
December 13th, 1860, four abnormal flowers of the common Indian cress
(Tropæolum majus), each presenting a supernumerary spur. On these he
remarked that "in Tropæolum the posterior part of the receptacle between
the insertion of the petals and that of the stamens is dilated so as to form the
spur which is so characteristic in the genus. The position of the spur in a
line with the posterior sepal has led many botanists to consider it as a
process of that sepal, but the fact of its being situated within the insertion of
the petals is conclusive as to its receptacular origin. In the flowers exhibited
It is probable that peloria may occur in any habitually irregular flower, and
that, if more attention were directed to the subject, illustrations might be
obtained from a larger number of natural families than can be done at
present. It is, however, necessary to exercise discrimination, and not to
attribute to peloria all the cases that at first sight appear to be so referable.
Thus, Professor Dickson exhibited at the Botanical Society of Edinburgh,
December 13th, 1860, four abnormal flowers of the common Indian cress
(Tropæolum majus), each presenting a supernumerary spur. On these he
remarked that "in Tropæolum the posterior part of the receptacle between
the insertion of the petals and that of the stamens is dilated so as to form the
spur which is so characteristic in the genus. The position of the spur in a
line with the posterior sepal has led many botanists to consider it as a
process of that sepal, but the fact of its being situated within the insertion of
the petals is conclusive as to its receptacular origin. In the flowers exhibited
Page 302
the supernumerary spur (as if to show its want of connection with any
sepal) was placed exactly between a lateral sepal and one of the anterior
sepals, sometimes on the one side of the flower and sometimes on the other.
These additional spurs were precisely similar to the normal ones, except
that they were a little shorter. This abnormality, although at first sight
seeming to indicate a pelorian tendency, is no approximation to regularity,
from the fact of the extra spur being differently placed, with regard to the
sepals, from the normal one."
Peloria of this kind, when perfect, is very often associated with other
alterations. Change of direction is one of the most common of these; the
usually drooping flower becomes erect, the stamens and style also are
changed in direction, while, not unfrequently, either the one or the other
(most often the stamens) are entirely suppressed. With this suppression an
increase in the size of the flower very generally coincides. The number of
parts is also frequently increased; thus, in Antirrhinum majus the corolla,
when subjected to peloria, is very generally six-parted, and has six stamens.
Fusion of one or more flowers is also a common accompaniment of peloria,
as in Digitalis purpurea, in which plant prolification often adds increased
complexity to the flower.
It has been stated by Moquin and others that the uppermost flower of an
inflorescence is the most subject to peloria; the uppermost flower of
Teucrium campanulatum, for instance, is very generally regular. In
Calceolaria it is the central terminal flower which is usually peloriated; on
the other hand, in Linaria and Antirrhinum the lower flowers, or those on
the secondary branches, are quite as often affected as the primary ones.
Cassini considered that the spur of Linaria was developed from the lower
petal rather than from the upper ones, because there is more room on the
side of the flower farthest from the stem than on the opposite side. With
reference to this point, M. Godron remarks that in habitually irregular
sepal) was placed exactly between a lateral sepal and one of the anterior
sepals, sometimes on the one side of the flower and sometimes on the other.
These additional spurs were precisely similar to the normal ones, except
that they were a little shorter. This abnormality, although at first sight
seeming to indicate a pelorian tendency, is no approximation to regularity,
from the fact of the extra spur being differently placed, with regard to the
sepals, from the normal one."
Peloria of this kind, when perfect, is very often associated with other
alterations. Change of direction is one of the most common of these; the
usually drooping flower becomes erect, the stamens and style also are
changed in direction, while, not unfrequently, either the one or the other
(most often the stamens) are entirely suppressed. With this suppression an
increase in the size of the flower very generally coincides. The number of
parts is also frequently increased; thus, in Antirrhinum majus the corolla,
when subjected to peloria, is very generally six-parted, and has six stamens.
Fusion of one or more flowers is also a common accompaniment of peloria,
as in Digitalis purpurea, in which plant prolification often adds increased
complexity to the flower.
It has been stated by Moquin and others that the uppermost flower of an
inflorescence is the most subject to peloria; the uppermost flower of
Teucrium campanulatum, for instance, is very generally regular. In
Calceolaria it is the central terminal flower which is usually peloriated; on
the other hand, in Linaria and Antirrhinum the lower flowers, or those on
the secondary branches, are quite as often affected as the primary ones.
Cassini considered that the spur of Linaria was developed from the lower
petal rather than from the upper ones, because there is more room on the
side of the flower farthest from the stem than on the opposite side. With
reference to this point, M. Godron remarks that in habitually irregular
Page 303
flowers the apex of the peduncle is oblique, and hence the flowers are bent
downwards or spread horizontally, but if the receptacle be quite flat and
level then the flower is regular. The oblique position causes some of the
organs to press on others, and hence induces abortion and suppression of
some parts and increased growth in others that are not subjected to pressure.
In a terminal peloriated flower of aconite, described by this naturalist, the
flower was removed so far from the nearest bracts that all its parts had the
chance of growing regularly. In ordinary cases M. Godron considers that the
compression of the lateral bracts is the cause of the irregularity of the
andrœcium and of the receptacle.[237]
It has also been somewhat too generally stated that peloria occurs
principally on luxuriant vigorous plants. It seems quite as often to happen in
plants characterised by their deficiencies in this respect. On this point M. de
Melicoq[238] says, referring to Linaria vulgaris affected with peloria, that on
the weakest plants the peloriated flower was at the top of the stem; while in
stronger plants, with more numerous flowers and larger foliage, the
peloriated flowers were principally to be found in the centre and at the base
of the inflorescence, and their pedicels were much longer than usual.
Linné, as has been already stated, considered these flowers to be sterile, and
only capable of multiplication by division of the root, but Willdenow
obtained seeds from the Linaria which reproduced the anomaly when sown
in rich soil. Baron Melicoq obtained similar results.[239] Mr. Darwin[240]
raised sixteen seedling plants of a peloric Antirrhinum, artificially fertilised
by its own pollen, all of which were as perfectly peloric as the parent plant.
On the other hand, the same observer alludes to the tendency that these
peloric plants have to revert to the usual form, as shown by the fact that
when the peloric flowers were crossed with pollen from flowers of the
ordinary shape, and vice versâ, not one of the seedlings, in either case, bore
peloric flowers. Hence, says Mr. Darwin, there is in these flowers "a strong
downwards or spread horizontally, but if the receptacle be quite flat and
level then the flower is regular. The oblique position causes some of the
organs to press on others, and hence induces abortion and suppression of
some parts and increased growth in others that are not subjected to pressure.
In a terminal peloriated flower of aconite, described by this naturalist, the
flower was removed so far from the nearest bracts that all its parts had the
chance of growing regularly. In ordinary cases M. Godron considers that the
compression of the lateral bracts is the cause of the irregularity of the
andrœcium and of the receptacle.[237]
It has also been somewhat too generally stated that peloria occurs
principally on luxuriant vigorous plants. It seems quite as often to happen in
plants characterised by their deficiencies in this respect. On this point M. de
Melicoq[238] says, referring to Linaria vulgaris affected with peloria, that on
the weakest plants the peloriated flower was at the top of the stem; while in
stronger plants, with more numerous flowers and larger foliage, the
peloriated flowers were principally to be found in the centre and at the base
of the inflorescence, and their pedicels were much longer than usual.
Linné, as has been already stated, considered these flowers to be sterile, and
only capable of multiplication by division of the root, but Willdenow
obtained seeds from the Linaria which reproduced the anomaly when sown
in rich soil. Baron Melicoq obtained similar results.[239] Mr. Darwin[240]
raised sixteen seedling plants of a peloric Antirrhinum, artificially fertilised
by its own pollen, all of which were as perfectly peloric as the parent plant.
On the other hand, the same observer alludes to the tendency that these
peloric plants have to revert to the usual form, as shown by the fact that
when the peloric flowers were crossed with pollen from flowers of the
ordinary shape, and vice versâ, not one of the seedlings, in either case, bore
peloric flowers. Hence, says Mr. Darwin, there is in these flowers "a strong
Page 304
latent tendency to become peloric, and there is also a still greater tendency
in all peloric plants to reacquire their normal irregular structure." So that
there are two opposed latent tendencies in the same plant. A similar remark
has been made with reference to malformations in general by other
observers.
It would be very interesting if some competent naturalist would collect
information as to whether any variations in degree of fertility exist in the
three forms of flowers in Linaria, viz. the ordinary one-spurred form, which
is intermediate between the spur-less and the five-spurred form. It must be
remembered, however, that in the latter cases the stamens are often
deficient. In the Compositæ, where there are regular flowers in the disc and
irregular ones in the ray, sexual differences, as is well known, accompany
the diversities in form.
To Mr. Darwin the author is indebted for the communication of some
flowers of Corydalis tuberosa (figs. 124, 125), provided with two spurs of
nearly equal size. To these flowers allusion is made in the work already
quoted[241] in the following terms:—"Corydalis tuberosa properly has one
of its two nectaries colourless, destitute of nectar, only half the size of the
other, and therefore to a certain extent in a rudimentary state; the pistil is
curved towards the perfect nectary, and the hood formed of the inner petals
slips off the pistil and stamens in one direction alone, so that when a bee
sucks the perfect nectary the stigma and stamens are exposed and rubbed
against the insect's body. In several closely allied genera, as in Dielytra,
there are two perfect nectaries; the pistil is straight, and the hood slips off
on either side, according as the bee sucks either nectary." In the flowers of
Corydalis, which were provided with two perfect nectaries containing
nectar, Mr. Darwin considers that there has been a redevelopment of a
partially aborted organ, accompanied by a change in the direction of the
pistil, which becomes straight, while the hood formed by the petals slips off
in all peloric plants to reacquire their normal irregular structure." So that
there are two opposed latent tendencies in the same plant. A similar remark
has been made with reference to malformations in general by other
observers.
It would be very interesting if some competent naturalist would collect
information as to whether any variations in degree of fertility exist in the
three forms of flowers in Linaria, viz. the ordinary one-spurred form, which
is intermediate between the spur-less and the five-spurred form. It must be
remembered, however, that in the latter cases the stamens are often
deficient. In the Compositæ, where there are regular flowers in the disc and
irregular ones in the ray, sexual differences, as is well known, accompany
the diversities in form.
To Mr. Darwin the author is indebted for the communication of some
flowers of Corydalis tuberosa (figs. 124, 125), provided with two spurs of
nearly equal size. To these flowers allusion is made in the work already
quoted[241] in the following terms:—"Corydalis tuberosa properly has one
of its two nectaries colourless, destitute of nectar, only half the size of the
other, and therefore to a certain extent in a rudimentary state; the pistil is
curved towards the perfect nectary, and the hood formed of the inner petals
slips off the pistil and stamens in one direction alone, so that when a bee
sucks the perfect nectary the stigma and stamens are exposed and rubbed
against the insect's body. In several closely allied genera, as in Dielytra,
there are two perfect nectaries; the pistil is straight, and the hood slips off
on either side, according as the bee sucks either nectary." In the flowers of
Corydalis, which were provided with two perfect nectaries containing
nectar, Mr. Darwin considers that there has been a redevelopment of a
partially aborted organ, accompanied by a change in the direction of the
pistil, which becomes straight, while the hood formed by the petals slips off
Page 305
in either direction, "so that these flowers have acquired the perfect structure,
so well adapted for insect agency, of Dielytra and its allies."
Fig. 124.—Two-spurred flowers of Corydalis.
Fig. 125.—Section through two-spurred flowers of Corydalis, Magnified.
Peloria, then, is especially interesting physiologically as well as
morphologically; it is also of value in a systematic point of view, as
showing how closely the deviations from the ordinary form of one plant
represent the ordinary condition of another; thus, the peloric Calceolarias
resemble the flowers of Fabiana, and De Candolle,[242] comparing the
peloric flowers of Scrophulariaceæ with those of Solanaceæ, concluded
that the former natural order was only an habitual alteration from the type
so well adapted for insect agency, of Dielytra and its allies."
Fig. 124.—Two-spurred flowers of Corydalis.
Fig. 125.—Section through two-spurred flowers of Corydalis, Magnified.
Peloria, then, is especially interesting physiologically as well as
morphologically; it is also of value in a systematic point of view, as
showing how closely the deviations from the ordinary form of one plant
represent the ordinary condition of another; thus, the peloric Calceolarias
resemble the flowers of Fabiana, and De Candolle,[242] comparing the
peloric flowers of Scrophulariaceæ with those of Solanaceæ, concluded
that the former natural order was only an habitual alteration from the type
Page 306
of the latter. Peloric flowers of Papilionaceæ in this way are
indistinguishable from those of Rosaceæ. In like manner we may trace an
analogy between the normal one-spurred Delphinium and the five-spurred
columbine (Aquilegia), an analogy strengthened by such a case as that of
the five-spurred flower of Delphinium elatum described by Godron.[243]
The Corydalis, before referred to, is another illustration of the same fact,
the structure being the same as in Dielytra, &c.
The ordinary irregular flowers may possibly be degenerated descendants of
a more completely organized ancestor, and some of the cases of peloria may
therefore be instances of reversion; some ancient Linaria may, perhaps,
have had all its petals spur-shaped, and the cases of irregular peloria now
found may be reversions to that original form. When both regular and
irregular forms of peloria occur on the same plant, as they frequently do in
Linaria, the one may be perhaps considered as a reversion to a very early
condition, the other to a later state, when all the petals were irregularly
formed. But before we can assert the truth of this surmise we must have
better evidence as to what the original condition really was than we have at
present.
The proximate cause of irregular peloria has been considered to be excess
of nourishment, but evidence as to this point is very conflicting. Willdenow
states that "radices peloriæ, solo sterili plantatæ, degenerant in Linariam,"
('Sp. Plant.,' iii, p. 254); but this opinion is counterbalanced by that of
others, while the frequent existence of both forms on the same plant, at the
same time, seems to negative the supposition of any direct effect from
external circumstances.
The following are the plants in which irregular peloria has been most often
observed:
Aconitum Napellus.
indistinguishable from those of Rosaceæ. In like manner we may trace an
analogy between the normal one-spurred Delphinium and the five-spurred
columbine (Aquilegia), an analogy strengthened by such a case as that of
the five-spurred flower of Delphinium elatum described by Godron.[243]
The Corydalis, before referred to, is another illustration of the same fact,
the structure being the same as in Dielytra, &c.
The ordinary irregular flowers may possibly be degenerated descendants of
a more completely organized ancestor, and some of the cases of peloria may
therefore be instances of reversion; some ancient Linaria may, perhaps,
have had all its petals spur-shaped, and the cases of irregular peloria now
found may be reversions to that original form. When both regular and
irregular forms of peloria occur on the same plant, as they frequently do in
Linaria, the one may be perhaps considered as a reversion to a very early
condition, the other to a later state, when all the petals were irregularly
formed. But before we can assert the truth of this surmise we must have
better evidence as to what the original condition really was than we have at
present.
The proximate cause of irregular peloria has been considered to be excess
of nourishment, but evidence as to this point is very conflicting. Willdenow
states that "radices peloriæ, solo sterili plantatæ, degenerant in Linariam,"
('Sp. Plant.,' iii, p. 254); but this opinion is counterbalanced by that of
others, while the frequent existence of both forms on the same plant, at the
same time, seems to negative the supposition of any direct effect from
external circumstances.
The following are the plants in which irregular peloria has been most often
observed:
Aconitum Napellus.
Page 307
Delphinium elatum!
Corydalis tuberosa.
*Viola odorata!
hirta.
Impatiens Balsamina.
Clitoria Ternatea.
Cytisus Laburnum!
Trifolium repens!
Lupinus polyphyllus!
*Gloxinia, var. cult.!
*Linaria vulgaris!
spuria.
Elatine.
triphylla.
æruginea.
triornithophora.
pilosa.
chalepensis.
cymbalaria!
purpurea!
decumbens.
Pelisseriana.
origanifolia.
Digitalis orientalis.
*purpurea!
Calceolaria crenatiflora.
rugosa.
*var. cult.!
Chelone barbata.
*Antirrhinum majus!
Rhinanthus crista galli.
Pedicularis sylvatica.
Pedicularis euphrasioides.
Scrophularia aquatica!
Sesamum indicum.
Lamium.
Mentha.
Sideritis.
Nepeta diffusa.
Corydalis tuberosa.
*Viola odorata!
hirta.
Impatiens Balsamina.
Clitoria Ternatea.
Cytisus Laburnum!
Trifolium repens!
Lupinus polyphyllus!
*Gloxinia, var. cult.!
*Linaria vulgaris!
spuria.
Elatine.
triphylla.
æruginea.
triornithophora.
pilosa.
chalepensis.
cymbalaria!
purpurea!
decumbens.
Pelisseriana.
origanifolia.
Digitalis orientalis.
*purpurea!
Calceolaria crenatiflora.
rugosa.
*var. cult.!
Chelone barbata.
*Antirrhinum majus!
Rhinanthus crista galli.
Pedicularis sylvatica.
Pedicularis euphrasioides.
Scrophularia aquatica!
Sesamum indicum.
Lamium.
Mentha.
Sideritis.
Nepeta diffusa.
Page 308
Galeopsis Ladanum.
Tetrahit.
Galeobdolon luteum.
Teucrium campanulatum!
Plectranthus fruticosus.
Cleonia lusitanica.
Dracocephalum austriacum.
Phlomis fruticosa!
Vitex incisa.
Aristolochia, sp.!
Ophrys aranifera!
Orchis simia.
pyramidalis!
latifolia!
morio!
papilionacea.
mascula.
latiflora.
conopsea.
Habenaria bifolia.
Corallorhiza innata.
Aceras anthropophora.
Cattleya Moasiæ!
Phalænopsis equestris!
Pogonia ophioglossoides!
The literature of peloria is very extensive. The following are the principal
papers, not already mentioned, which relate to the subject, arranged under
the genera, placing those first which are most subject to this anomaly (see
also Regular Peloria).
Linaria.—Adanson, 'Fam. Plant.,' t. i, p. 110. Jussien, 'Gen. Plant.,' p. 120. Poiret, 'Encycl.
Method, Suppl.,' t. iii, Jaeger, 'Missbilld. der Gewachs.,' pp. 94, 97, and 313. Cassini, 'Op.
Phytol.,' t. ii, p. 331. Ratzebourg, 'Animadv. ad pelor. spectand.,' 1825. Turpin. 'Ic. Veget.,'
tab. xx, f. 16. Curtis, 'Flor. Londin.,' i, 118. Hopkirk, 'Flora Anom.,' pl. vii, figs. 1, 2, 3.
Haller, 'Act. Helvet.,' 2, p. 25, t. iv. De Candolle, 'Flore Franc.,' t. iii, p. 583. Sowerby,
'Engl. Bot.,' iv, 260, ed. Syme, tab. 963. Chavannes, 'Mon. Antirrhin.' Delavaud, 'Bull. Soc.
Bot. France,' 1858, p. 689; id., 1860, p. 175. Heufler, 'Linnæa,' xvii, tab. ii. Weber,
'Verhandl. des Nat. Hist. Vereins. f. d. Rh. Preuss.,' 1850, tab. i, figs. 1–8. 'Verh. Nat. Hist.
Tetrahit.
Galeobdolon luteum.
Teucrium campanulatum!
Plectranthus fruticosus.
Cleonia lusitanica.
Dracocephalum austriacum.
Phlomis fruticosa!
Vitex incisa.
Aristolochia, sp.!
Ophrys aranifera!
Orchis simia.
pyramidalis!
latifolia!
morio!
papilionacea.
mascula.
latiflora.
conopsea.
Habenaria bifolia.
Corallorhiza innata.
Aceras anthropophora.
Cattleya Moasiæ!
Phalænopsis equestris!
Pogonia ophioglossoides!
The literature of peloria is very extensive. The following are the principal
papers, not already mentioned, which relate to the subject, arranged under
the genera, placing those first which are most subject to this anomaly (see
also Regular Peloria).
Linaria.—Adanson, 'Fam. Plant.,' t. i, p. 110. Jussien, 'Gen. Plant.,' p. 120. Poiret, 'Encycl.
Method, Suppl.,' t. iii, Jaeger, 'Missbilld. der Gewachs.,' pp. 94, 97, and 313. Cassini, 'Op.
Phytol.,' t. ii, p. 331. Ratzebourg, 'Animadv. ad pelor. spectand.,' 1825. Turpin. 'Ic. Veget.,'
tab. xx, f. 16. Curtis, 'Flor. Londin.,' i, 118. Hopkirk, 'Flora Anom.,' pl. vii, figs. 1, 2, 3.
Haller, 'Act. Helvet.,' 2, p. 25, t. iv. De Candolle, 'Flore Franc.,' t. iii, p. 583. Sowerby,
'Engl. Bot.,' iv, 260, ed. Syme, tab. 963. Chavannes, 'Mon. Antirrhin.' Delavaud, 'Bull. Soc.
Bot. France,' 1858, p. 689; id., 1860, p. 175. Heufler, 'Linnæa,' xvii, tab. ii. Weber,
'Verhandl. des Nat. Hist. Vereins. f. d. Rh. Preuss.,' 1850, tab. i, figs. 1–8. 'Verh. Nat. Hist.
Page 309
Ver. Rh. Preus.,' 1849, vol. vi, p. 290, tab. xiii.—Antirrhinum, Clos, 'Mém. Acad. Toulous.,'
vi, 1862. Chavannes, 'Mon. Antirrh.,' p. 62. Fresenius, 'Mus. Senkenb.,' ii, t. iv, fig. 10.
'Bot. Soc. Edinb.,' 1851, July 10.—Calceolaria, Chamisso, 'Linnæa,' t. vii, p. 206.
Guillemin, 'Archiv. Bot.,' t. ii, p. 1 et 136. Schlechtendal, 'Linnæa,' xii, p. 686. Ernst Meyer,
'Linnæa,' xvi, 26, tab. iii. Morren, 'Bull. Acad. Belg.,' t. xv, n. 7, et t. xviii, p. 583. 'Gard.
Chron.,' 1850, p. 389; ibid., 1866, p. 612.—Viola, Leers, 'Flor. Herborn.,' p. 145. De
Candolle, 'Organ. Veget.,' t. i, p. 519, pl. xlv. Forbes, 'Proc. Linn. Soc.,' June 6, 1848, p.
382. Hildebrand, 'Bot. Zeit.,' 1862, vol. xx, tab. viii.—Orchidaceæ, His, 'Jourl. Phys.,' 65,
p. 241. Wydler, 'Arch. Bot.,' t. ii, p. 310, tab. xvi. R. Brown, 'Obs. organ. Orchid.,' p. 698.
A. Richard, 'Mém. soc. d'hist. nat.,' t. i, p. 212. Greville, 'Flora Edinens.,' p. 87
(Corallorhiza). Curtis, 'Flora Londinensis,' t. lxxxii. Morren, C., 'Bull. Acad. Roy. Belg.,' t.
xix, part ii, p. 171. Clos, 'Mém. Acad. Sc. Toulous.,' 5 ser., vol. iii. Caspary, 'Schrift. K.
Gesellsch. Königsberg,' 1860, i, 59. Masters, 'Jourl. Linn. Soc.,' vol. viii, p. 208 (Ophrys,
Pogonia). Duchartre, 'Bull. Soc. Bot. Fr.,' vol. vii, 1860, p. 26, Cattleya. Cramer,
'Bildungsabweich.'—Limosella, Baillon, 'Adansonia,' i, p. 305. (Flower normally irregular,
becoming regular "à force d'irregularité.")—Chelone, Chamisso, 'Linnæa,' vii, p. 206,—
Clitoria, Bonavia, 'Gard. Chron.,' 1868, p. 1013. In this latter communication, published as
this sheet is passing through the press, the author gives an interesting account of the
transitional stages between the ordinary papilionaceous condition and the regular form
which is like that of a Rosaceous plant. The peloric form is stated to be transmitted by
seed.
For other references see Moq.-Tandon, 'El. Terat. Veget.,' p. 186. Hallier, 'Phytopathol.,' p.
151.
FOOTNOTES:
[232] 'Amœn. Acad.,' i, p. 55, t. iii (1744):—The following note refers to Linné's notion that
these forms were due to hybridization. It is extracted from Gmelin's edition of the 'Systema
Naturæ,' 1791, p. 931. "Linariæ proles hybrida, ejusdemque qualitatis et constans, radicibus
infinite sese multiplicans charactere fructificationis diversissima, corolla regulari, quinque-
corniculata, pentandra, ut genus proprium absolute constitueret et distinctissimum, nisi fructus
frequentissime abortiret. Naturæ prodigium. Ita quidem a Linné. Verisimilor autem videtur ea
opinio, quæ peloriam pro peculiari degeneratione monstrosa floris habet, in quam inclinare hoc
genus (Linaria) præ aliis, similis a forma deflexio in aliis speciebus, e.g. spurio Elatine,
cymbalaria, observata, ... Merk., 'Goett. gel. Anz.,' 1774, n. 121. Linck, 'Annal. Naturg.,' i, p.
32."
[233] 'Bull. Acad. Belg.,' xviii, part i, p. 591. Lobelia, p. 137.
[234] See also Seringe, 'Esquisse d'une Monogr. du genre Aconitum,' p. 124.
[235] Schlotterbec, 'Act. Helvet.,' t. ii, pl. i, Roeper. Balsam, p. 10, note.
vi, 1862. Chavannes, 'Mon. Antirrh.,' p. 62. Fresenius, 'Mus. Senkenb.,' ii, t. iv, fig. 10.
'Bot. Soc. Edinb.,' 1851, July 10.—Calceolaria, Chamisso, 'Linnæa,' t. vii, p. 206.
Guillemin, 'Archiv. Bot.,' t. ii, p. 1 et 136. Schlechtendal, 'Linnæa,' xii, p. 686. Ernst Meyer,
'Linnæa,' xvi, 26, tab. iii. Morren, 'Bull. Acad. Belg.,' t. xv, n. 7, et t. xviii, p. 583. 'Gard.
Chron.,' 1850, p. 389; ibid., 1866, p. 612.—Viola, Leers, 'Flor. Herborn.,' p. 145. De
Candolle, 'Organ. Veget.,' t. i, p. 519, pl. xlv. Forbes, 'Proc. Linn. Soc.,' June 6, 1848, p.
382. Hildebrand, 'Bot. Zeit.,' 1862, vol. xx, tab. viii.—Orchidaceæ, His, 'Jourl. Phys.,' 65,
p. 241. Wydler, 'Arch. Bot.,' t. ii, p. 310, tab. xvi. R. Brown, 'Obs. organ. Orchid.,' p. 698.
A. Richard, 'Mém. soc. d'hist. nat.,' t. i, p. 212. Greville, 'Flora Edinens.,' p. 87
(Corallorhiza). Curtis, 'Flora Londinensis,' t. lxxxii. Morren, C., 'Bull. Acad. Roy. Belg.,' t.
xix, part ii, p. 171. Clos, 'Mém. Acad. Sc. Toulous.,' 5 ser., vol. iii. Caspary, 'Schrift. K.
Gesellsch. Königsberg,' 1860, i, 59. Masters, 'Jourl. Linn. Soc.,' vol. viii, p. 208 (Ophrys,
Pogonia). Duchartre, 'Bull. Soc. Bot. Fr.,' vol. vii, 1860, p. 26, Cattleya. Cramer,
'Bildungsabweich.'—Limosella, Baillon, 'Adansonia,' i, p. 305. (Flower normally irregular,
becoming regular "à force d'irregularité.")—Chelone, Chamisso, 'Linnæa,' vii, p. 206,—
Clitoria, Bonavia, 'Gard. Chron.,' 1868, p. 1013. In this latter communication, published as
this sheet is passing through the press, the author gives an interesting account of the
transitional stages between the ordinary papilionaceous condition and the regular form
which is like that of a Rosaceous plant. The peloric form is stated to be transmitted by
seed.
For other references see Moq.-Tandon, 'El. Terat. Veget.,' p. 186. Hallier, 'Phytopathol.,' p.
151.
FOOTNOTES:
[232] 'Amœn. Acad.,' i, p. 55, t. iii (1744):—The following note refers to Linné's notion that
these forms were due to hybridization. It is extracted from Gmelin's edition of the 'Systema
Naturæ,' 1791, p. 931. "Linariæ proles hybrida, ejusdemque qualitatis et constans, radicibus
infinite sese multiplicans charactere fructificationis diversissima, corolla regulari, quinque-
corniculata, pentandra, ut genus proprium absolute constitueret et distinctissimum, nisi fructus
frequentissime abortiret. Naturæ prodigium. Ita quidem a Linné. Verisimilor autem videtur ea
opinio, quæ peloriam pro peculiari degeneratione monstrosa floris habet, in quam inclinare hoc
genus (Linaria) præ aliis, similis a forma deflexio in aliis speciebus, e.g. spurio Elatine,
cymbalaria, observata, ... Merk., 'Goett. gel. Anz.,' 1774, n. 121. Linck, 'Annal. Naturg.,' i, p.
32."
[233] 'Bull. Acad. Belg.,' xviii, part i, p. 591. Lobelia, p. 137.
[234] See also Seringe, 'Esquisse d'une Monogr. du genre Aconitum,' p. 124.
[235] Schlotterbec, 'Act. Helvet.,' t. ii, pl. i, Roeper. Balsam, p. 10, note.
Page 310
[236] Masters. "Peloria, &c., Ophrys aranifera," 'Journ. Linn. Soc.,' viii, p. 207.
[237] Godron, "Mém. sur les Fumarieès à fl. irreg.," 'Ann. Sc. Nat.,' sér. 5, vol. ii, tab. xvii, p.
280.
[238] 'Bull. Soc. Bot. France,' vol. v, 1858, p. 701.
[239] 'Bull. Soc. Bot. France,' vol. vi, 1859, p. 717.
[240] 'Variation of Anim. and Plants,' ii, p. 70.
[241] Loc. cit., p. 59.
[242] 'Théor. Elém.,' ed. 2, p. 266.
[243] Cited in 'Bull. Soc. Bot. France,' vol. xiii (Rev. Bibl.), p. 81.
PART III.
METAMORPHY.
Much of the objection with which Goethe's famous essay on the
'Metamorphosis of Plants' was met on its publication may be traced to a
misapprehension of the sense in which Goethe employed the word. As used
by him, it had nearly the same signification as now applied to the word
development by organogenists. It does not necessarily imply that there has
been a change in any particular organ, but rather that there has been, to
some extent, a change in the plan of construction, in accordance with which
a deviation from the customary form results. The particular organ was never
anything else than what it is; it has not been metamorphosed in the ordinary
sense of the word; for instance, in a double flower, where the stamens are,
as it is said, changed or metamorphosed into petals, no absolute change
really has taken place—the petal was never a stamen, although it occupies
the position of the latter, and may be considered a substitute for it.
[237] Godron, "Mém. sur les Fumarieès à fl. irreg.," 'Ann. Sc. Nat.,' sér. 5, vol. ii, tab. xvii, p.
280.
[238] 'Bull. Soc. Bot. France,' vol. v, 1858, p. 701.
[239] 'Bull. Soc. Bot. France,' vol. vi, 1859, p. 717.
[240] 'Variation of Anim. and Plants,' ii, p. 70.
[241] Loc. cit., p. 59.
[242] 'Théor. Elém.,' ed. 2, p. 266.
[243] Cited in 'Bull. Soc. Bot. France,' vol. xiii (Rev. Bibl.), p. 81.
PART III.
METAMORPHY.
Much of the objection with which Goethe's famous essay on the
'Metamorphosis of Plants' was met on its publication may be traced to a
misapprehension of the sense in which Goethe employed the word. As used
by him, it had nearly the same signification as now applied to the word
development by organogenists. It does not necessarily imply that there has
been a change in any particular organ, but rather that there has been, to
some extent, a change in the plan of construction, in accordance with which
a deviation from the customary form results. The particular organ was never
anything else than what it is; it has not been metamorphosed in the ordinary
sense of the word; for instance, in a double flower, where the stamens are,
as it is said, changed or metamorphosed into petals, no absolute change
really has taken place—the petal was never a stamen, although it occupies
the position of the latter, and may be considered a substitute for it.
Page 311
The term metamorphosis, then, really implies an alteration in the organizing
force, taking effect at a very early period of the life of the flower, at or
before the period when the primitive aggregation of cells, of which it is at
that time composed, becomes separated or "differentiated" into the several
parts of the flower. In other words, the "development" of the flower pursues
a different course from what is usual. In the preceding sections the effects
of arrest and of excess in this process have been partly treated of; other
deviations arising from similar causes will be mentioned elsewhere, but,
under the present heading, are specially included cases not of merely
diminished or increased, but of perverted development; the natural process
is here not necessarily checked or enhanced, but it is changed. Hence, in the
present work, the term metamorphy is employed to distinguish cases where
the ordinary course of development has been perverted or changed. As it is
applied solely for teratological purposes, the ordinary acceptation of the
term, as nearly synonymous with "development," is not interfered with.
In order to avoid other possible misapprehensions, the terms retrograde and
progressive metamorphosis employed by Goethe are not herein used, their
place being, to a great extent, supplied by the more intelligible expressions
arrest or excess of development.[244]
FOOTNOTES:
[244] See Goethe, 'Versuch. der Metam. der Pflanzen,' 1790. English translation by Emily M.
Cox, in Seemann's 'Journal of Botany,' vol. i, 1863, p. 327. For a brief sketch of the origin and
progress of the theory of vegetable morphology, prior to the publications of Wolff, Linné, and
Goethe, as well as for an attempt to show what share each of these authors had in the
establishment of the doctrine, the reader is referred to an article in the 'Brit. and For. Medico-
Chirurgical Review,' January, 1862, entitled "Vegetable Morphology: its History and Present
Condition," by Maxwell T. Masters.
force, taking effect at a very early period of the life of the flower, at or
before the period when the primitive aggregation of cells, of which it is at
that time composed, becomes separated or "differentiated" into the several
parts of the flower. In other words, the "development" of the flower pursues
a different course from what is usual. In the preceding sections the effects
of arrest and of excess in this process have been partly treated of; other
deviations arising from similar causes will be mentioned elsewhere, but,
under the present heading, are specially included cases not of merely
diminished or increased, but of perverted development; the natural process
is here not necessarily checked or enhanced, but it is changed. Hence, in the
present work, the term metamorphy is employed to distinguish cases where
the ordinary course of development has been perverted or changed. As it is
applied solely for teratological purposes, the ordinary acceptation of the
term, as nearly synonymous with "development," is not interfered with.
In order to avoid other possible misapprehensions, the terms retrograde and
progressive metamorphosis employed by Goethe are not herein used, their
place being, to a great extent, supplied by the more intelligible expressions
arrest or excess of development.[244]
FOOTNOTES:
[244] See Goethe, 'Versuch. der Metam. der Pflanzen,' 1790. English translation by Emily M.
Cox, in Seemann's 'Journal of Botany,' vol. i, 1863, p. 327. For a brief sketch of the origin and
progress of the theory of vegetable morphology, prior to the publications of Wolff, Linné, and
Goethe, as well as for an attempt to show what share each of these authors had in the
establishment of the doctrine, the reader is referred to an article in the 'Brit. and For. Medico-
Chirurgical Review,' January, 1862, entitled "Vegetable Morphology: its History and Present
Condition," by Maxwell T. Masters.
Page 312
CHAPTER I.
PHYLLODY.
This condition, wherein true leaves are substituted for some other organs,
[245] must be distinguished from Virescence, q. v., in which the parts
affected have simply the green colour of leaves, without their form or
structure. The appearance of perfect leaves, in place of other organs, is
frequently looked on as due to retrograde metamorphosis, or to an arrest of
development. But this is not strictly correct; for instance, suppose a petal,
which is very generally merely the sheath of a leaf, with the addition of
colouring matter, to be replaced by a perfect leaf, one in which all three
constituent parts, sheath, stalk, and blade, are present, it surely can hardly
be said that there has been any retrogression or arrest of development in the
formation of a complete in place of an incomplete organ. The term
retrograde here is used in a purely theoretical sense, and cannot be held to
imply any actual degradation. Morphologically, as has been stated, the case
is one of advance rather than the reverse, and hence the assignment of
instances of this nature to a perversion of development, rather than to a
diminution or to an exaltation of that process, seems most consistent with
truth. The affected organs have really undergone no actual change, simply
the direction of the organising force has been altered at a very early state, so
that the usual differentiation of parts has not taken place.
PHYLLODY.
This condition, wherein true leaves are substituted for some other organs,
[245] must be distinguished from Virescence, q. v., in which the parts
affected have simply the green colour of leaves, without their form or
structure. The appearance of perfect leaves, in place of other organs, is
frequently looked on as due to retrograde metamorphosis, or to an arrest of
development. But this is not strictly correct; for instance, suppose a petal,
which is very generally merely the sheath of a leaf, with the addition of
colouring matter, to be replaced by a perfect leaf, one in which all three
constituent parts, sheath, stalk, and blade, are present, it surely can hardly
be said that there has been any retrogression or arrest of development in the
formation of a complete in place of an incomplete organ. The term
retrograde here is used in a purely theoretical sense, and cannot be held to
imply any actual degradation. Morphologically, as has been stated, the case
is one of advance rather than the reverse, and hence the assignment of
instances of this nature to a perversion of development, rather than to a
diminution or to an exaltation of that process, seems most consistent with
truth. The affected organs have really undergone no actual change, simply
the direction of the organising force has been altered at a very early state, so
that the usual differentiation of parts has not taken place.
Page 313
Fig. 126.—'Rose plantain,' Plantago media var., spike contracted; bracts leafy.
Phyllody of the bracts.—As bracts are very generally imperfect organs, so
their replacement by perfect leaves is not attributable to arrest of
development or retrograde metamorphosis, but the reverse. The bracts of
some species of Plantago[246] are very subject to this change. Thus, in the
rose plantain of gardens, P. media (fig. 126), the bracts are leafy and the
axis depressed or not elongated, so that it is surmounted by a rosette of
small leafy organs. A similar condition of the bracts, unattended with arrest
of growth in the axis, is common in P. major (fig. 127) and in P. lanceolata
(see p. 108). It also occurs in the bracts of Corydalis solida, Amorpha
fruticosa, Ajuga reptans, Parthenium inodorum, Centaurea Jacea, in the
involucral bracts of the dandelion, the daisy, and many other composites. In
the 'Gardeners Chronicle,' 1852, p. 579, is figured a dahlia in which the
bracts of the involucre and the scales of the receptacle had all assumed the
form, texture, and venation of leaves.[247]
Phyllody of the bracts.—As bracts are very generally imperfect organs, so
their replacement by perfect leaves is not attributable to arrest of
development or retrograde metamorphosis, but the reverse. The bracts of
some species of Plantago[246] are very subject to this change. Thus, in the
rose plantain of gardens, P. media (fig. 126), the bracts are leafy and the
axis depressed or not elongated, so that it is surmounted by a rosette of
small leafy organs. A similar condition of the bracts, unattended with arrest
of growth in the axis, is common in P. major (fig. 127) and in P. lanceolata
(see p. 108). It also occurs in the bracts of Corydalis solida, Amorpha
fruticosa, Ajuga reptans, Parthenium inodorum, Centaurea Jacea, in the
involucral bracts of the dandelion, the daisy, and many other composites. In
the 'Gardeners Chronicle,' 1852, p. 579, is figured a dahlia in which the
bracts of the involucre and the scales of the receptacle had all assumed the
form, texture, and venation of leaves.[247]
Page 314
Fig. 127.—Leaf-like bracts in Plantago major.
Page 315
Fig. 128.—Dahlia. Scales of receptacle leafy.
In Umbelliferæ the substitution of leaves for involucral bracts is not
infrequent. It has been observed among other plants in Angelica Razoulzii,
Carum carui, Daucus Carota, &c. The scales of the hop (Humulus Lupulus)
not infrequently manifest this change, as do also the bracts of many
amentaceous plants, e.g. in the male catkins of the walnut, the female
catkins of the alder,[248] of some willows,[249] &c. The bracts of some
Euphorbiaceæ, as E. pusilla, E. Lathyris, E. Cyparissias, have been
observed to undergo a similar alteration.[250]
Amongst monocotyledons an analogous change occurs not unfrequently, as
in some commelynaceous plants, e.g. Tradescantia, in Musa, &c.
The spathe of Arum maculatum is sometimes represented by a stalked leaf
similar to that which occurs, under ordinary circumstances, in
Spathiphyllum, but in which genus the spadix is more or less adherent to the
leaf-like spathe.[251] In Schœnus cephalotes a similar exaggerated
development of the bracts is figured by Rottboell.[252]
Phyllody in inflorescence of Conifers.—This demands passing notice by
reason of the interest attaching to the morphological construction of these
plants. The elongation of the axis which occurs in the female cones has
been already alluded to under the head of prolification of the inflorescence.
This change is frequently associated with a more or less foliaceous
condition of the bracts, which, indeed, may be seen to be serially
continuous, both above and below, with the ordinary leaves. The scales, too,
become notched and bipartite, and show, between the lobes, the rudiment of
a bud, which in a further stage becomes developed into a shoot bearing
leaves. Such a change has been described by Parlatore in Abies Brunoniana,
and examples may frequently be met with in the larch (Larix europæa), and
specially in Cryptomeria japonica.[253] The scales of the male catkins of
In Umbelliferæ the substitution of leaves for involucral bracts is not
infrequent. It has been observed among other plants in Angelica Razoulzii,
Carum carui, Daucus Carota, &c. The scales of the hop (Humulus Lupulus)
not infrequently manifest this change, as do also the bracts of many
amentaceous plants, e.g. in the male catkins of the walnut, the female
catkins of the alder,[248] of some willows,[249] &c. The bracts of some
Euphorbiaceæ, as E. pusilla, E. Lathyris, E. Cyparissias, have been
observed to undergo a similar alteration.[250]
Amongst monocotyledons an analogous change occurs not unfrequently, as
in some commelynaceous plants, e.g. Tradescantia, in Musa, &c.
The spathe of Arum maculatum is sometimes represented by a stalked leaf
similar to that which occurs, under ordinary circumstances, in
Spathiphyllum, but in which genus the spadix is more or less adherent to the
leaf-like spathe.[251] In Schœnus cephalotes a similar exaggerated
development of the bracts is figured by Rottboell.[252]
Phyllody in inflorescence of Conifers.—This demands passing notice by
reason of the interest attaching to the morphological construction of these
plants. The elongation of the axis which occurs in the female cones has
been already alluded to under the head of prolification of the inflorescence.
This change is frequently associated with a more or less foliaceous
condition of the bracts, which, indeed, may be seen to be serially
continuous, both above and below, with the ordinary leaves. The scales, too,
become notched and bipartite, and show, between the lobes, the rudiment of
a bud, which in a further stage becomes developed into a shoot bearing
leaves. Such a change has been described by Parlatore in Abies Brunoniana,
and examples may frequently be met with in the larch (Larix europæa), and
specially in Cryptomeria japonica.[253] The scales of the male catkins of
Page 316
conifers likewise occasionally assume the appearance of leaves; this may be
seen in monstrous catkins of Araucaria, as also in Podocarpeæ and
Cupressineæ (Eichler).
Phyllody of the calyx.—Sepals under ordinary circumstances are so like
leaves, that it is not wonderful that they are often replaced by those organs.
[254] A singular instance of this has been mentioned as occurring in Cakile
maritima, wherein the sepals were found by M. Fournier to be pinnatifid
like the ordinary leaves of the plant.[255] The sepals of Ranunculaceæ and
Rosaceæ, for example, Rosa, Geum, are particularly liable to this change.
Fig. 129.—Flower of rose, sepals replaced by five perfect leaves; axis prolonged through the flower
in the form of a leafy branch.
In a species of Geranium recently examined the sepals presented
themselves in the form of three-lobed leaflets; so in fuchsias and in
Epilobium hirsutum the sepals occasionally are not distinguishable from
seen in monstrous catkins of Araucaria, as also in Podocarpeæ and
Cupressineæ (Eichler).
Phyllody of the calyx.—Sepals under ordinary circumstances are so like
leaves, that it is not wonderful that they are often replaced by those organs.
[254] A singular instance of this has been mentioned as occurring in Cakile
maritima, wherein the sepals were found by M. Fournier to be pinnatifid
like the ordinary leaves of the plant.[255] The sepals of Ranunculaceæ and
Rosaceæ, for example, Rosa, Geum, are particularly liable to this change.
Fig. 129.—Flower of rose, sepals replaced by five perfect leaves; axis prolonged through the flower
in the form of a leafy branch.
In a species of Geranium recently examined the sepals presented
themselves in the form of three-lobed leaflets; so in fuchsias and in
Epilobium hirsutum the sepals occasionally are not distinguishable from
Page 317
ordinary leaves (fig. 130). In roses, the change in question is a very frequent
accompaniment of prolification (fig. 129). In the peach also this
replacement of the sepals is sometimes carried to such an extent, that five
perfect, bistipulate leaves occur in the place of the calyx, but when this is
the case it usually happens that the pistil is abortive.
Fig. 130.—Fuchsia, with one of the sepals leaf-like.
De Candolle[256] figures a curious instance wherein the pappus of
Podospermum laciniatum was replaced by five linear, foliaceous lobes. A
similar change has been noticed in other composites, as in Tragopogon
pratense. Engelmann mentions as subject to this hypertrophy of the pappus,
as it may be termed, Scorzonera octangularis and Senecio vulgaris. Wigand
has observed a similar transformation in a species of Centranthus
(Valerianaceæ).
accompaniment of prolification (fig. 129). In the peach also this
replacement of the sepals is sometimes carried to such an extent, that five
perfect, bistipulate leaves occur in the place of the calyx, but when this is
the case it usually happens that the pistil is abortive.
Fig. 130.—Fuchsia, with one of the sepals leaf-like.
De Candolle[256] figures a curious instance wherein the pappus of
Podospermum laciniatum was replaced by five linear, foliaceous lobes. A
similar change has been noticed in other composites, as in Tragopogon
pratense. Engelmann mentions as subject to this hypertrophy of the pappus,
as it may be termed, Scorzonera octangularis and Senecio vulgaris. Wigand
has observed a similar transformation in a species of Centranthus
(Valerianaceæ).
Page 318
In some cases the phyllody of the sepals has a special interest, as bearing on
the question whether what is termed calyx-tube is or is not a portion of the
calyx, and whether the sepals are modifications of the blade or of the sheath
of the leaf. Thus in the primrose the phyllodic sepals seem to show clearly
that the sepals are in that plant of a laminar nature (fig. 131). The so-called
calyx-tube of roses is elsewhere alluded to. The leaf-like organs sometimes
seen at the apex of a cucumber would seem to support the view that there
was really a calyx-tube in Cucurbitaceæ adherent to the carpels. It is also
shown in the cut, fig. 132, borrowed from the 'Gardeners' Chronicle,' 1859,
p. 654.
Fig. 131.—Primrose. Calyx of foliaceous segments.
the question whether what is termed calyx-tube is or is not a portion of the
calyx, and whether the sepals are modifications of the blade or of the sheath
of the leaf. Thus in the primrose the phyllodic sepals seem to show clearly
that the sepals are in that plant of a laminar nature (fig. 131). The so-called
calyx-tube of roses is elsewhere alluded to. The leaf-like organs sometimes
seen at the apex of a cucumber would seem to support the view that there
was really a calyx-tube in Cucurbitaceæ adherent to the carpels. It is also
shown in the cut, fig. 132, borrowed from the 'Gardeners' Chronicle,' 1859,
p. 654.
Fig. 131.—Primrose. Calyx of foliaceous segments.
Page 319
Fig. 132.—Leafy calyx of melon.
Under ordinary circumstances, the sepals may be considered as the
representatives of the sheath of the leaf (cataphyllary) or of the blade
(euphyllary), the arrangement of the veins being different in the two cases;
thus, in the vagina or sheath, there are generally several large veins of about
equal size, either convergent towards the apex, or divergent; on the other
hand, in the blade, there is usually but one central vein, the midrib, larger
than the rest, and the smaller veins come off at a less acute angle, and are
more reticulated.[257]
Now, when phyllomorphy occurs in sepals which ordinarily are vaginal, it
is obvious that the case is one, not merely of increased relative growth, but
also of the appearance or development of an organ habitually suppressed;
on the other hand, when phyllomorphy occurs in sepals which usually are
laminar in form and nervation, the case is one of unusual growth or
Under ordinary circumstances, the sepals may be considered as the
representatives of the sheath of the leaf (cataphyllary) or of the blade
(euphyllary), the arrangement of the veins being different in the two cases;
thus, in the vagina or sheath, there are generally several large veins of about
equal size, either convergent towards the apex, or divergent; on the other
hand, in the blade, there is usually but one central vein, the midrib, larger
than the rest, and the smaller veins come off at a less acute angle, and are
more reticulated.[257]
Now, when phyllomorphy occurs in sepals which ordinarily are vaginal, it
is obvious that the case is one, not merely of increased relative growth, but
also of the appearance or development of an organ habitually suppressed;
on the other hand, when phyllomorphy occurs in sepals which usually are
laminar in form and nervation, the case is one of unusual growth or
Page 320
hypertrophy, and not of the development of an organ habitually suppressed,
so that the amount of change is greater in the former than in the latter
instance.
Under normal circumstances it will be found that laminar venation is most
common in gamosepalous and vaginal venation in polysepalous calyces.
And the same holds good in cases where the calyx is abnormally leafy. The
complete leaf development shows itself more frequently among the
monosepalous plants than in the polysepalous ones, as shown even in the
subjoined list of species. This statement would be more fully verified were
it possible to state the frequency with which the condition occurred in
individual plants, when it would be found that phyllody of the calyx occurs
much more often in individual gamosepalous plants than in polysepalous
ones.
Phyllody of the calyx has been most often observed in the following plants:
Ranunculus acris!
Delphinium Ajacis.
Caltha palustris.
Anemone Pulsatilla.
sylvestris!
nemorosa!
hortensis!
coronaria!
*Papaver orientale.
Escholtzia crocea.
Cakile maritima.
Diplotaxis tenuifolia.
Thlaspi arvense.
Cheiranthus Cheiri.
incanus.
Sinapis arvensis.
Brassica oleracea!
Peltaria alliacea.
*Sisymbrium officinale.
so that the amount of change is greater in the former than in the latter
instance.
Under normal circumstances it will be found that laminar venation is most
common in gamosepalous and vaginal venation in polysepalous calyces.
And the same holds good in cases where the calyx is abnormally leafy. The
complete leaf development shows itself more frequently among the
monosepalous plants than in the polysepalous ones, as shown even in the
subjoined list of species. This statement would be more fully verified were
it possible to state the frequency with which the condition occurred in
individual plants, when it would be found that phyllody of the calyx occurs
much more often in individual gamosepalous plants than in polysepalous
ones.
Phyllody of the calyx has been most often observed in the following plants:
Ranunculus acris!
Delphinium Ajacis.
Caltha palustris.
Anemone Pulsatilla.
sylvestris!
nemorosa!
hortensis!
coronaria!
*Papaver orientale.
Escholtzia crocea.
Cakile maritima.
Diplotaxis tenuifolia.
Thlaspi arvense.
Cheiranthus Cheiri.
incanus.
Sinapis arvensis.
Brassica oleracea!
Peltaria alliacea.
*Sisymbrium officinale.
Page 321
Caryophyllaceæ,[258] sp. pl.
Geranium, sp.!
*Fuchsia, var. hort.!
Epilobium hirsutum!
Cucurbita Pepo!
*Rosa, var. hort.!
Potentilla nepalensis.
Fragaria sp.
Geum rivale.
Amygdalus communis.
Persica vulgaris.
Cerasus!
Pyrus Malus.
Daucus Carota.
Athamanta Cervaria.
*Trifolium repens!
Centranthus macrosiphon.
Tragopogon pratense.
orientale.
Scorzonera octangularis.
Hypochæris radicata.
*Senecio vulgaris!
Podospermum laciniatum.
Cirsium arvense.
Carduus heterophyllus
tataricus.
Campanula, sp.
Convolvulus sepium.
*Primula officinalis, var. cult!
acaulis.
elatior.
Gentiana campestris.
*Petunia violacea!
Lycium europæum.
Laurus Sassafras.
Tulipa Gesneriana.
Convallaria maialis.
Colchicum autumnale! (virescent?)
Geranium, sp.!
*Fuchsia, var. hort.!
Epilobium hirsutum!
Cucurbita Pepo!
*Rosa, var. hort.!
Potentilla nepalensis.
Fragaria sp.
Geum rivale.
Amygdalus communis.
Persica vulgaris.
Cerasus!
Pyrus Malus.
Daucus Carota.
Athamanta Cervaria.
*Trifolium repens!
Centranthus macrosiphon.
Tragopogon pratense.
orientale.
Scorzonera octangularis.
Hypochæris radicata.
*Senecio vulgaris!
Podospermum laciniatum.
Cirsium arvense.
Carduus heterophyllus
tataricus.
Campanula, sp.
Convolvulus sepium.
*Primula officinalis, var. cult!
acaulis.
elatior.
Gentiana campestris.
*Petunia violacea!
Lycium europæum.
Laurus Sassafras.
Tulipa Gesneriana.
Convallaria maialis.
Colchicum autumnale! (virescent?)
Page 322
Consult also Turpin, 'Atlas de Goethe,' t. iv, f. 12, Lycium. Engelmann, 'De Anthol.,' § 35,
p. 31. This author figures phyllodic sepals in Senecio vulgaris, tab. v, figs. 24–26;
Campanula, tab. iii, f. 15, 16; Athamanta cervaria, tab. v, f. 14. Lindley, 'Elements of
Botany,' 1847, pp. 64, 73, &c. 'Gard. Chron.,' 1858, p. 685; 1859, p. 654, Cucurbita.
Petunnikoff, 'Bull. Soc. Imp. Moscow,' 1862, Cirsium. Braun, 'Rejuvenescence,' Ray
Society's Transl. See succeeding paragraphs.
Phyllody of the corolla.—The petals also are frequently replaced by
leaves, though in many of the recorded instances the change has been one
of colour only; these latter are strictly cases of virescence. M. Seringe[259]
speaks of a flower of Peltaria alliacea in which the calyx was petal-like,
while the corolla was leafy as if there had been transposition of the two
organs, a very rare, if not unparalleled, instance. In a flower of Campanula
Medium, provided, as is often the case, with a double corolla, the outer
corolla was slit down on one side, the edges of the cleft being leafy.
Fig. 133.—Sepals and petals to leaves. Geranium.
The frondescent petals are very often completely disjoined, as in Verbascum
nigrum, and Lonicera Periclymenum, in which, moreover, median
prolification generally coexists. In the case of Tropæolum majus, the
ordinary leaves of which are peltate and orbicular, the petals when
frondescent have not the peltate arrangement, but are spathulate, and
provided with very long, narrow stalks, so that, in some cases, they are,
more properly speaking, enlarged virescent petals than true leaves; in other
p. 31. This author figures phyllodic sepals in Senecio vulgaris, tab. v, figs. 24–26;
Campanula, tab. iii, f. 15, 16; Athamanta cervaria, tab. v, f. 14. Lindley, 'Elements of
Botany,' 1847, pp. 64, 73, &c. 'Gard. Chron.,' 1858, p. 685; 1859, p. 654, Cucurbita.
Petunnikoff, 'Bull. Soc. Imp. Moscow,' 1862, Cirsium. Braun, 'Rejuvenescence,' Ray
Society's Transl. See succeeding paragraphs.
Phyllody of the corolla.—The petals also are frequently replaced by
leaves, though in many of the recorded instances the change has been one
of colour only; these latter are strictly cases of virescence. M. Seringe[259]
speaks of a flower of Peltaria alliacea in which the calyx was petal-like,
while the corolla was leafy as if there had been transposition of the two
organs, a very rare, if not unparalleled, instance. In a flower of Campanula
Medium, provided, as is often the case, with a double corolla, the outer
corolla was slit down on one side, the edges of the cleft being leafy.
Fig. 133.—Sepals and petals to leaves. Geranium.
The frondescent petals are very often completely disjoined, as in Verbascum
nigrum, and Lonicera Periclymenum, in which, moreover, median
prolification generally coexists. In the case of Tropæolum majus, the
ordinary leaves of which are peltate and orbicular, the petals when
frondescent have not the peltate arrangement, but are spathulate, and
provided with very long, narrow stalks, so that, in some cases, they are,
more properly speaking, enlarged virescent petals than true leaves; in other
Page 323
instances, however, the arrangement of the veins is more like that of the true
leaves than that of the petals.
As might be expected, frondescence of the petals is frequently accompanied
by other changes of a similar nature in other parts of the flower, and
sometimes by the abortion of the sexual organs. Thus, in Actæa spicata, as
observed by Fresenius, the petals were replaced by true petiolate,
palminerved, lobed leaves, the stamens and pistils being abortive. In
Ranunculus the leaves that appear in the place of the petals have no scale at
their base, and in Tropæolum the calyx (or receptacle) is free from the usual
spur.
The absolute frequency of this occurrence seems to be greatest in those
flowers which are normally polypetalous. The petals of these flowers, as a
general rule, are more like the leaf-sheaths than the leaf-blades as to their
venation, hence it would seem that the phyllomorphic condition in these
petals is a manifestation of a greater degree of organizing force than that
which occurs in those cases where the petals are normally present in the
form of contracted blades or laminæ. (See the remarks in the preceding
section.)
Frondescence of the petals has been observed most frequently in the
following cases; some, perhaps, were cases merely of virescence, q. v.; see
also under Chloranthy, Prolification.
Ranunculus repens!
Delphinium Ajacis.
crassicaule.
Aquilegia vulgaris.
Actæa spicata.
*Brassica oleracea!
Diplotaxis muralis.
Hesperis matronalis.
Thlaspi bursa pastoris.
leaves than that of the petals.
As might be expected, frondescence of the petals is frequently accompanied
by other changes of a similar nature in other parts of the flower, and
sometimes by the abortion of the sexual organs. Thus, in Actæa spicata, as
observed by Fresenius, the petals were replaced by true petiolate,
palminerved, lobed leaves, the stamens and pistils being abortive. In
Ranunculus the leaves that appear in the place of the petals have no scale at
their base, and in Tropæolum the calyx (or receptacle) is free from the usual
spur.
The absolute frequency of this occurrence seems to be greatest in those
flowers which are normally polypetalous. The petals of these flowers, as a
general rule, are more like the leaf-sheaths than the leaf-blades as to their
venation, hence it would seem that the phyllomorphic condition in these
petals is a manifestation of a greater degree of organizing force than that
which occurs in those cases where the petals are normally present in the
form of contracted blades or laminæ. (See the remarks in the preceding
section.)
Frondescence of the petals has been observed most frequently in the
following cases; some, perhaps, were cases merely of virescence, q. v.; see
also under Chloranthy, Prolification.
Ranunculus repens!
Delphinium Ajacis.
crassicaule.
Aquilegia vulgaris.
Actæa spicata.
*Brassica oleracea!
Diplotaxis muralis.
Hesperis matronalis.
Thlaspi bursa pastoris.
Page 324
Sisymbrium tenuifolium.
Turritis glabra.
Raphanus sativus.
Peltaria alliacea.
Alyssum incanum.
Erysimum Barbarea.
officinale!
cheiranthoides.
Cheiranthus Cheiri.
*Dictamnus Fraxinella!
Lychnis sylvestris.
dioica!
Alsine media.
Cerastium vulgatum!
triviale.
Reseda lutea.
Phyteuma.
Malva sylvestris.
*Tropæolum majus!
Geranium, sp.!
Triumfetta, sp.!
Epilobium hirsutum!
Œnothera striata.
Rubus, sp.
*Rosa, var. cult.!
*Trifolium repens!
Spiræa oblongifolia.
Amygdalus communis.
*Rosa!
Cerasus vulgaris!
Persica vulgaris!
Potentilla nepalensis.
Geum rivale.
Daucus Carota!
Heracleum Sphondylium.
Torilis Anthriscus.
Echinophora maritima.
Campanula rapunculoides.
glomerata.
Turritis glabra.
Raphanus sativus.
Peltaria alliacea.
Alyssum incanum.
Erysimum Barbarea.
officinale!
cheiranthoides.
Cheiranthus Cheiri.
*Dictamnus Fraxinella!
Lychnis sylvestris.
dioica!
Alsine media.
Cerastium vulgatum!
triviale.
Reseda lutea.
Phyteuma.
Malva sylvestris.
*Tropæolum majus!
Geranium, sp.!
Triumfetta, sp.!
Epilobium hirsutum!
Œnothera striata.
Rubus, sp.
*Rosa, var. cult.!
*Trifolium repens!
Spiræa oblongifolia.
Amygdalus communis.
*Rosa!
Cerasus vulgaris!
Persica vulgaris!
Potentilla nepalensis.
Geum rivale.
Daucus Carota!
Heracleum Sphondylium.
Torilis Anthriscus.
Echinophora maritima.
Campanula rapunculoides.
glomerata.
Page 325
Phyteuma spicatum.
Calendula officinalis.
Cirsium tricephalodes.
Senecio vulgaris.
Scabiosa columbaria.
agrestis.
Lonicera xylosteum.
Periclymenum.
Gentiana Amarella.
Gilia glomeriflora.
*Symphytum officinale.
Petunia violacea!
Verbascum, sp.
Antirrhinum majus!
Stachys sylvatica.
*Anagallis phœnicea?
Primula sinensis!
Polemonium cœruleum.
See Moquin-Tandon, 'El. Terat. Veg.,' p. 203. Engelmann, 'De Anthol.,' § 38 et seq.; tab. ii,
figs. 8–14, Gilia; tab. v, 23–26, Senecio; tab. v, f. 1–13, Torilis; tab. iv, f. 3, Erysimum.
'Bull. Soc. Bot. Fr.,' vol. ii, 1855, p. 479, Primula sinensis. Giraud, 'Edinb. Phil. Magazine,'
1839, Antirrhinum. Jaeger, 'Act. Acad. Cæs. Nat. Cur.,' vol. xiii, 2, p. 1, tab. xli,
Tropæolum. Bischoff, 'Lehrbuch,' 11, 2, p. 27, note, Tropæolum. Fresenius, 'Mus.
Senkenb.,' ii, 35, tab. 4, fig. 5, Actæa. See also succeeding paragraphs and sections in
Chloranthy, Virescence, &c.
Phyllody of the stamens happens less frequently than the corresponding
condition in the neighbouring organs. The structure of the anther is so much
removed from that of the leaf, that the change of the stamen from its
ordinary condition to that of a leaf must be regarded as indicating a greater
degree of perverted development than that which occurs in those cases
where less highly differentiated organs, such as the sepals, petals, and
pistils, are thus altered.[260]
In all cases it is desirable to ascertain, if possible, what parts of the stamen
are thus transformed. In some Petunias the filaments are unchanged, but in
Calendula officinalis.
Cirsium tricephalodes.
Senecio vulgaris.
Scabiosa columbaria.
agrestis.
Lonicera xylosteum.
Periclymenum.
Gentiana Amarella.
Gilia glomeriflora.
*Symphytum officinale.
Petunia violacea!
Verbascum, sp.
Antirrhinum majus!
Stachys sylvatica.
*Anagallis phœnicea?
Primula sinensis!
Polemonium cœruleum.
See Moquin-Tandon, 'El. Terat. Veg.,' p. 203. Engelmann, 'De Anthol.,' § 38 et seq.; tab. ii,
figs. 8–14, Gilia; tab. v, 23–26, Senecio; tab. v, f. 1–13, Torilis; tab. iv, f. 3, Erysimum.
'Bull. Soc. Bot. Fr.,' vol. ii, 1855, p. 479, Primula sinensis. Giraud, 'Edinb. Phil. Magazine,'
1839, Antirrhinum. Jaeger, 'Act. Acad. Cæs. Nat. Cur.,' vol. xiii, 2, p. 1, tab. xli,
Tropæolum. Bischoff, 'Lehrbuch,' 11, 2, p. 27, note, Tropæolum. Fresenius, 'Mus.
Senkenb.,' ii, 35, tab. 4, fig. 5, Actæa. See also succeeding paragraphs and sections in
Chloranthy, Virescence, &c.
Phyllody of the stamens happens less frequently than the corresponding
condition in the neighbouring organs. The structure of the anther is so much
removed from that of the leaf, that the change of the stamen from its
ordinary condition to that of a leaf must be regarded as indicating a greater
degree of perverted development than that which occurs in those cases
where less highly differentiated organs, such as the sepals, petals, and
pistils, are thus altered.[260]
In all cases it is desirable to ascertain, if possible, what parts of the stamen
are thus transformed. In some Petunias the filaments are unchanged, but in
Page 326
place of the anther is a small lamina, representing precisely the blade of an
ordinary leaf. Sometimes the connective only is replaced by a leaf. One of
the most interesting cases of this kind that has fallen under the writer's
observation was in Euphorbia geniculata, in which, in addition to other
changes mentioned under prolification of the inflorescence, some of the
stamens were partly frondescent, half the anther being perfect, the other half
leaf-like. Another filament bore just above the usual joint three leaflets, two
lateral ones, somewhat conduplicate, and a third central one, half anther,
half leaflet.
Fig. 134.—Flower of a Petunia, opened to show the stamens partially replaced by stalked leaves.
In the case of frondescent flowers of Tropæolum majus the stamens are
usually absent or atrophied, but in other instances the filament is present as
usual, representing the stalk of the leaf, and surmounted by a small lamina,
but this latter, in place of being nearly flat, is pinched up in the centre from
back to front, and surmounted by a two-lobed anther, so that the general
ordinary leaf. Sometimes the connective only is replaced by a leaf. One of
the most interesting cases of this kind that has fallen under the writer's
observation was in Euphorbia geniculata, in which, in addition to other
changes mentioned under prolification of the inflorescence, some of the
stamens were partly frondescent, half the anther being perfect, the other half
leaf-like. Another filament bore just above the usual joint three leaflets, two
lateral ones, somewhat conduplicate, and a third central one, half anther,
half leaflet.
Fig. 134.—Flower of a Petunia, opened to show the stamens partially replaced by stalked leaves.
In the case of frondescent flowers of Tropæolum majus the stamens are
usually absent or atrophied, but in other instances the filament is present as
usual, representing the stalk of the leaf, and surmounted by a small lamina,
but this latter, in place of being nearly flat, is pinched up in the centre from
back to front, and surmounted by a two-lobed anther, so that the general
Page 327
appearance of the whole structure is that of a central anther, supported at the
base on each side by two concave leaf-lobes, or it might be compared with a
three-lobed leaf, the terminal lobe represented by the anther.
In Jatropha Pohliana, Müll. (Adenorophium luxurians, Pohl.), a singular
condition has been observed by M. Müller (Argov.). In this flower the
anther, in place of being represented by the flat blade of a single leaf, had
the appearance as if two such blades were present and coherent one with the
other by their midribs, along their upper or inner surfaces, which were
directed towards the centre of the flower (fig. 136), thus resembling the
cases of adhesion of leaves by their surfaces already referred to (p. 33). In
other cases, in the same plant, the anther appeared as if formed by two
collateral leaves, the faces looking towards the circumference of the flower,
and their margins so folded together as to represent an open anther lobe (fig.
135). These cases are apparently due, not to the formation and adhesion of
two leaves, but rather to the exuberant development of one leaf into two
blades.[261] The bearings of these and other similar malformations on the
morphology of the anther are alluded to under the head of petalody of the
anther.
base on each side by two concave leaf-lobes, or it might be compared with a
three-lobed leaf, the terminal lobe represented by the anther.
In Jatropha Pohliana, Müll. (Adenorophium luxurians, Pohl.), a singular
condition has been observed by M. Müller (Argov.). In this flower the
anther, in place of being represented by the flat blade of a single leaf, had
the appearance as if two such blades were present and coherent one with the
other by their midribs, along their upper or inner surfaces, which were
directed towards the centre of the flower (fig. 136), thus resembling the
cases of adhesion of leaves by their surfaces already referred to (p. 33). In
other cases, in the same plant, the anther appeared as if formed by two
collateral leaves, the faces looking towards the circumference of the flower,
and their margins so folded together as to represent an open anther lobe (fig.
135). These cases are apparently due, not to the formation and adhesion of
two leaves, but rather to the exuberant development of one leaf into two
blades.[261] The bearings of these and other similar malformations on the
morphology of the anther are alluded to under the head of petalody of the
anther.
Page 328
Fig. 135.—Phylloid anther of Jatropha, after Müller (Arg.).
Page 329
Fig. 136.—Leaf-like anther of Jatropha Pohliana, after Müller.
Phyllody of the stamens has been most often observed in the following
plants:
Anemone nemorosa.
coronaria.
Delphinium crassicaule.
Nymphæa dentata.
Tropæolum majus!
Dictamnus albus.
*Trifolium repens!
Torilis anthriscus.
Heracleum Sphondylium.
Daucus Carota
Epilobium hirsutum!
*Rosa, var. cult.!
Lonicera Periclymenum.
Anagallis arvensis.
Primula sinensis!
Petunia, var. cult.
Jatropha Pohliana.
Euphorbia goniculata.
In addition to the foregoing there are very numerous instances of similar
substitution in chloranthic flowers. In the above list only those cases are
given wherein the leafy change is confined to the stamens, or, at least, to a
few only of the other parts of the flower.
Phyllody of the pistils.[262]—This is of more common occurrence than is
the corresponding change in the case of the stamens. It is of interest, as it
sometimes serves to illustrate the morphological nature of the pistil. Of this
the double-flowering cherry is a well-known illustration, the pistil being
here represented by two small foliar laminæ, whose midribs are prolonged
with a short style, terminated by an imperfect stigma. It is usually the basal
portion of the pistil, the ovary, which is thus specially affected, the margins
Phyllody of the stamens has been most often observed in the following
plants:
Anemone nemorosa.
coronaria.
Delphinium crassicaule.
Nymphæa dentata.
Tropæolum majus!
Dictamnus albus.
*Trifolium repens!
Torilis anthriscus.
Heracleum Sphondylium.
Daucus Carota
Epilobium hirsutum!
*Rosa, var. cult.!
Lonicera Periclymenum.
Anagallis arvensis.
Primula sinensis!
Petunia, var. cult.
Jatropha Pohliana.
Euphorbia goniculata.
In addition to the foregoing there are very numerous instances of similar
substitution in chloranthic flowers. In the above list only those cases are
given wherein the leafy change is confined to the stamens, or, at least, to a
few only of the other parts of the flower.
Phyllody of the pistils.[262]—This is of more common occurrence than is
the corresponding change in the case of the stamens. It is of interest, as it
sometimes serves to illustrate the morphological nature of the pistil. Of this
the double-flowering cherry is a well-known illustration, the pistil being
here represented by two small foliar laminæ, whose midribs are prolonged
with a short style, terminated by an imperfect stigma. It is usually the basal
portion of the pistil, the ovary, which is thus specially affected, the margins
Page 330
being also often disunited so as to expose the ovules. These latter organs
may be absent or they may themselves be the subjects of foliaceous
development. Moquin[263] relates having found in the neighbourhood of
Montpellier a flower of a tulip the ovary of which was represented by true
leaves, which bore on their margins the ovules, and thus presented a striking
analogy with the carpels of those Sterculias, like S. platanifolia, which are
foliaceous in texture and open very early in the course of their development.
A similar occurrence has also been frequently noticed in the Columbine and
also in Cruciferæ and Umbelliferæ. M. Germain de St. Pierre mentions an
instance wherein the carpels of Salix Babylonica were converted into two
leaves, provided with stipules. All the flowers of the catkins were similarly
changed, so that it became permanent, and resembled a branch.
Fig. 137.—Rose, in which the axial portion of the flower was elongated and the carpels were more or
less replaced by leaves.
may be absent or they may themselves be the subjects of foliaceous
development. Moquin[263] relates having found in the neighbourhood of
Montpellier a flower of a tulip the ovary of which was represented by true
leaves, which bore on their margins the ovules, and thus presented a striking
analogy with the carpels of those Sterculias, like S. platanifolia, which are
foliaceous in texture and open very early in the course of their development.
A similar occurrence has also been frequently noticed in the Columbine and
also in Cruciferæ and Umbelliferæ. M. Germain de St. Pierre mentions an
instance wherein the carpels of Salix Babylonica were converted into two
leaves, provided with stipules. All the flowers of the catkins were similarly
changed, so that it became permanent, and resembled a branch.
Fig. 137.—Rose, in which the axial portion of the flower was elongated and the carpels were more or
less replaced by leaves.
Page 331
Substitutions of this kind form the green "eyes" or centres of certain
varieties of Ranunculus and Anemone.
In proliferous roses, or in cases where the central axis of the flower is
prolonged, it frequently happens that the pistils are more or less replaced by
leaves. Fig. 137, from a specimen of Dr. Bell Salter's, given in the
'Gardeners' Chronicle,' shows the passage, from below upwards, of the
ordinary carpels to perfect leaves; the so-called calyx-tube being completely
deficient and the ovaries entirely superior. Like most similar specimens, this
one bears out the notion that what is called the calyx-tube in roses is really
an expansion and dilatation of the top of the flower-stalk.
Fig. 138.—Cucumber with leaf attached.
Fig. 138, for which I am indebted to Mr. S. J. Salter, represents a very
singular conformation in the cucumber, described by that gentleman in
'Henfrey's Botanical Gazette,' i, p. 208, and considered by him to be due to
varieties of Ranunculus and Anemone.
In proliferous roses, or in cases where the central axis of the flower is
prolonged, it frequently happens that the pistils are more or less replaced by
leaves. Fig. 137, from a specimen of Dr. Bell Salter's, given in the
'Gardeners' Chronicle,' shows the passage, from below upwards, of the
ordinary carpels to perfect leaves; the so-called calyx-tube being completely
deficient and the ovaries entirely superior. Like most similar specimens, this
one bears out the notion that what is called the calyx-tube in roses is really
an expansion and dilatation of the top of the flower-stalk.
Fig. 138.—Cucumber with leaf attached.
Fig. 138, for which I am indebted to Mr. S. J. Salter, represents a very
singular conformation in the cucumber, described by that gentleman in
'Henfrey's Botanical Gazette,' i, p. 208, and considered by him to be due to
Page 332
the foliaceous condition of one of the three carpels of which the fruit is
composed. The portion near the peduncle was binary, while the distal
extremity of the fruit was ternary. The main difficulties attending the
acceptance of this explanation reside in the peculiar reversed position of the
leaf, and in the fact that the fruit of the Cucurbitaceæ is probably of axial
nature, the dilated and succulent end of the peduncle adhering to and
usually concealing the carpels; in some cases, however, these latter project
beyond the axial portion, leaving no doubt as to the true nature of the
structure in these particular instances.
Admitting the axial nature of the fruit, it might be supposed that in Mr.
Salter's cucumber an adventitious leaf had been given off from the axis, but
even on that supposition the reversed position offers a difficulty, and there
still remains to be explained the fact that the proximal part of the fruit was
binary in its constitution, the distal end ternary.
M. Norman[264] mentions a case wherein the carpels of Anchusa ochroleuca
were replaced by two leaves; from this he draws the inference that the pistil
of borages and labiates is really composed of two leaves, placed fore and
aft, the margins of the leaves being congenitally fused. This tallies well with
the account given of the development of these plants by Payer, Germain de
St. Pierre, and others.
In an Indian species of Triumfetta, not only were the petals virescent, but
the ovary also was much enlarged, and in some flowers it was divided half
way down into five lanceolate leaves (fig. 139), the sepals and stamens
being in their normal condition.
In the preceding instances the foliaceous condition has pervaded the entire
pistil, or at any rate the basal portion or ovary, and it may be noticed that
composed. The portion near the peduncle was binary, while the distal
extremity of the fruit was ternary. The main difficulties attending the
acceptance of this explanation reside in the peculiar reversed position of the
leaf, and in the fact that the fruit of the Cucurbitaceæ is probably of axial
nature, the dilated and succulent end of the peduncle adhering to and
usually concealing the carpels; in some cases, however, these latter project
beyond the axial portion, leaving no doubt as to the true nature of the
structure in these particular instances.
Admitting the axial nature of the fruit, it might be supposed that in Mr.
Salter's cucumber an adventitious leaf had been given off from the axis, but
even on that supposition the reversed position offers a difficulty, and there
still remains to be explained the fact that the proximal part of the fruit was
binary in its constitution, the distal end ternary.
M. Norman[264] mentions a case wherein the carpels of Anchusa ochroleuca
were replaced by two leaves; from this he draws the inference that the pistil
of borages and labiates is really composed of two leaves, placed fore and
aft, the margins of the leaves being congenitally fused. This tallies well with
the account given of the development of these plants by Payer, Germain de
St. Pierre, and others.
In an Indian species of Triumfetta, not only were the petals virescent, but
the ovary also was much enlarged, and in some flowers it was divided half
way down into five lanceolate leaves (fig. 139), the sepals and stamens
being in their normal condition.
In the preceding instances the foliaceous condition has pervaded the entire
pistil, or at any rate the basal portion or ovary, and it may be noticed that
Page 333
the ovary is thus shown to consist in some cases of the sheath of the leaf, as
in Aquilegia; in other cases of the blade, as in Cerasus, Daucus, &c.
Fig. 139.—Flower of Triumfetta, sp., carpels represented by five leaves.
There are cases, however, in which a part only of the pistillary structure
thus becomes foliaceous. Linnæus, 'Prolepsis,' § 9, mentions some flowers
of Carduus heterophyllus and C. tataricus in which the style had grown into
two green leaflets, and in which the calyx and corolla were also leaf-like. A
very singular instance is recorded by Baillon,[265] wherein the pistil of
Trifolium repens consisted of three carpels, either separate, or combined so
as to form a one-celled ovary with three parietal, pluri-ovulate placentæ; the
ovary in these flowers was formed of the basal vaginiform part of the leaf;
the three styles were formed by the petioles, while the stigmas were
represented by trifoliolate leaves. The back of the leaf in these cases is
usually directed away from the centre of the flower. When this change
in Aquilegia; in other cases of the blade, as in Cerasus, Daucus, &c.
Fig. 139.—Flower of Triumfetta, sp., carpels represented by five leaves.
There are cases, however, in which a part only of the pistillary structure
thus becomes foliaceous. Linnæus, 'Prolepsis,' § 9, mentions some flowers
of Carduus heterophyllus and C. tataricus in which the style had grown into
two green leaflets, and in which the calyx and corolla were also leaf-like. A
very singular instance is recorded by Baillon,[265] wherein the pistil of
Trifolium repens consisted of three carpels, either separate, or combined so
as to form a one-celled ovary with three parietal, pluri-ovulate placentæ; the
ovary in these flowers was formed of the basal vaginiform part of the leaf;
the three styles were formed by the petioles, while the stigmas were
represented by trifoliolate leaves. The back of the leaf in these cases is
usually directed away from the centre of the flower. When this change
Page 334
occurs it is commonly attended by an increased number of parts, as in the
trefoil just mentioned, or in the double cherry, where usually two foliaceous
carpels may be met with, and sometimes more.
The change is also of interest when it affects such orders as the
Umbelliferæ, which have their ovaries inferior under ordinary
circumstances; but when these organs assume a leafy condition they
become superior also, i.e. they are detached from the calyx.
As regards the position of the ovules in these foliaceous pistils, they may be
placed, as in Aquilegia, Delphinium, &c., on the edges of the carpel or on
the surface, as in some flowers of Ranunculus repens and R. Ficaria. A
similar position of the ovules is recorded in the case of the vine (Vitis),
where the pistil consisted of leaves bearing the ovules on their inner
surface.[266] The supposed causes of this and other similar malformations
are alluded to under the head of chloranthy, but it may be here remarked
that semi-double flowers, fertilised by the pollen of similar flowers, are said
to produce flowers with a centre of small green leaves, this central tuft
resulting from the expansion and frondescence of the pistils.
As this condition rarely occurs without corresponding changes in other
parts of the flower, further remarks on this subject will be found in the
chapter relating to Chloranthy.
Phyllody of the pistil has been most frequently recorded in the following
plants:
Pæonia officinalis.
Ranunculus repens!
*Aquilegia vulgaris!
Delphinium elatum.
crassicaule.
Ajacis.
trefoil just mentioned, or in the double cherry, where usually two foliaceous
carpels may be met with, and sometimes more.
The change is also of interest when it affects such orders as the
Umbelliferæ, which have their ovaries inferior under ordinary
circumstances; but when these organs assume a leafy condition they
become superior also, i.e. they are detached from the calyx.
As regards the position of the ovules in these foliaceous pistils, they may be
placed, as in Aquilegia, Delphinium, &c., on the edges of the carpel or on
the surface, as in some flowers of Ranunculus repens and R. Ficaria. A
similar position of the ovules is recorded in the case of the vine (Vitis),
where the pistil consisted of leaves bearing the ovules on their inner
surface.[266] The supposed causes of this and other similar malformations
are alluded to under the head of chloranthy, but it may be here remarked
that semi-double flowers, fertilised by the pollen of similar flowers, are said
to produce flowers with a centre of small green leaves, this central tuft
resulting from the expansion and frondescence of the pistils.
As this condition rarely occurs without corresponding changes in other
parts of the flower, further remarks on this subject will be found in the
chapter relating to Chloranthy.
Phyllody of the pistil has been most frequently recorded in the following
plants:
Pæonia officinalis.
Ranunculus repens!
*Aquilegia vulgaris!
Delphinium elatum.
crassicaule.
Ajacis.
Page 335
amænum.
Nymphæa dentata.
Sinapis arvensis!
Diplotaxis tenuifolia.
*Brassica oleracea!
*Sisymbrium officinale!
Dianthus. sp
Reseda Phyteuma.
Triumfetta, sp.!
Lychnis dioica.
Cerastium, sp.!
*Dictamnus Fraxinella!
Cerasus avium.
vulgaris!
*Rosa, var. cult.!
*Daucus Carota!
Heracleum, sp.
Epilobium hirsutum!
Lathyrus latifolius.
*Trifolium repens!
hybridum.
Melilotus, sp.
Medicago, sp.
Lonicera Periclymenum.
Carduus heterophyllus.
tataricus.
Scrophularia aquatica.
Symphytum officinale.
Anchusa ochroleuca.
paniculata.
*Primula sinensis!
Salix babylonica.
Hyacinthus, sp.
Tulipa, sp.
Some of the above are probably cases of mere virescence rather than of
phyllody. For further illustrations, references to authorities, &c., see under
Chloranthy, Virescence, Prolification, &c.
Nymphæa dentata.
Sinapis arvensis!
Diplotaxis tenuifolia.
*Brassica oleracea!
*Sisymbrium officinale!
Dianthus. sp
Reseda Phyteuma.
Triumfetta, sp.!
Lychnis dioica.
Cerastium, sp.!
*Dictamnus Fraxinella!
Cerasus avium.
vulgaris!
*Rosa, var. cult.!
*Daucus Carota!
Heracleum, sp.
Epilobium hirsutum!
Lathyrus latifolius.
*Trifolium repens!
hybridum.
Melilotus, sp.
Medicago, sp.
Lonicera Periclymenum.
Carduus heterophyllus.
tataricus.
Scrophularia aquatica.
Symphytum officinale.
Anchusa ochroleuca.
paniculata.
*Primula sinensis!
Salix babylonica.
Hyacinthus, sp.
Tulipa, sp.
Some of the above are probably cases of mere virescence rather than of
phyllody. For further illustrations, references to authorities, &c., see under
Chloranthy, Virescence, Prolification, &c.
Page 336
Phyllody of the ovules.—Pending the settlement of the existing differences
of opinion with reference to the morphological nature of the ovule and its
component parts, much interest attaches to the malformations to which they
are occasionally subject. Considered purely in a teratological point of view,
it seems clear that the ovular coats are usually, if not always, of foliar
nature, while the central nucleus is an axial organ; but if this be so there still
remains the question whether the leafy coats of the ovule are processes of
the carpel itself, or distinct independent formations, like the scales of a leaf-
bud; as to this latter point, the evidence is at present very conflicting. Prof.
Al. Braun, who has devoted much attention to the subject, describes and
figures ovules of Nigella and Adonis, wherein the outer coat of the ovule
was converted into a leafy, lobed mass, like the ordinary leaves, and these
he considers to be a portion, not of the carpel, but of the ovular bud; he,
however, hesitates to pronounce an opinion on the nature of the pedicel of
the ovule. In Primulaceæ, wherein ovular changes are very common, the
leafy coat of the ovule would seem, from the nature of the placenta, to be
independent of the carpel. Morren, who studied the changes in the ovules of
Primula sinensis, applied the term lepyrophylly (λεπυρον, a scale) to the
foliaceous condition of the testa in this plant. Unger[267] describes a series
of malformations in Primula sinensis, consisting chiefly of reversions of the
part of the flower to leaves. The carpels were entirely absent in this case,
and the place of the free central placenta was occupied by a circle of leaves,
sometimes bearing imperfect ovules on their edges. An instance of a similar
kind has been described by A. de Candolle.[268]
In these flowers the placenta seemed to be composed of several funiculi
soldered together, and bearing imperfect ovules. In other cases no traces of
ovules are visible, but the funiculi are in a foliaceous condition. Moquin
also alludes to a case of the same nature in Cortusa Mathioli, in which the
funiculi bore little rounded leaves. Brongniart has described some
malformations of Primula sinensis in which the ovules were transformed
of opinion with reference to the morphological nature of the ovule and its
component parts, much interest attaches to the malformations to which they
are occasionally subject. Considered purely in a teratological point of view,
it seems clear that the ovular coats are usually, if not always, of foliar
nature, while the central nucleus is an axial organ; but if this be so there still
remains the question whether the leafy coats of the ovule are processes of
the carpel itself, or distinct independent formations, like the scales of a leaf-
bud; as to this latter point, the evidence is at present very conflicting. Prof.
Al. Braun, who has devoted much attention to the subject, describes and
figures ovules of Nigella and Adonis, wherein the outer coat of the ovule
was converted into a leafy, lobed mass, like the ordinary leaves, and these
he considers to be a portion, not of the carpel, but of the ovular bud; he,
however, hesitates to pronounce an opinion on the nature of the pedicel of
the ovule. In Primulaceæ, wherein ovular changes are very common, the
leafy coat of the ovule would seem, from the nature of the placenta, to be
independent of the carpel. Morren, who studied the changes in the ovules of
Primula sinensis, applied the term lepyrophylly (λεπυρον, a scale) to the
foliaceous condition of the testa in this plant. Unger[267] describes a series
of malformations in Primula sinensis, consisting chiefly of reversions of the
part of the flower to leaves. The carpels were entirely absent in this case,
and the place of the free central placenta was occupied by a circle of leaves,
sometimes bearing imperfect ovules on their edges. An instance of a similar
kind has been described by A. de Candolle.[268]
In these flowers the placenta seemed to be composed of several funiculi
soldered together, and bearing imperfect ovules. In other cases no traces of
ovules are visible, but the funiculi are in a foliaceous condition. Moquin
also alludes to a case of the same nature in Cortusa Mathioli, in which the
funiculi bore little rounded leaves. Brongniart has described some
malformations of Primula sinensis in which the ovules were transformed
Page 337
wholly or partially into small leaves with three to five lobes.[269] Dr.
Marchand[270] mentions similar changes in Anagallis arvensis and Lonicera
Periclymenum.
Cramer[271] figures ovules of Primula sinensis in the form of stalked leaves,
often becoming infolded at the margins, and giving origin to a small
nucleus on their inner surface.
M. Tassi[272] records an instance in Symphytum officinale wherein the
ovules were replaced by two small linear leaves arising entirely from the
axis, and not from the carpels.
In most of the foregoing illustrations the foliar portion of the ovule must
have been independent of the carpel; this independence is less manifest,
though probably as real in the cases now to be mentioned. In Sinapis and in
Brassica oleracea foliaceous ovules may occasionally be seen, attached to
the placenta by long stalks. No trace of the nucleus is visible in these
specimens.
Marchand[270] mentions similar changes in Anagallis arvensis and Lonicera
Periclymenum.
Cramer[271] figures ovules of Primula sinensis in the form of stalked leaves,
often becoming infolded at the margins, and giving origin to a small
nucleus on their inner surface.
M. Tassi[272] records an instance in Symphytum officinale wherein the
ovules were replaced by two small linear leaves arising entirely from the
axis, and not from the carpels.
In most of the foregoing illustrations the foliar portion of the ovule must
have been independent of the carpel; this independence is less manifest,
though probably as real in the cases now to be mentioned. In Sinapis and in
Brassica oleracea foliaceous ovules may occasionally be seen, attached to
the placenta by long stalks. No trace of the nucleus is visible in these
specimens.
Page 338
Fig. 140.—Sinapis, replum and ovules; the dotted line shows the position of the carpels.
Griffith, in alluding to a similar case in Sinapis,[273] describes the ovules as
foliaceous, and having their backs turned away from the axis, the raphe
being next to the axis and representing the midrib the funicle corresponding
to the petiole. The outer tegument of the ovule, according to Griffith, is a
leaf united along its margins, but always more or less open at its apex. No
inversion can, therefore, really take place in anatropous ovules, but the
blade of the leaf is bent back on the funicle, with which its margins also
cohere.
Caspary, in an elaborate paper on phyllomorphy occurring in Trifolium
repens, figures foliaceous ovules springing from the edge of an open, leafy
carpel. The nucleus of the ovule, in these cases, appears to originate as a
little bud from the surface of the leafy ovule (figs. 141, 142).
Griffith, in alluding to a similar case in Sinapis,[273] describes the ovules as
foliaceous, and having their backs turned away from the axis, the raphe
being next to the axis and representing the midrib the funicle corresponding
to the petiole. The outer tegument of the ovule, according to Griffith, is a
leaf united along its margins, but always more or less open at its apex. No
inversion can, therefore, really take place in anatropous ovules, but the
blade of the leaf is bent back on the funicle, with which its margins also
cohere.
Caspary, in an elaborate paper on phyllomorphy occurring in Trifolium
repens, figures foliaceous ovules springing from the edge of an open, leafy
carpel. The nucleus of the ovule, in these cases, appears to originate as a
little bud from the surface of the leafy ovule (figs. 141, 142).
Page 339
Fig. 141.—Leafy ovules, &c., Trifolium repens.
In a species of Triumfetta (see p. 260), of which I examined dried
specimens, the ovary was open and partly foliaceous; it bore on its infolded
margins ten erect leaflets, representing so many ovules; each leaflet was
conduplicate, the back being turned towards the placenta.
Fig. 142.—Leafy ovules of Trifolium repens, showing formation of nucleus, &c. After Caspary.
In a species of Triumfetta (see p. 260), of which I examined dried
specimens, the ovary was open and partly foliaceous; it bore on its infolded
margins ten erect leaflets, representing so many ovules; each leaflet was
conduplicate, the back being turned towards the placenta.
Fig. 142.—Leafy ovules of Trifolium repens, showing formation of nucleus, &c. After Caspary.
Page 340
On the other hand, there are cases in which the leafy coat of the ovule, in
place of being a distinct organ, seems to originate from the margin of the
carpellary leaf itself—to be, as it were, a lobule or small process of the
carpel, and not an absolutely new growth. Thus, Planchon[274], from an
examination of some monstrous flowers of Drosera intermedia, was led to
the inference that the ovules are analogous to hairs on the margins of the
leaves. This acute botanist was enabled to trace all the gradations between
the simple cup formed by the confluence of four glanduliferous hairs and
the concave leaf and the perfect ovule.
Brongniart[275] records ovules of Delphinium elatum existing in the form of
marginal lobes of the carpellary leaf itself; so that each ovule corresponds to
a lobe or large tooth of this leaf, the funiculus, as well as the raphe, being
formed by the median nerve of the lateral lobe. M. Clos[276] mentions a
similar instance in Aquilegia Skinneri; and another is figured in Lindley's
'Elements of Botany,' p. 88, f. 180.
place of being a distinct organ, seems to originate from the margin of the
carpellary leaf itself—to be, as it were, a lobule or small process of the
carpel, and not an absolutely new growth. Thus, Planchon[274], from an
examination of some monstrous flowers of Drosera intermedia, was led to
the inference that the ovules are analogous to hairs on the margins of the
leaves. This acute botanist was enabled to trace all the gradations between
the simple cup formed by the confluence of four glanduliferous hairs and
the concave leaf and the perfect ovule.
Brongniart[275] records ovules of Delphinium elatum existing in the form of
marginal lobes of the carpellary leaf itself; so that each ovule corresponds to
a lobe or large tooth of this leaf, the funiculus, as well as the raphe, being
formed by the median nerve of the lateral lobe. M. Clos[276] mentions a
similar instance in Aquilegia Skinneri; and another is figured in Lindley's
'Elements of Botany,' p. 88, f. 180.
Page 341
Fig. 143.—Portion of an open foliaceous carpel of Delphinium, with ovules on the lobules.
Cramer[277], from an examination of several ovular malformations, as well
as from the investigation of the mode of evolution of the ovules, is led to a
similar conclusion with reference to the production of ovules from the
modified lobes of the carpellary leaf. Figs. 143–145, copied from Cramer,
show how the nucleus of the ovule is formed as a new growth from the
surface of the lobes of the leaf in Delphinium elatum.
Cramer[277], from an examination of several ovular malformations, as well
as from the investigation of the mode of evolution of the ovules, is led to a
similar conclusion with reference to the production of ovules from the
modified lobes of the carpellary leaf. Figs. 143–145, copied from Cramer,
show how the nucleus of the ovule is formed as a new growth from the
surface of the lobes of the leaf in Delphinium elatum.
Page 342
Fig. 144.—Section through marginal lobe of carpel (Delphinium), showing the nucleus (n).
Page 343
Fig. 145.—Section through marginal lobe of carpel, showing nucleus and tegument (Delphinium).
Page 344
Fig. 146.—1. Placenta of Dianthus, bearing ovules and carpels. 2. One of the ovaries separated.
Fig. 147.—Ovules of Dianthus passing into carpels.
One of the most singular instances of ovular malformation in record is that
cited by the Rev. M. J. Berkeley, in the 'Gardener's Chronicle,' September
28th, 1850, p. 612. The plant was a carnation, and its placenta bore, not
only ovules, but also carpels (fig. 146), the latter originating in a perverted
development of the former, so that many intermediate stages could be
traced between the ordinary ovule and the ovary (fig. 147, 1, a, 2, b). Some
of these carpels, thus derived from the ovules, themselves bore secondary
ovules on a marginal placenta, as shown in the sections at c, d, e. Could
such a change occur in the animal kingdom, there would be the unfertilised
ovum converted into an ovary, and this again bearing Graafian vesicles! In
Mr. Berkeley's carnation the change was not so great, seeing that the
nucleus of the ovule was not developed, and sufficient evidence has been
above given as to the foliar nature of the primine, while for a leaf to be
folded up so as to form a carpel is an ordinary occurrence.
It is worthy of remark that in these foliaceous ovules there is never more
than one coat, the secondine and other integuments do not make their
Fig. 147.—Ovules of Dianthus passing into carpels.
One of the most singular instances of ovular malformation in record is that
cited by the Rev. M. J. Berkeley, in the 'Gardener's Chronicle,' September
28th, 1850, p. 612. The plant was a carnation, and its placenta bore, not
only ovules, but also carpels (fig. 146), the latter originating in a perverted
development of the former, so that many intermediate stages could be
traced between the ordinary ovule and the ovary (fig. 147, 1, a, 2, b). Some
of these carpels, thus derived from the ovules, themselves bore secondary
ovules on a marginal placenta, as shown in the sections at c, d, e. Could
such a change occur in the animal kingdom, there would be the unfertilised
ovum converted into an ovary, and this again bearing Graafian vesicles! In
Mr. Berkeley's carnation the change was not so great, seeing that the
nucleus of the ovule was not developed, and sufficient evidence has been
above given as to the foliar nature of the primine, while for a leaf to be
folded up so as to form a carpel is an ordinary occurrence.
It is worthy of remark that in these foliaceous ovules there is never more
than one coat, the secondine and other integuments do not make their
Page 345
appearance in these cases, and that very generally the change in question
accompanies a similar foliaceous condition in the carpel, the margins of
which are more or less disunited.
Prof. A. Braun remarks that up to this date no such change has been
observed in the ovules of Monocotyledons.
Changes in the nucleus of the ovule.—The preceding remarks have had
reference especially to the ovular coats, but it is desirable also to allude to
certain points connected with the nucleus. Very frequently, when the coat of
the ovule is phylloid, as before described, the nucleus is altogether wanting,
though sometimes it is present as a small cellular papilla; very rarely is it to
be found in its perfect state. Occasionally the nucleus is present in the guise
of a small elongated branch. Wigand cites ovular buds in every stage of
progress into a branch, sometimes even bearing indications of anthers.
Wydler has observed a similar occurrence in ovules of Alliaria officinalis,
and Schimper has described and figured specimens of Nigella damascena in
which the outer coats of the ovule were but little changed, while the nucleus
was replaced by a leafy shoot. On one of the leaves of this latter was found
an imperfect ovule—an ovule on an ovule!
Fig. 148 shows a floret of a species of Gaillardia, in which the ovule was
replaced by a leafy shoot which had made its way through a chink in the
ovary. In this specimen, however, there was no evidence to show whether
the shoot in question was a perverted development of the nucleus, or
whether it was wholly independent of the ovule.
accompanies a similar foliaceous condition in the carpel, the margins of
which are more or less disunited.
Prof. A. Braun remarks that up to this date no such change has been
observed in the ovules of Monocotyledons.
Changes in the nucleus of the ovule.—The preceding remarks have had
reference especially to the ovular coats, but it is desirable also to allude to
certain points connected with the nucleus. Very frequently, when the coat of
the ovule is phylloid, as before described, the nucleus is altogether wanting,
though sometimes it is present as a small cellular papilla; very rarely is it to
be found in its perfect state. Occasionally the nucleus is present in the guise
of a small elongated branch. Wigand cites ovular buds in every stage of
progress into a branch, sometimes even bearing indications of anthers.
Wydler has observed a similar occurrence in ovules of Alliaria officinalis,
and Schimper has described and figured specimens of Nigella damascena in
which the outer coats of the ovule were but little changed, while the nucleus
was replaced by a leafy shoot. On one of the leaves of this latter was found
an imperfect ovule—an ovule on an ovule!
Fig. 148 shows a floret of a species of Gaillardia, in which the ovule was
replaced by a leafy shoot which had made its way through a chink in the
ovary. In this specimen, however, there was no evidence to show whether
the shoot in question was a perverted development of the nucleus, or
whether it was wholly independent of the ovule.
Page 346
Fig. 148.—Floret of Gaillardia, showing leafy shoot occupying the place of the ovule.
From this occasional elongation of the nucleus, as well as from the foliar
nature of the ovular coats, Prof. Alex. Braun arrives at the conclusion that
the ovule is to be looked on as a bud, the ovular coatings, so often variable
in number, representing the scales of the bud, the nucleus corresponding to
the end of the axis or growing point. Griffith had previously expressed the
same opinion from his observations on malformed ovules of Sinapis and
Lonicera, while Caspary's conclusions from the foliaceous ovules of
Trifolum repens are somewhat similar. The latter observer considers that the
funiculus, with the integuments, is the equivalent of a leaflet, the petiolule
or midrib of which answers to the funiculus, and its hollow expansion to the
integument. The nucleus itself is considered to be a new formation
analogous to a shoot.
From this occasional elongation of the nucleus, as well as from the foliar
nature of the ovular coats, Prof. Alex. Braun arrives at the conclusion that
the ovule is to be looked on as a bud, the ovular coatings, so often variable
in number, representing the scales of the bud, the nucleus corresponding to
the end of the axis or growing point. Griffith had previously expressed the
same opinion from his observations on malformed ovules of Sinapis and
Lonicera, while Caspary's conclusions from the foliaceous ovules of
Trifolum repens are somewhat similar. The latter observer considers that the
funiculus, with the integuments, is the equivalent of a leaflet, the petiolule
or midrib of which answers to the funiculus, and its hollow expansion to the
integument. The nucleus itself is considered to be a new formation
analogous to a shoot.
Page 347
M. van Tieghem's conclusion[278] from the examination, of flowers of
Tropæolum majus, in which the ovules were replaced by perfect peltate
leaves, is that the ovules are foliar productions springing, not directly from
a prolonged floral axis, as in Primulaceæ, but from branches of the axis
arising from the axils of the carpellary leaves.
Phyllody of the ovules has been met with most often in the following
species:
*Aquilegia vulgaris!
Skinneri.
Delphinium crassicaule.
elatum.
dictyocarpum.
Ajacis.
Nigella damascena.
Adonis autumnalis.
Cheiranthus Cheiri!
Nasturtium, sp.
Sisymbrium officinale!
Brassica napus!
*olcracea!
*Alliaria officinalis!
Sinapis arvensis!
Turritis, sp.
Thlaspi arvense.
Erucastrum Pollichii.
Stellaria media.
*Reseda lutea.
Drosera intermedia.
Agrostemma Githago.
Stellaria media.
Triumfetta, sp.!
Tropæolum majus!
Dictamnus albus.
Fraxinella!
Caram carui
Tropæolum majus, in which the ovules were replaced by perfect peltate
leaves, is that the ovules are foliar productions springing, not directly from
a prolonged floral axis, as in Primulaceæ, but from branches of the axis
arising from the axils of the carpellary leaves.
Phyllody of the ovules has been met with most often in the following
species:
*Aquilegia vulgaris!
Skinneri.
Delphinium crassicaule.
elatum.
dictyocarpum.
Ajacis.
Nigella damascena.
Adonis autumnalis.
Cheiranthus Cheiri!
Nasturtium, sp.
Sisymbrium officinale!
Brassica napus!
*olcracea!
*Alliaria officinalis!
Sinapis arvensis!
Turritis, sp.
Thlaspi arvense.
Erucastrum Pollichii.
Stellaria media.
*Reseda lutea.
Drosera intermedia.
Agrostemma Githago.
Stellaria media.
Triumfetta, sp.!
Tropæolum majus!
Dictamnus albus.
Fraxinella!
Caram carui
Page 348
Pastinaca sativa.
Torilis anthriscus.
Thysselinum palustre.
Epilobium palustre.
Rosa, sp.
Fragaria alpina.
*Trifolium repens!
Medicago maculata.
Desmodium canadense.
Melilotus macrorhiza.
Lonicera, sp.
Gaillardia!
Crepis, sp.
Phyteuma odorata.
Symphytum Zeyheri.
*officinale.
Stachys sylvatica.
Anagallia arvensis.
phœnicea.
Lysimachia ephemerum.
*Primula sinensis!
Auricula.
prænitens.
Gilia glomeruliflora.
Rumex arifolius.
scutatus.
Salix capræa.
The following list of publications relating to ovular malformations is copied
from A. Braun, 'Ueber Polyembryonie und Keimung von Cælobogyne'
(Appendix),[279] to which are also added some others not alluded to by that
author and not specially referred to in the preceding pages:
Jaeger, 'Missbilld. d. Gewächse,' p. 78, 79, f. 47. Rœper, 'Enum. Euphorb.,' 1824. p. 45,
Delphinium.—Schimper, 'Flora,' 1829, pp. 437–8, et 'Mag. fur Pharmacie de Geiger,'
1829–30, pl. iv-vi, text wanting, Primula, Reseda, Cheiranthus.—Engelmann, 'De
Antholysi,' 1832.—Valentin, 'Act. Acad. Nat. Cur.,' 1839, p. 225, Lysimachia.—Unger,
'Act. Acad. Nat. Cur.,' xxii, 11, 1850, p. 543, t. 5 B, Primula.—'Flora (B. Z.)', 1842, p. 369,
t. ii, Trifolium.—Brongniart, 'Ann. Sc. Nat.,' 1834, ii, p. 308; also 'Archives Mus. d'Hist.
Torilis anthriscus.
Thysselinum palustre.
Epilobium palustre.
Rosa, sp.
Fragaria alpina.
*Trifolium repens!
Medicago maculata.
Desmodium canadense.
Melilotus macrorhiza.
Lonicera, sp.
Gaillardia!
Crepis, sp.
Phyteuma odorata.
Symphytum Zeyheri.
*officinale.
Stachys sylvatica.
Anagallia arvensis.
phœnicea.
Lysimachia ephemerum.
*Primula sinensis!
Auricula.
prænitens.
Gilia glomeruliflora.
Rumex arifolius.
scutatus.
Salix capræa.
The following list of publications relating to ovular malformations is copied
from A. Braun, 'Ueber Polyembryonie und Keimung von Cælobogyne'
(Appendix),[279] to which are also added some others not alluded to by that
author and not specially referred to in the preceding pages:
Jaeger, 'Missbilld. d. Gewächse,' p. 78, 79, f. 47. Rœper, 'Enum. Euphorb.,' 1824. p. 45,
Delphinium.—Schimper, 'Flora,' 1829, pp. 437–8, et 'Mag. fur Pharmacie de Geiger,'
1829–30, pl. iv-vi, text wanting, Primula, Reseda, Cheiranthus.—Engelmann, 'De
Antholysi,' 1832.—Valentin, 'Act. Acad. Nat. Cur.,' 1839, p. 225, Lysimachia.—Unger,
'Act. Acad. Nat. Cur.,' xxii, 11, 1850, p. 543, t. 5 B, Primula.—'Flora (B. Z.)', 1842, p. 369,
t. ii, Trifolium.—Brongniart, 'Ann. Sc. Nat.,' 1834, ii, p. 308; also 'Archives Mus. d'Hist.
Page 349
Nat.,' 1844, t. iv, p. 43, pl. iv, v, Primula.—Reissek, 'Linnæa,' xvii, 1843, Alliaria.—
Wydler, 'Denkshrift. d. Regensb. Bot. Gesell.,' 1855, iv, s. 77, t. vii, Alliaria.—Wigand.
'Grundlegung der Pflanzen Teratol.,' 1850, p. 39, Turritis.—Wigand, 'Bot.
Untersuchungen,' 1853, p. 23, Rosa, Turritis, Crepis.—Germain de St. Pierre, 'L'lnstitut,'
1853, n. 1051, p. 351.—Rossmann, "Entwicklung der Eiknospen aus dem Fruchtblatte,"
&c., 'Flora,' 1855, pp. 647 and 705.—Dareste, 'Ann. Sc. Nat.,' 1842, p. 220, Delphinium.—
Fresenius, 'Mus. Senkenb.,' ii, p. 39, t. iv, f. 9, Primula.—Schultz, 'Flora o. d. Bot. Zeit.,'
1834, xvii, p. 121, Nasturtium.—Seringe and Heyland, 'Bull. Bot.,' 1–7, Diplotaxis.—Clos,
'Mem. Acad. Toulouse,' vi, 1862, Delphinium.—Morren, C., 'Bull. Acad. Belg.,' xix, part ii,
p. 519, Primula.—Caspary, 'Schrift. d. Physik. Œk. Gesell. zu Königsberg,' band ii, p. 51,
tabs. ii, iii. Fleischer, 'Ueber Missbildungen Verschiedener Cultur Pflanzen.,' &c.,
Esslingen, 1862. Cramer, 'Bildungsabweich,' p. 68, &c. &c., Trifolium.—Moquin-Tandon,
'El. Terat. Veg.,' p. 206, Cortusa.—Guillard, 'Bull. Soc. Bot. Fr.,' 1857, vol. iv, p. 761,
Stellaria.—Moelkenboer, 'Tijdschrift v. Natuurl. Geschied.,' 1843, p. 355, t. vi, vii,
Primula.—Van Tieghem, 'Bull. Soc. Bot. Fr.,' 1865, p, 411, Tropæolum.
Phyllody in accessory organs.—In addition to the ordinary organs of the
plant, what are termed the accessory organs, such as hairs, spines, &c.,
sometimes become foliaceous. It is not to be wondered at that spines, when
they represent the framework of a leaf, become sometimes clothed with
cellular tissue, and thus become indeed true leaves. This happens
occasionally in Berberis; a similar thing occurs in the stipules of some
Leguminosæ; the scales of some begonias; the tendrils of Bignonia, Cobæa,
&c.
The presence of two small green laminæ on the outer side of the two
posterior stamens in Antirrhinum majus has also been met with. The
adventitious organs appeared as if they were developments from the
thalamus—a kind of foliaceous disc, in fact.
Wydler, 'Denkshrift. d. Regensb. Bot. Gesell.,' 1855, iv, s. 77, t. vii, Alliaria.—Wigand.
'Grundlegung der Pflanzen Teratol.,' 1850, p. 39, Turritis.—Wigand, 'Bot.
Untersuchungen,' 1853, p. 23, Rosa, Turritis, Crepis.—Germain de St. Pierre, 'L'lnstitut,'
1853, n. 1051, p. 351.—Rossmann, "Entwicklung der Eiknospen aus dem Fruchtblatte,"
&c., 'Flora,' 1855, pp. 647 and 705.—Dareste, 'Ann. Sc. Nat.,' 1842, p. 220, Delphinium.—
Fresenius, 'Mus. Senkenb.,' ii, p. 39, t. iv, f. 9, Primula.—Schultz, 'Flora o. d. Bot. Zeit.,'
1834, xvii, p. 121, Nasturtium.—Seringe and Heyland, 'Bull. Bot.,' 1–7, Diplotaxis.—Clos,
'Mem. Acad. Toulouse,' vi, 1862, Delphinium.—Morren, C., 'Bull. Acad. Belg.,' xix, part ii,
p. 519, Primula.—Caspary, 'Schrift. d. Physik. Œk. Gesell. zu Königsberg,' band ii, p. 51,
tabs. ii, iii. Fleischer, 'Ueber Missbildungen Verschiedener Cultur Pflanzen.,' &c.,
Esslingen, 1862. Cramer, 'Bildungsabweich,' p. 68, &c. &c., Trifolium.—Moquin-Tandon,
'El. Terat. Veg.,' p. 206, Cortusa.—Guillard, 'Bull. Soc. Bot. Fr.,' 1857, vol. iv, p. 761,
Stellaria.—Moelkenboer, 'Tijdschrift v. Natuurl. Geschied.,' 1843, p. 355, t. vi, vii,
Primula.—Van Tieghem, 'Bull. Soc. Bot. Fr.,' 1865, p, 411, Tropæolum.
Phyllody in accessory organs.—In addition to the ordinary organs of the
plant, what are termed the accessory organs, such as hairs, spines, &c.,
sometimes become foliaceous. It is not to be wondered at that spines, when
they represent the framework of a leaf, become sometimes clothed with
cellular tissue, and thus become indeed true leaves. This happens
occasionally in Berberis; a similar thing occurs in the stipules of some
Leguminosæ; the scales of some begonias; the tendrils of Bignonia, Cobæa,
&c.
The presence of two small green laminæ on the outer side of the two
posterior stamens in Antirrhinum majus has also been met with. The
adventitious organs appeared as if they were developments from the
thalamus—a kind of foliaceous disc, in fact.
Page 350
Fig. 149.—Leafy petal of Epilobium.
Fig. 150.—Chloranthy, &c. Epilobium hirsutum.
Fig. 150.—Chloranthy, &c. Epilobium hirsutum.
Page 351
Chloranthy.—The term phyllomorphy is applied to the individual parts of
the flower which assume the form and appearance of leaves. By chloranthy
it is to be understood that all, or the great majority of the organs of the
flower assume these conditions.[280] In chloranthy, as here defined, there is
no unusual number of buds, as there is in prolification, but the appearance
of the flower-bud is so changed as to make it resemble more closely a leaf-
bud than a flower-bud. There is not necessarily any increase in the number,
or any alteration in the position of the buds, but the form and appearance of
the latter differ from what is usual. Chloranthy, then, is a more complete
form of frondescence. Owing to the vagueness with which the word has
been applied by various authors, it becomes very difficult to ascertain
whether the recorded instances of chloranthy were really illustrations of
what is here meant by that term, or whether they were cases of mere
virescence (green colour, without other perceptible change), or of
prolification (formation of adventitious buds). It is, therefore, quite possible
that some of the instances to be now mentioned were not strictly cases of
chloranthy.
the flower which assume the form and appearance of leaves. By chloranthy
it is to be understood that all, or the great majority of the organs of the
flower assume these conditions.[280] In chloranthy, as here defined, there is
no unusual number of buds, as there is in prolification, but the appearance
of the flower-bud is so changed as to make it resemble more closely a leaf-
bud than a flower-bud. There is not necessarily any increase in the number,
or any alteration in the position of the buds, but the form and appearance of
the latter differ from what is usual. Chloranthy, then, is a more complete
form of frondescence. Owing to the vagueness with which the word has
been applied by various authors, it becomes very difficult to ascertain
whether the recorded instances of chloranthy were really illustrations of
what is here meant by that term, or whether they were cases of mere
virescence (green colour, without other perceptible change), or of
prolification (formation of adventitious buds). It is, therefore, quite possible
that some of the instances to be now mentioned were not strictly cases of
chloranthy.
Page 352
Fig. 151.—a. Open leafy carpel of "green rose," with two deformed ovules. b. Ovule separate. c.
Primine removed. d. Secondine and nucleus, with the bulbous end that projects through the
micropyle.
Seringe[281] has described a malformation in Diplotaxis tenuifolia in which
all the floral organs were replaced by sixteen distinct leaflets which had
preserved their proper relative position. The Cruciferæ, of which family the
last-named plant is a member, are particularly liable to this malformation, as
also are the Rosaceæ, as will be seen from the following illustrations. Roses
indeed often exhibit alterations of this kind as the commencement of
prolification. There is also in cultivation a rose[282] called the green rose,
"Rose bengale à fleurs vertes," in which all the parts of the flower are
represented by leaves. One of the most remarkable features in this plant is,
that the carpels have often two ovules on their margins. Now, Payer, in his
"Organogénie," has shown that at a certain period of the development of the
ordinary rose flower the ovary contains two collateral ovules, of which one
Primine removed. d. Secondine and nucleus, with the bulbous end that projects through the
micropyle.
Seringe[281] has described a malformation in Diplotaxis tenuifolia in which
all the floral organs were replaced by sixteen distinct leaflets which had
preserved their proper relative position. The Cruciferæ, of which family the
last-named plant is a member, are particularly liable to this malformation, as
also are the Rosaceæ, as will be seen from the following illustrations. Roses
indeed often exhibit alterations of this kind as the commencement of
prolification. There is also in cultivation a rose[282] called the green rose,
"Rose bengale à fleurs vertes," in which all the parts of the flower are
represented by leaves. One of the most remarkable features in this plant is,
that the carpels have often two ovules on their margins. Now, Payer, in his
"Organogénie," has shown that at a certain period of the development of the
ordinary rose flower the ovary contains two collateral ovules, of which one
Page 353
becomes in process of time suppressed.[283] Geum coccineum has been
found by Wigand with its flowers in this condition.[284]
Lindley[285] figures a very interesting illustration in Potentilla nepalensis, in
which some of the flowers have their component parts leafy, in others the
receptacle lengthens, till in extreme cases the whole of the floral apparatus
is represented by a branch bearing a rosette of leaves.
A particular variety of the Alpine strawberry is also described as
occasionally subject to this transformation. In these flowers the calyx
remains normal, while all the other parts of the flower, even to the coating
of the ovule, assume a leaf-like condition.[286]
Among Leguminosæ a partial leafy condition (frondescence), or a more
complete degree of the same change, (chloranthy) is not infrequent,
particularly in Trifolium repens. In this species the changes are so common,
so various and important, that they may be alluded to in some little detail.
M. Germain de Saint Pierre,[287] in commenting on the frequency with
which the flowers of this plant are more or less frondescent, remarks that
although all the flowers on one plant may be affected, they are all changed
in the same manner, but on different specimens different degrees of
transformation are found. In all the corolla and stamens are comparatively
little removed from the ordinary form, the calyx and pistil, however, have a
particular tendency to assume a foliar condition. The author just cited
arranges the malformations of this plant under three heads, as follows:
1. Calyx-teeth larger than usual, sometimes dentate at the margin; petals more or less
regular and disposed to run away from the papilionaceous form; filaments free; anthers
normal; carpel transformed into a true leaf with a long stalk provided at the base, with two
stipules, terminal leaflet, solitary, green, with no trace of ovules. Sometimes a second
carpellary leaf, similar to the first, is formed; in other cases the central axis of the flower is
occasionally prolonged into a head of young flowers—median prolification. In some few
instances the calyx is not at all altered, but the carpellary leaf is trifoliolate, or even
found by Wigand with its flowers in this condition.[284]
Lindley[285] figures a very interesting illustration in Potentilla nepalensis, in
which some of the flowers have their component parts leafy, in others the
receptacle lengthens, till in extreme cases the whole of the floral apparatus
is represented by a branch bearing a rosette of leaves.
A particular variety of the Alpine strawberry is also described as
occasionally subject to this transformation. In these flowers the calyx
remains normal, while all the other parts of the flower, even to the coating
of the ovule, assume a leaf-like condition.[286]
Among Leguminosæ a partial leafy condition (frondescence), or a more
complete degree of the same change, (chloranthy) is not infrequent,
particularly in Trifolium repens. In this species the changes are so common,
so various and important, that they may be alluded to in some little detail.
M. Germain de Saint Pierre,[287] in commenting on the frequency with
which the flowers of this plant are more or less frondescent, remarks that
although all the flowers on one plant may be affected, they are all changed
in the same manner, but on different specimens different degrees of
transformation are found. In all the corolla and stamens are comparatively
little removed from the ordinary form, the calyx and pistil, however, have a
particular tendency to assume a foliar condition. The author just cited
arranges the malformations of this plant under three heads, as follows:
1. Calyx-teeth larger than usual, sometimes dentate at the margin; petals more or less
regular and disposed to run away from the papilionaceous form; filaments free; anthers
normal; carpel transformed into a true leaf with a long stalk provided at the base, with two
stipules, terminal leaflet, solitary, green, with no trace of ovules. Sometimes a second
carpellary leaf, similar to the first, is formed; in other cases the central axis of the flower is
occasionally prolonged into a head of young flowers—median prolification. In some few
instances the calyx is not at all altered, but the carpellary leaf is trifoliolate, or even
Page 354
quinquefoliolate, the corolla being then absent. The heads of flowers in this first form have
the aspect of little tufts of leaves.
2. Each of the teeth of the calyx is represented by a long stalk, terminated by a single
articulated leaflet, the bi-labiate form of the calyx is still recognisable; the two upper petals
are united, the three lower separate; the tube of the calyx is not deformed and seems to be
formed of the petioles of the sepals united by their stipules. In this second class of cases the
corolla is papilionaceous, the filaments free, the carpellary leaf on a long stalk provided
with stipules, its blade more or less like the usual carpel, with its margins disunited or more
commonly united with the ovules in the interior, sometimes represented by a foliaceous,
dentate primine only. In one case the carpel was closed above, gaping below, where it gave
origin to several leaflets, the lower ones oval, dentate, like ordinary leaflets, the upper ones
merely lanceolate, leafy lobes, representing the primine reduced to a foliaceous condition.
Inflorescence—a head with leafy flowers on long stalks, which are longer at the
circumference than in the centre.
3. Calyx-teeth lance-shaped, acuminate; corolla more or less regular, arrested in its
development and scarcely exceeding the tube of the calyx within which it is crumpled up;
stamens but little changed; carpellary leaf on a short stalk, not exceeding the calyx tube,
but the ovarian portion very long, and provided with abortive ovules.
These three groups will be found to include most of the forms under which frondescence of
the clover blossoms occurs, but there are, of course, intermediate forms not readily to be
grouped under either of the above heads. Such are the cases brought under the notice of the
British Association at Birmingham in 1849 by Mr. R. Austen, in some of which the petals
and stamens even were represented by leaves.
Although, on the whole, chloranthy is most frequent in the families already
alluded to, yet it is by no means confined to them, as the examples now to
be given amply show. Specimens of Nymphæa Lotus have been seen in
which all the parts of the flower, even to the stigmas, were leafy, while the
ovules were entirely wanting.
Planchon[288] figures and describes a flower of Drosera intermedia that had
passed into a chloranthic condition, excepting the calyx, which was
unchanged; the petals, like the valves of the ovary, were provided with
stipules, and were circinate in vernation.
the aspect of little tufts of leaves.
2. Each of the teeth of the calyx is represented by a long stalk, terminated by a single
articulated leaflet, the bi-labiate form of the calyx is still recognisable; the two upper petals
are united, the three lower separate; the tube of the calyx is not deformed and seems to be
formed of the petioles of the sepals united by their stipules. In this second class of cases the
corolla is papilionaceous, the filaments free, the carpellary leaf on a long stalk provided
with stipules, its blade more or less like the usual carpel, with its margins disunited or more
commonly united with the ovules in the interior, sometimes represented by a foliaceous,
dentate primine only. In one case the carpel was closed above, gaping below, where it gave
origin to several leaflets, the lower ones oval, dentate, like ordinary leaflets, the upper ones
merely lanceolate, leafy lobes, representing the primine reduced to a foliaceous condition.
Inflorescence—a head with leafy flowers on long stalks, which are longer at the
circumference than in the centre.
3. Calyx-teeth lance-shaped, acuminate; corolla more or less regular, arrested in its
development and scarcely exceeding the tube of the calyx within which it is crumpled up;
stamens but little changed; carpellary leaf on a short stalk, not exceeding the calyx tube,
but the ovarian portion very long, and provided with abortive ovules.
These three groups will be found to include most of the forms under which frondescence of
the clover blossoms occurs, but there are, of course, intermediate forms not readily to be
grouped under either of the above heads. Such are the cases brought under the notice of the
British Association at Birmingham in 1849 by Mr. R. Austen, in some of which the petals
and stamens even were represented by leaves.
Although, on the whole, chloranthy is most frequent in the families already
alluded to, yet it is by no means confined to them, as the examples now to
be given amply show. Specimens of Nymphæa Lotus have been seen in
which all the parts of the flower, even to the stigmas, were leafy, while the
ovules were entirely wanting.
Planchon[288] figures and describes a flower of Drosera intermedia that had
passed into a chloranthic condition, excepting the calyx, which was
unchanged; the petals, like the valves of the ovary, were provided with
stipules, and were circinate in vernation.
Page 355
M. A. Viaud-Grand-Marais[289] records an interesting example of
chloranthy, in which the sepals, petals, pistils, and ovules of Anagallis
arvensis were all foliaceous. Similar changes have not unfrequently been
met with in Dictamnus Fraxinella.
M. Germain de Saint Pierre has also recorded the following deviations in
the flowers of Rumex arifolius and R. scutatus; in these specimens the calyx
was normal, the petals large, foliaceous, shaped like the stem-leaves, the
stamens were absent, the three carpels fused into a triangular leafy pod, as
long again as the perianth, the stigmas normal or wanting, the ovule
represented by a thick funicle, terminated by a foliaceous appendage
analogous to the primine.[290]
In grasses it frequently happens that the flowers are replaced by leaf-buds;
this condition is alluded to elsewhere under the head of viviparous grasses,
but in this place may be mentioned a less degree of change, and which
seems to have been a genuine case of chloranthy in Glyceria fluitans, the
spikelet of which, as observed by Wigand,[291] consisted below of the
ordinary unchanged glumes, but the remaining paleæ as well as the lodicles
and stamens were represented by ligulate leaves. The plant, it is stated, was
affected by a parasitic fungus. On the other hand, General Munro, in his
valuable monograph of the Bambusaceæ,[292] refers to an illustration in
which "the lowest glumes generally, and the lowest paleæ occasionally, had
the appearance of miniature leaves, with vaginæ, ligules and cilia,
enveloping, however, perfect fertile spiculæ; as progress is made towards
the top of the spike, the ligule first, then the cilia, and finally, the leaf-like
extension disappears, and the uppermost glumes assume the ordinary shape
and form of those organs."
General remarks on chloranthy and frondescence.—Moquin remarks
with justice that the position of the flowers on the axis is of importance with
chloranthy, in which the sepals, petals, pistils, and ovules of Anagallis
arvensis were all foliaceous. Similar changes have not unfrequently been
met with in Dictamnus Fraxinella.
M. Germain de Saint Pierre has also recorded the following deviations in
the flowers of Rumex arifolius and R. scutatus; in these specimens the calyx
was normal, the petals large, foliaceous, shaped like the stem-leaves, the
stamens were absent, the three carpels fused into a triangular leafy pod, as
long again as the perianth, the stigmas normal or wanting, the ovule
represented by a thick funicle, terminated by a foliaceous appendage
analogous to the primine.[290]
In grasses it frequently happens that the flowers are replaced by leaf-buds;
this condition is alluded to elsewhere under the head of viviparous grasses,
but in this place may be mentioned a less degree of change, and which
seems to have been a genuine case of chloranthy in Glyceria fluitans, the
spikelet of which, as observed by Wigand,[291] consisted below of the
ordinary unchanged glumes, but the remaining paleæ as well as the lodicles
and stamens were represented by ligulate leaves. The plant, it is stated, was
affected by a parasitic fungus. On the other hand, General Munro, in his
valuable monograph of the Bambusaceæ,[292] refers to an illustration in
which "the lowest glumes generally, and the lowest paleæ occasionally, had
the appearance of miniature leaves, with vaginæ, ligules and cilia,
enveloping, however, perfect fertile spiculæ; as progress is made towards
the top of the spike, the ligule first, then the cilia, and finally, the leaf-like
extension disappears, and the uppermost glumes assume the ordinary shape
and form of those organs."
General remarks on chloranthy and frondescence.—Moquin remarks
with justice that the position of the flowers on the axis is of importance with
Page 356
reference to the existence of chloranthy. Terminal flowers are more subject
to it than lateral ones, and if the latter, by accident, become terminal, they
seem peculiarly liable to assume a foliaceous condition. Kirschleger says,
that in Rubus there are two sorts of chloranthy, according as the anomaly
affects the ordinary flowering branches, or the leafy shoots of the year, the
summits of which, instead of developing in the customary manner,
terminate each in one vast and long inflorescence, very loose and
indeterminate, and with axillary flowers.[293]
On the whole, taking in consideration cases of partial frondescence, as well
as those in which most of the parts of the flower are affected, phyllody
would seem to be most common in the petals and carpels, least so in the
case of the stamens and sepals. It is more common among polysepalous and
polypetalous plants than in those in which the sepals or petals are united
together.
The causes assigned for these phenomena are chiefly those of a nature to
debilitate or injure the plant; thus it has been frequently observed to follow
the puncture of an insect. M. Guillard[294] gives an instance in Stellaria
media where the condition appeared to be due to the attacks of an insect
Thrips fasciata. Still more commonly it arises from the attacks of parasitic
fungi, e.g. Uredo candida, in Crucifers, &c.
In other cases it has been observed when the plants have been growing in
very damp places, or in very wet seasons, or in the shade, or where the plant
has been much trampled on. This happens frequently with Trifolium repens.
The frequency with which the change is encountered in this particular
species is very remarkable; it is difficult to see why one species should be
so much more subject to the kind of change than another of nearly identical
conformation.
to it than lateral ones, and if the latter, by accident, become terminal, they
seem peculiarly liable to assume a foliaceous condition. Kirschleger says,
that in Rubus there are two sorts of chloranthy, according as the anomaly
affects the ordinary flowering branches, or the leafy shoots of the year, the
summits of which, instead of developing in the customary manner,
terminate each in one vast and long inflorescence, very loose and
indeterminate, and with axillary flowers.[293]
On the whole, taking in consideration cases of partial frondescence, as well
as those in which most of the parts of the flower are affected, phyllody
would seem to be most common in the petals and carpels, least so in the
case of the stamens and sepals. It is more common among polysepalous and
polypetalous plants than in those in which the sepals or petals are united
together.
The causes assigned for these phenomena are chiefly those of a nature to
debilitate or injure the plant; thus it has been frequently observed to follow
the puncture of an insect. M. Guillard[294] gives an instance in Stellaria
media where the condition appeared to be due to the attacks of an insect
Thrips fasciata. Still more commonly it arises from the attacks of parasitic
fungi, e.g. Uredo candida, in Crucifers, &c.
In other cases it has been observed when the plants have been growing in
very damp places, or in very wet seasons, or in the shade, or where the plant
has been much trampled on. This happens frequently with Trifolium repens.
The frequency with which the change is encountered in this particular
species is very remarkable; it is difficult to see why one species should be
so much more subject to the kind of change than another of nearly identical
conformation.
Page 357
It might at first be supposed that the same causes that bring about the
complete substitution of leaf-buds for flower-buds (see Heterotaxy) would
operate also in the partial substitution of leaves for other parts of the flower,
but it will be seen that the inducing cause, whether similar or not in the two
cases respectively, acts at different times; in the one case, it is not brought
into play until the rudiments of the flower are already formed, whereas in
the other the influence is exerted prior to the formation of the flower. So
that while the formation of leaf-buds in place of flower-buds may be and
generally is due to an excess of nutrition, inducing over activity of the
vegetative organs, the production of phyllomorphic or chloranthic flowers
may be owing rather to a perversion of development arising from injury or
from some debilitating agency. The discrepancies in the assigned causes for
the conditions above mentioned may, therefore, in great measure, be
attributed to the different periods at which the causes in question operate.
The following list may serve as a guide to the plants most frequently the
subjects of chloranthy, but reference should also be made to preceding and
subsequent sections, and to that relating to prolification of the
inflorescence.
Aquilegia vulgaris.
Chelidonium majus.
Corydalis aurea.
Nymphæa Lotus!
*Brassica oleracea!
Bunias.
Hesperis matronalis.
*Sinapis arvensis!
Sisymbrium officinale.
Erucastrum canariense.
Diplotaxis tenuifolia.
Lychnis dioica!
Cerastium glomeratum!
triviale.
complete substitution of leaf-buds for flower-buds (see Heterotaxy) would
operate also in the partial substitution of leaves for other parts of the flower,
but it will be seen that the inducing cause, whether similar or not in the two
cases respectively, acts at different times; in the one case, it is not brought
into play until the rudiments of the flower are already formed, whereas in
the other the influence is exerted prior to the formation of the flower. So
that while the formation of leaf-buds in place of flower-buds may be and
generally is due to an excess of nutrition, inducing over activity of the
vegetative organs, the production of phyllomorphic or chloranthic flowers
may be owing rather to a perversion of development arising from injury or
from some debilitating agency. The discrepancies in the assigned causes for
the conditions above mentioned may, therefore, in great measure, be
attributed to the different periods at which the causes in question operate.
The following list may serve as a guide to the plants most frequently the
subjects of chloranthy, but reference should also be made to preceding and
subsequent sections, and to that relating to prolification of the
inflorescence.
Aquilegia vulgaris.
Chelidonium majus.
Corydalis aurea.
Nymphæa Lotus!
*Brassica oleracea!
Bunias.
Hesperis matronalis.
*Sinapis arvensis!
Sisymbrium officinale.
Erucastrum canariense.
Diplotaxis tenuifolia.
Lychnis dioica!
Cerastium glomeratum!
triviale.
Page 358
Stellaria media.
Poterium polygamum.
Torilis anthriscus.
Seseli, sp.
Selinum caruifolium.
Epilobium hirsutum!
Begonia fuchsioides.
Gomphia, sp.
Scabiosa Columbaria.
Dipsacus fullonum.
Matricaria Parthenium.
Calendula officinalis.
Campanula pyramidalis.
Reseda odorata!
Vitis vinifera.
Dictamnus Fraxinella!
Triumfetta, sp.!
*Tropæolum majus!
Rhamnus Frangula.
*Trifolium repens!
Lupinus, sp.
Rosa diversifolia!
Potentilla nepalensis.
argentea.
Fragaria vesca!
Geum rivale.
Rubus fruticosus.
cæsius.
Saxifraga foliosa.
Verbascum phlomoides.
Scrophularia nodosa.
aquatica!
*Primula sinensis!
Lysimachia Ephemerum.
Anagallis arvensis.
Webbiana.
Nicotiana rustica.
Anchusa ochroleuca.
Myosotis cæspitosa.
Poterium polygamum.
Torilis anthriscus.
Seseli, sp.
Selinum caruifolium.
Epilobium hirsutum!
Begonia fuchsioides.
Gomphia, sp.
Scabiosa Columbaria.
Dipsacus fullonum.
Matricaria Parthenium.
Calendula officinalis.
Campanula pyramidalis.
Reseda odorata!
Vitis vinifera.
Dictamnus Fraxinella!
Triumfetta, sp.!
*Tropæolum majus!
Rhamnus Frangula.
*Trifolium repens!
Lupinus, sp.
Rosa diversifolia!
Potentilla nepalensis.
argentea.
Fragaria vesca!
Geum rivale.
Rubus fruticosus.
cæsius.
Saxifraga foliosa.
Verbascum phlomoides.
Scrophularia nodosa.
aquatica!
*Primula sinensis!
Lysimachia Ephemerum.
Anagallis arvensis.
Webbiana.
Nicotiana rustica.
Anchusa ochroleuca.
Myosotis cæspitosa.
Page 359
Stachys sylvatica.
Gilia capitata.
Euphorbia segetalis.
Rumex arifolius.
scutatus.
Juncus lampocarpus.
uliginosus.
In addition to the publications before cited the following may be named as
containing valuable information on the subject of this chapter.
Jæger, 'Missbild. Gewächs.,' 1814, p. 83, Trifolium repens. For other accounts of similar
malformations in the same plant, see Schmitz, 'Linnæa,' xv, p. 268. Unger, 'Flora' (B. Z.)
xxv, p. 369. Caspary, 'Schrift. der. Physik. ökon. Gesellsch. zu Königsberg,' 2, 1861, p. 51,
tabs. ii, iii. Fleischer, 'Missbilld. verschied. Cult. Pflanz.,' 1862, p. 55, &c., t. v, vii, &c. For
Primula see Brongniart, 'Ann. Sc. Nat.,' ser. 2, t. i, p. 308. A. P. and Alph. De Candolle in
'Neue Denkschrift.' Morren, C., 'Bull. Acad. Roy. Belg.,' xix, part 2, p. 539. Molkenboer,
'Tijdschr. voor Natuurl. Geschied.,' 1843, p. 355, tabs. vi, vii. Marchand, 'Adansonia,' iv, p.
167 and p. 159. Anagallis, p. 171, Lonicera, p. 83, Juncus. For other plants see Fresenius,
'Mus. Senk.,' 2, p. 35, &c. Norman, 'Ann. Sc. Nat.,' ser. 4, 1858, vol. ix, p. 220. Christ,
'Flora' (B. Z.) 1867, p. 376, tabs. v, vi, Stachys. Cramer, 'Bildungsabweich.,' p. 26, &c.
Baillon, 'Adansonia,' ii, p. 300. Moquin-Tandon, 'El. Ter. Veg.,' p. 230. Schauer's
translation, p. 220. Hallier, 'Phytopathologie,' p. 160.
FOOTNOTES:
[245] Engelmann makes use of the word frondescence in the same cases. 'De Anthol.,' p. 32,
§ 38, while Morren adopts the term Phyllomorphy, 'Lobelia,' p. 95.
[246] See Schlechtendal, 'Bot. Zeit.,' vol. xv, 1857, p. 873; also Marchand, 'Adansonia,' iv, p.
156.
[247] For instances of similar changes in Composites, see De Candolle, 'Prod.,' t. vi, p. 571,
Centaurea Jacea phyllocephala. Clos, 'Ann. Sc. Nat.,' ser. iii, tom. xvi, 1851, p. 41. 'Science
Gossip,' 1865, p. 104, &c.
[248] Kickx, 'Bull. Acad. Belg.,' t. xviii, part 2, p. 288.
[249] Weber, 'Verhandl. Nat. Hist. Vereins. f. Preuss.,' &c., 1860, p. 381.
[250] Weber, loc. cit.
Gilia capitata.
Euphorbia segetalis.
Rumex arifolius.
scutatus.
Juncus lampocarpus.
uliginosus.
In addition to the publications before cited the following may be named as
containing valuable information on the subject of this chapter.
Jæger, 'Missbild. Gewächs.,' 1814, p. 83, Trifolium repens. For other accounts of similar
malformations in the same plant, see Schmitz, 'Linnæa,' xv, p. 268. Unger, 'Flora' (B. Z.)
xxv, p. 369. Caspary, 'Schrift. der. Physik. ökon. Gesellsch. zu Königsberg,' 2, 1861, p. 51,
tabs. ii, iii. Fleischer, 'Missbilld. verschied. Cult. Pflanz.,' 1862, p. 55, &c., t. v, vii, &c. For
Primula see Brongniart, 'Ann. Sc. Nat.,' ser. 2, t. i, p. 308. A. P. and Alph. De Candolle in
'Neue Denkschrift.' Morren, C., 'Bull. Acad. Roy. Belg.,' xix, part 2, p. 539. Molkenboer,
'Tijdschr. voor Natuurl. Geschied.,' 1843, p. 355, tabs. vi, vii. Marchand, 'Adansonia,' iv, p.
167 and p. 159. Anagallis, p. 171, Lonicera, p. 83, Juncus. For other plants see Fresenius,
'Mus. Senk.,' 2, p. 35, &c. Norman, 'Ann. Sc. Nat.,' ser. 4, 1858, vol. ix, p. 220. Christ,
'Flora' (B. Z.) 1867, p. 376, tabs. v, vi, Stachys. Cramer, 'Bildungsabweich.,' p. 26, &c.
Baillon, 'Adansonia,' ii, p. 300. Moquin-Tandon, 'El. Ter. Veg.,' p. 230. Schauer's
translation, p. 220. Hallier, 'Phytopathologie,' p. 160.
FOOTNOTES:
[245] Engelmann makes use of the word frondescence in the same cases. 'De Anthol.,' p. 32,
§ 38, while Morren adopts the term Phyllomorphy, 'Lobelia,' p. 95.
[246] See Schlechtendal, 'Bot. Zeit.,' vol. xv, 1857, p. 873; also Marchand, 'Adansonia,' iv, p.
156.
[247] For instances of similar changes in Composites, see De Candolle, 'Prod.,' t. vi, p. 571,
Centaurea Jacea phyllocephala. Clos, 'Ann. Sc. Nat.,' ser. iii, tom. xvi, 1851, p. 41. 'Science
Gossip,' 1865, p. 104, &c.
[248] Kickx, 'Bull. Acad. Belg.,' t. xviii, part 2, p. 288.
[249] Weber, 'Verhandl. Nat. Hist. Vereins. f. Preuss.,' &c., 1860, p. 381.
[250] Weber, loc. cit.
Page 360
[251] Sauter, 'Flora v. Bot. Zeit.,' 1831, p. 11.
[252] 'Descr. et Icon. Plant.' tab. 20.
[253] For references see p. 115; see also to Eichler, 'Excurs. Morpholog. de format. flor.
Gymnosperm.,' in "Mart. Flor. Brasil," abstracted in English in 'Natural History Review,' April,
1864.
[254] "Calyx tunc plane non differt a foliis proxime ipsi præcedentibus." Wolff, 'Theor.
Gener.,' § 114. Linn., 'Proleps.,' § 6. Goethe, 'Versuch.,' §§ 31–38.
[255] 'Bull. Soc. Bot. France,' vol. viii, 1861, p. 697.
[256] 'Organ. Véget.,' t. i, p. 492, pl. xxxii, f. 6.
[257] This distinction between laminar and vaginal venation is well seen in cases like
Mussaenda, Calycophyllum, or Dipterocarpus, where the enlarged calycine segment has a
strictly vaginal arrangement of its veins, very different from that which occurs in the true leaf-
blades. These are cases, therefore, where the sheath of the leaf is unusually enlarged, and are
not to be referred, as is often done, to metamorphosis of one or more sepals to perfect leaves.
Prolified roses, cherries, &c., furnish frequently parallel cases. With reference to Mussaenda,
C. Morren held the view that the petal-like sepal was really a bract adherent to the calyx, and
incorporating with itself one of the calycine lobes—"soudée au calice et ayant dévorée, en
englobant dans sa propre masse, un lobe calicinal." The Belgian savant considers this
somewhat improbable explanation as supported by a case wherein there were five calyx lobes
of uniform size, and a detached feather-veined leaf proceeding from the side of the ovary
lower down ('Bull. Acad. Belg.,' xvii, p. 17, Fuchsia, p. 169).
[258] In this order Agrostemma Githago offers an illustration of a normally leafy calyx.
[259] 'Bull. Bot.,' i, p. 6.
[260] Wolff's original opinion was that the stamens were equivalent to so many buds placed
in the axil of the petals or sepals (see 'Theoria Generationis,' 1759, § 114)—an opinion which
more recently has received the support of Agardh and Endlicher. Wolff himself, however,
seems to have abandoned his original notion, for in his memoir, "De formatione intestinorum
præcipue tum et de amnio spurio aliisque partibus embryonis gallinacei, nondum visis," &c.,
in 'Comm. Acad. Petrop.,' xii, p. 403, anno 1766, he considers the stamens as essentially
leaves. See also Linn. 'Prolepsis,' § viii; Goethe, 'Metam.,' § 46.
[261] Müller (Argov.), in 'Mém. Soc. Phys. et d'Hist. Nat. Genev.,' t. xvii.
[262] "If we keep in view the observations which have now been made, we shall not fail to
recognise the leaf in all seed-vessels, notwithstanding their manifold forms, their variable
[252] 'Descr. et Icon. Plant.' tab. 20.
[253] For references see p. 115; see also to Eichler, 'Excurs. Morpholog. de format. flor.
Gymnosperm.,' in "Mart. Flor. Brasil," abstracted in English in 'Natural History Review,' April,
1864.
[254] "Calyx tunc plane non differt a foliis proxime ipsi præcedentibus." Wolff, 'Theor.
Gener.,' § 114. Linn., 'Proleps.,' § 6. Goethe, 'Versuch.,' §§ 31–38.
[255] 'Bull. Soc. Bot. France,' vol. viii, 1861, p. 697.
[256] 'Organ. Véget.,' t. i, p. 492, pl. xxxii, f. 6.
[257] This distinction between laminar and vaginal venation is well seen in cases like
Mussaenda, Calycophyllum, or Dipterocarpus, where the enlarged calycine segment has a
strictly vaginal arrangement of its veins, very different from that which occurs in the true leaf-
blades. These are cases, therefore, where the sheath of the leaf is unusually enlarged, and are
not to be referred, as is often done, to metamorphosis of one or more sepals to perfect leaves.
Prolified roses, cherries, &c., furnish frequently parallel cases. With reference to Mussaenda,
C. Morren held the view that the petal-like sepal was really a bract adherent to the calyx, and
incorporating with itself one of the calycine lobes—"soudée au calice et ayant dévorée, en
englobant dans sa propre masse, un lobe calicinal." The Belgian savant considers this
somewhat improbable explanation as supported by a case wherein there were five calyx lobes
of uniform size, and a detached feather-veined leaf proceeding from the side of the ovary
lower down ('Bull. Acad. Belg.,' xvii, p. 17, Fuchsia, p. 169).
[258] In this order Agrostemma Githago offers an illustration of a normally leafy calyx.
[259] 'Bull. Bot.,' i, p. 6.
[260] Wolff's original opinion was that the stamens were equivalent to so many buds placed
in the axil of the petals or sepals (see 'Theoria Generationis,' 1759, § 114)—an opinion which
more recently has received the support of Agardh and Endlicher. Wolff himself, however,
seems to have abandoned his original notion, for in his memoir, "De formatione intestinorum
præcipue tum et de amnio spurio aliisque partibus embryonis gallinacei, nondum visis," &c.,
in 'Comm. Acad. Petrop.,' xii, p. 403, anno 1766, he considers the stamens as essentially
leaves. See also Linn. 'Prolepsis,' § viii; Goethe, 'Metam.,' § 46.
[261] Müller (Argov.), in 'Mém. Soc. Phys. et d'Hist. Nat. Genev.,' t. xvii.
[262] "If we keep in view the observations which have now been made, we shall not fail to
recognise the leaf in all seed-vessels, notwithstanding their manifold forms, their variable
Page 361
structure, and different combinations."—(Goethe, 'Metam.,' § 78.) Wolff, 'N. Comm. Acad.
Petrop.,' 1766, xii, p. 403, expresses precisely the same opinion as to the nature of the seed-
vessel.
[263] 'El. Terat. Veg.,' p. 205.
[264] 'Ann. Sc. Nat.,' 4th series, vol. ix, p. 209.
[265] 'Adansonia,' iv, p. 70. A similar deviation has been observed by M. van Tieghem in the
ovary of Tropæolum majus, 'Bull. Soc. Bot. Fr.,' 1865, p. 411.
[266] Planchon et Marès, 'Ann. Sc. Nat.,' ser. 5, vol. vi, 1866, p. 228, tab. xii.
[267] 'Act. Acad. Nat. Cur.,' 22, 11. 1850, p. 543, t. v, vi.
[268] 'Neue Denkschrift der allg. Schweiz. Gesellsch.,' band v. p. 9, tab. 3, 4.
[269] 'Ann. Sc. Nat.,' 2 ser., vol. i, p. 308, pl. ix, c.
[270] 'Adansonia,' vol. iv, pp. 159, 171.
[271] 'Bildungsabweichungen,' &c., tab. iv, figs. 1, 2, 21, 28, 29, &c.
[272] 'Bull. Soc. Bot. France,' viii, p. 395.
[273] 'Notulæ,' p. 125, atlas, pl. xxxv; and 'Journals of Travels,' 1847, p. 475, Lonicera.
[274] 'Ann. Science Nat.,' 3rd ser., vol. ix, p. 86, tabs. 5, 6.
[275] 'Comptes Rendus,' vol. xviii, March 25th, 1864, and 'Ann. Sc. Nat.,' 3 ser., vol. ii, p.
32.
[276] 'Mém. Acad. Sc. Toulous.,' ser. 5, vol. iii.
[277] 'Bildungsabweich. Pflanz. Famil.,' p. 89, tab. xi.
[278] 'Bull. Soc. Bot. Fr.,' 1865, p. 411.
[279] Translated in 'Ann. Sc. Nat.,' 4th series, t. xiv, p. 24.
[280] The calyx is not unfrequently excepted.
[281] 'Bull. Bot.,' t. i, p. 6.
[282] Lindley, 'Theor. Horticult.,' ed. 2, p. 84, f. 17.
Petrop.,' 1766, xii, p. 403, expresses precisely the same opinion as to the nature of the seed-
vessel.
[263] 'El. Terat. Veg.,' p. 205.
[264] 'Ann. Sc. Nat.,' 4th series, vol. ix, p. 209.
[265] 'Adansonia,' iv, p. 70. A similar deviation has been observed by M. van Tieghem in the
ovary of Tropæolum majus, 'Bull. Soc. Bot. Fr.,' 1865, p. 411.
[266] Planchon et Marès, 'Ann. Sc. Nat.,' ser. 5, vol. vi, 1866, p. 228, tab. xii.
[267] 'Act. Acad. Nat. Cur.,' 22, 11. 1850, p. 543, t. v, vi.
[268] 'Neue Denkschrift der allg. Schweiz. Gesellsch.,' band v. p. 9, tab. 3, 4.
[269] 'Ann. Sc. Nat.,' 2 ser., vol. i, p. 308, pl. ix, c.
[270] 'Adansonia,' vol. iv, pp. 159, 171.
[271] 'Bildungsabweichungen,' &c., tab. iv, figs. 1, 2, 21, 28, 29, &c.
[272] 'Bull. Soc. Bot. France,' viii, p. 395.
[273] 'Notulæ,' p. 125, atlas, pl. xxxv; and 'Journals of Travels,' 1847, p. 475, Lonicera.
[274] 'Ann. Science Nat.,' 3rd ser., vol. ix, p. 86, tabs. 5, 6.
[275] 'Comptes Rendus,' vol. xviii, March 25th, 1864, and 'Ann. Sc. Nat.,' 3 ser., vol. ii, p.
32.
[276] 'Mém. Acad. Sc. Toulous.,' ser. 5, vol. iii.
[277] 'Bildungsabweich. Pflanz. Famil.,' p. 89, tab. xi.
[278] 'Bull. Soc. Bot. Fr.,' 1865, p. 411.
[279] Translated in 'Ann. Sc. Nat.,' 4th series, t. xiv, p. 24.
[280] The calyx is not unfrequently excepted.
[281] 'Bull. Bot.,' t. i, p. 6.
[282] Lindley, 'Theor. Horticult.,' ed. 2, p. 84, f. 17.
Page 362
[283] Gris, 'Bull. Soc. Bot. Fr.,' 1858, vol. v, p. 261, and 'Ann. Sc. Nat.,' ser. 4, vol. ix, p. 80.
Planchon, 'Flore des Serres,' vol. i, 1856, p. 129.
[284] 'Flora,' 1856, p. 711.
[285] 'Theory of Horticult.,' ed. 2, p. 90, f. 25.
[286] As considerable interest attaches to the "Plymouth strawberry," and very little is known
of it in this country, or on the continent, the author gladly avails himself of this opportunity of
inserting an account of it, for which he is indebted to the kindness of Dr. Robert Hogg.—The
Plymouth Strawberry (Fragaria vesca fructu hispido) is a sort of botanical Dodo upon which
many have written, and which few have seen. Many years have elapsed since it was first
discovered; and although a century and a half have passed since there was any evidence of its
existence, it serves still as an illustration for students in morphology of one of those strange
abnormal structures with which the vegetable kingdom abounds.
It is to old John Tradescant we are indebted for the earliest record of this plant. Johnson, in his
edition of 'Gerard,' says; "Mr. John Tradescant hath told me that he was the first that tooke
notice of this strawberry, and that in a woman's garden at Plimouth, whose daughter had
gathered and set the roots in her garden, in stead of the common strawberry; but she, finding
the fruit not to answer her expectation, intended to throw it away; which labour he spared her
in taking it and bestowing it among the louers of such varieties, in whose garden it is yet
preserved." Doubtless one of those "lovers" was his friend John Parkinson, who, in the year
1629, thus wrote concerning it: "One strawberry more I promised to shew you, which,
although it be a wilde kinde, and of no vse for meate, yet I would not let this discourse passe
without giuing you the knowledge of it. It is in leafe much like vnto the ordinary, but differeth
in that the flower, if it haue any, is greene, or rather it beareth a small head of greene leaues,
many set thicke together like vnto a double ruffe, in the midst whereof standeth the fruit,
which, when it is ripe, sheweth to be soft and somewhat reddish, like vnto a strawberry, but
with many small harmlesse prickles on them which may be eaten and chewed in the mouth
without any maner of offence and is somewhat pleasant as a strawberry; it is no great bearer,
but those it doth beare, are set at the toppes of the stalks close together, pleasant to behold, and
fit for a gentlewoman to weare on her arme, &c., as a rairitie in stead of a flower."
Merret, in his 'Pinax.' published in 1667, says he found it growing in the woods of Hyde Park
and Hampstead, and Zanoni was the first to figure it (with the exception of Parkinson's rude
woodcut) in his 'Istoria Botanica,' published in 1675. It is mentioned by Morison and also by
Ray, the latter of whom inserts it in his Synopsis, but without any habitat; though in his
'Historia Plantarum' he says: "Cantabrigiæ in horto per aliquot annos colui." From this time
henceforth the Plymouth strawberry has become a botanical Dodo, nothing more having been
seen or heard of it except the mere record of the name. In 1766, M. Duchesne informed the
world of the generosity of "M. Monti, Docteur de Philosophie et de Médecine à Boulogne en
Planchon, 'Flore des Serres,' vol. i, 1856, p. 129.
[284] 'Flora,' 1856, p. 711.
[285] 'Theory of Horticult.,' ed. 2, p. 90, f. 25.
[286] As considerable interest attaches to the "Plymouth strawberry," and very little is known
of it in this country, or on the continent, the author gladly avails himself of this opportunity of
inserting an account of it, for which he is indebted to the kindness of Dr. Robert Hogg.—The
Plymouth Strawberry (Fragaria vesca fructu hispido) is a sort of botanical Dodo upon which
many have written, and which few have seen. Many years have elapsed since it was first
discovered; and although a century and a half have passed since there was any evidence of its
existence, it serves still as an illustration for students in morphology of one of those strange
abnormal structures with which the vegetable kingdom abounds.
It is to old John Tradescant we are indebted for the earliest record of this plant. Johnson, in his
edition of 'Gerard,' says; "Mr. John Tradescant hath told me that he was the first that tooke
notice of this strawberry, and that in a woman's garden at Plimouth, whose daughter had
gathered and set the roots in her garden, in stead of the common strawberry; but she, finding
the fruit not to answer her expectation, intended to throw it away; which labour he spared her
in taking it and bestowing it among the louers of such varieties, in whose garden it is yet
preserved." Doubtless one of those "lovers" was his friend John Parkinson, who, in the year
1629, thus wrote concerning it: "One strawberry more I promised to shew you, which,
although it be a wilde kinde, and of no vse for meate, yet I would not let this discourse passe
without giuing you the knowledge of it. It is in leafe much like vnto the ordinary, but differeth
in that the flower, if it haue any, is greene, or rather it beareth a small head of greene leaues,
many set thicke together like vnto a double ruffe, in the midst whereof standeth the fruit,
which, when it is ripe, sheweth to be soft and somewhat reddish, like vnto a strawberry, but
with many small harmlesse prickles on them which may be eaten and chewed in the mouth
without any maner of offence and is somewhat pleasant as a strawberry; it is no great bearer,
but those it doth beare, are set at the toppes of the stalks close together, pleasant to behold, and
fit for a gentlewoman to weare on her arme, &c., as a rairitie in stead of a flower."
Merret, in his 'Pinax.' published in 1667, says he found it growing in the woods of Hyde Park
and Hampstead, and Zanoni was the first to figure it (with the exception of Parkinson's rude
woodcut) in his 'Istoria Botanica,' published in 1675. It is mentioned by Morison and also by
Ray, the latter of whom inserts it in his Synopsis, but without any habitat; though in his
'Historia Plantarum' he says: "Cantabrigiæ in horto per aliquot annos colui." From this time
henceforth the Plymouth strawberry has become a botanical Dodo, nothing more having been
seen or heard of it except the mere record of the name. In 1766, M. Duchesne informed the
world of the generosity of "M. Monti, Docteur de Philosophie et de Médecine à Boulogne en
Page 363
Italie," who divided with him a dried specimen taken from his own herbarium, "Ce présent
prétieux m'ôte toute incertitude sur la nature de ce Fraisier et sur ses caractères monstrueux. Il
paroît ne pas avoir aujourd'hui plus d'existence."
[287] 'Bull. Soc. Bot. France,' 1856, vol. iii, p. 477.
[288] 'Ann. Sc. Nat.,' 3 ser., vol. ix, p. 86, tabs. v, vi.
[289] 'Bull. Soc. Bot. France,' vol. viii, 1861, p. 695.
[290] Ibid., vol. iii, 1856, p. 475.
[291] 'Flora,' 1856, p. 712.
[292] 'Trans. Linn. Soc.,' vol. xxvi, p. 37.
[293] 'Bull. Soc. Bot. France,' 1862, vol. ix, p. 36, tab. i, and also p. 291.
[294] Ibid., 1857, vol. iv, p. 761.
CHAPTER II.
METAMORPHY OF THE FLORAL ORGANS.
One of the main arguments adduced by Goethe and others in support of the
now generally received doctrine of the essential morphological identity of
the various whorls of the flower is derived from the frequent appearance of
one organ in the guise of another. The several parts of the flower become, as
it is said, metamorphosed; sometimes the change is complete, while at other
times there may be every conceivable intermediate condition between one
form and another. The sense in which the terms metamorphosis,
substitution, transformation, and the like, are herein used has already been
explained. For the convenience of arrangement, metamorphosis of the parts
of the flower may be divided into several subdivisions, according to the
particular organ affected, and according to the special kind or degree of
prétieux m'ôte toute incertitude sur la nature de ce Fraisier et sur ses caractères monstrueux. Il
paroît ne pas avoir aujourd'hui plus d'existence."
[287] 'Bull. Soc. Bot. France,' 1856, vol. iii, p. 477.
[288] 'Ann. Sc. Nat.,' 3 ser., vol. ix, p. 86, tabs. v, vi.
[289] 'Bull. Soc. Bot. France,' vol. viii, 1861, p. 695.
[290] Ibid., vol. iii, 1856, p. 475.
[291] 'Flora,' 1856, p. 712.
[292] 'Trans. Linn. Soc.,' vol. xxvi, p. 37.
[293] 'Bull. Soc. Bot. France,' 1862, vol. ix, p. 36, tab. i, and also p. 291.
[294] Ibid., 1857, vol. iv, p. 761.
CHAPTER II.
METAMORPHY OF THE FLORAL ORGANS.
One of the main arguments adduced by Goethe and others in support of the
now generally received doctrine of the essential morphological identity of
the various whorls of the flower is derived from the frequent appearance of
one organ in the guise of another. The several parts of the flower become, as
it is said, metamorphosed; sometimes the change is complete, while at other
times there may be every conceivable intermediate condition between one
form and another. The sense in which the terms metamorphosis,
substitution, transformation, and the like, are herein used has already been
explained. For the convenience of arrangement, metamorphosis of the parts
of the flower may be divided into several subdivisions, according to the
particular organ affected, and according to the special kind or degree of
Page 364
change manifested, the main subdivisions being here classed as Sepalody,
Petalody, Staminody, and Pistillody.
Sepalody of the petals.—This change, spoken of by most authors as
retrograde metamorphosis of the petals into sepals, or as a substitution of
sepals for petals, is obviously a condition that is in most cases hardly
distinguishable from virescence of the corolla, or from multiplication of the
sepals. Nor is this of much consequence unless there are some special
structural features which render the discrimination a matter of importance,
in which case the difficulty is generally easily surmounted. The flower of
the Saint-Valèry Apple may perhaps be cited under this head. In the flower
in question there are neither stamens nor petals, unless the second or inner
of sepals be considered as sepaloid petals (fig. 152).
Fig. 152.—Flower of St. Valèry apple, with sepaloid petals.
M. Alph. de Candolle[295] describes an instance in Primula Auricula in
which the corolla had assumed the appearance of the calyx, but neither
calyx nor corolla in this case possessed perfect stomata.
This malformation is much less common than the converse one of
calycanthemy. Many of the recorded instances of so-called metamorphosis
Petalody, Staminody, and Pistillody.
Sepalody of the petals.—This change, spoken of by most authors as
retrograde metamorphosis of the petals into sepals, or as a substitution of
sepals for petals, is obviously a condition that is in most cases hardly
distinguishable from virescence of the corolla, or from multiplication of the
sepals. Nor is this of much consequence unless there are some special
structural features which render the discrimination a matter of importance,
in which case the difficulty is generally easily surmounted. The flower of
the Saint-Valèry Apple may perhaps be cited under this head. In the flower
in question there are neither stamens nor petals, unless the second or inner
of sepals be considered as sepaloid petals (fig. 152).
Fig. 152.—Flower of St. Valèry apple, with sepaloid petals.
M. Alph. de Candolle[295] describes an instance in Primula Auricula in
which the corolla had assumed the appearance of the calyx, but neither
calyx nor corolla in this case possessed perfect stomata.
This malformation is much less common than the converse one of
calycanthemy. Many of the recorded instances of so-called metamorphosis
Page 365
of the parts of the flower to sepals have occurred in monocotyledonous
plants, or others in which the calyx and corolla are of the same colour, and
constitute what is frequently termed the perianth; and as this is usually
brightly coloured (not green) it is more convenient to group the
metamorphoses in question under the general term Petalody, which thus
includes all those cases in which the organs of the flower appear in the form
of coloured petal-like organs, whether they be true petals or segments of a
coloured perianth. As the morphological difference between the organs is
one of position merely, there is little objection to be raised to this course,
the less so as the term petalody merely conveys an idea of resemblance and
not of absolute identity.
Petaloid coloration of the ordinary leaves, or of the bracts, is mentioned
under the chapter relating to colour.
Petalody of the calyx—Calycanthemy.—As with the bracts, so the calyx
in certain instances is naturally coloured, as in Delphinium, Tropæolum, and
others. In Mussænda, Calycophyllum, Usteria, &c., one or more of the
calyx lobes become enlarged normally. Considered teratologically, petaloid
coloration of the sepals is either general or partial; in the latter case the
nerves retain their green colour longest. There is in cultivation a variety of
the primrose called Primula calycanthema, in which the upper part of the
calyx becomes coloured, so that the flower seems to have two corollas
placed one within the other; a similar thing happens in Mimulus, in which
plant, as the calyx is permanent while the corolla is deciduous, the coloured
calyx is a great advantage in a horticultural point of view. Morren[296] says
that in order to produce the fine colour of the calyx of Primula officinalis
(var. smaragdina) the Belgian gardeners cut away the corolla in a very early
stage, and that in consequence the colouring matter proper to the corolla is
developed in the tube of the calyx, the edges of the limb remaining green,
the middle of the limb being purple (Primula tricolor).
plants, or others in which the calyx and corolla are of the same colour, and
constitute what is frequently termed the perianth; and as this is usually
brightly coloured (not green) it is more convenient to group the
metamorphoses in question under the general term Petalody, which thus
includes all those cases in which the organs of the flower appear in the form
of coloured petal-like organs, whether they be true petals or segments of a
coloured perianth. As the morphological difference between the organs is
one of position merely, there is little objection to be raised to this course,
the less so as the term petalody merely conveys an idea of resemblance and
not of absolute identity.
Petaloid coloration of the ordinary leaves, or of the bracts, is mentioned
under the chapter relating to colour.
Petalody of the calyx—Calycanthemy.—As with the bracts, so the calyx
in certain instances is naturally coloured, as in Delphinium, Tropæolum, and
others. In Mussænda, Calycophyllum, Usteria, &c., one or more of the
calyx lobes become enlarged normally. Considered teratologically, petaloid
coloration of the sepals is either general or partial; in the latter case the
nerves retain their green colour longest. There is in cultivation a variety of
the primrose called Primula calycanthema, in which the upper part of the
calyx becomes coloured, so that the flower seems to have two corollas
placed one within the other; a similar thing happens in Mimulus, in which
plant, as the calyx is permanent while the corolla is deciduous, the coloured
calyx is a great advantage in a horticultural point of view. Morren[296] says
that in order to produce the fine colour of the calyx of Primula officinalis
(var. smaragdina) the Belgian gardeners cut away the corolla in a very early
stage, and that in consequence the colouring matter proper to the corolla is
developed in the tube of the calyx, the edges of the limb remaining green,
the middle of the limb being purple (Primula tricolor).
Page 366
Fig. 153.—Flower of Mimulus, with petaloid calyx.
Under this head may be mentioned the occurrence of tubular sepals in place
of the ordinary flat ones in Helleborus olympicus; only two of the sepals
were thus affected in a specimen recently observed—a third exhibited an
intermediate condition.
The normal coloration of the calyx occurs most frequently in polysepalous
calyces; teratological coloration, on the other hand, occurs especially in
gamosepalous flowers. This assertion is borne out by the frequency of the
change in the plants already mentioned, and also in the following:—
Campanula persicifolia, Anagallis arvensis, Gloxinia, Syringa persica,[297]
Calceolaria, &c. &c. In the last-named plant one or more of the lobes of the
calyx may frequently be seen replaced by a slipper-like petal.
Among polysepalous plants petaloid sepals have been observed in
Ranunculus auricomus, Rubus cæsius, &c. Fleischer also describes a case
Under this head may be mentioned the occurrence of tubular sepals in place
of the ordinary flat ones in Helleborus olympicus; only two of the sepals
were thus affected in a specimen recently observed—a third exhibited an
intermediate condition.
The normal coloration of the calyx occurs most frequently in polysepalous
calyces; teratological coloration, on the other hand, occurs especially in
gamosepalous flowers. This assertion is borne out by the frequency of the
change in the plants already mentioned, and also in the following:—
Campanula persicifolia, Anagallis arvensis, Gloxinia, Syringa persica,[297]
Calceolaria, &c. &c. In the last-named plant one or more of the lobes of the
calyx may frequently be seen replaced by a slipper-like petal.
Among polysepalous plants petaloid sepals have been observed in
Ranunculus auricomus, Rubus cæsius, &c. Fleischer also describes a case
Page 367
of this kind in Carum carui.[298]
It will be seen from the above that in the majority of cases there is no real
metamorphosis or substitution of petal for calyx, but simply an alteration in
colour; nevertheless, a change in form may accompany a change of colour:
this happens especially if there has been any displacement of organs. Thus,
if, in an orchidaceous plant, a sepal be displaced from any cause, or a petal
be twisted out of its natural position to occupy the place of an absent sepal,
that petal will be sepal-like in form, and vice versâ.
Petalody of the stamens.—A petaloid condition of the stamens is one of
the commonest of all malformations. A large number of so-called double
flowers (flores pleni)[299] owe their peculiar appearance to this
circumstance.
It is necessary to distinguish carefully this petaloid development of the
stamens from the corresponding condition of the pistils, and from that kind
of doubling which is a result of multiplication of the corolla, as in Datura,
Campanula, Primula, &c. (flores duplices, triplices, &c.), or from that
produced by true median prolification (flores geminati, &c.).
In cases of true petaloid development of the stamens there are usually
numerous intermediate forms between that of the true petals and that of the
perfect stamens; indeed, in Nymphæa, Canna, and in some other plants,
such a transition occurs normally. Petalody of the stamens may occur either
without material change in the flower or it may exist in combination or in
conjunction with an increased development of parts (Multiplication), or
with a similar change in the carpels, and it is either partial or complete.
Among the flowers in which petaloid development of the stamens happens
most frequently may be mentioned those in which the calyx is normally
coloured, as in Nigella damascena, Aguilegia, and Delphinium.
It will be seen from the above that in the majority of cases there is no real
metamorphosis or substitution of petal for calyx, but simply an alteration in
colour; nevertheless, a change in form may accompany a change of colour:
this happens especially if there has been any displacement of organs. Thus,
if, in an orchidaceous plant, a sepal be displaced from any cause, or a petal
be twisted out of its natural position to occupy the place of an absent sepal,
that petal will be sepal-like in form, and vice versâ.
Petalody of the stamens.—A petaloid condition of the stamens is one of
the commonest of all malformations. A large number of so-called double
flowers (flores pleni)[299] owe their peculiar appearance to this
circumstance.
It is necessary to distinguish carefully this petaloid development of the
stamens from the corresponding condition of the pistils, and from that kind
of doubling which is a result of multiplication of the corolla, as in Datura,
Campanula, Primula, &c. (flores duplices, triplices, &c.), or from that
produced by true median prolification (flores geminati, &c.).
In cases of true petaloid development of the stamens there are usually
numerous intermediate forms between that of the true petals and that of the
perfect stamens; indeed, in Nymphæa, Canna, and in some other plants,
such a transition occurs normally. Petalody of the stamens may occur either
without material change in the flower or it may exist in combination or in
conjunction with an increased development of parts (Multiplication), or
with a similar change in the carpels, and it is either partial or complete.
Among the flowers in which petaloid development of the stamens happens
most frequently may be mentioned those in which the calyx is normally
coloured, as in Nigella damascena, Aguilegia, and Delphinium.
Page 368
M. Alph. de Candolle, in the 'Neue Denkschriften,' 1841, described and
figured a singular form of Viola odorata, known under the name of
"Bruneau," in Switzerland, in which the stamens are absent, and their place
supplied by a second row of petals, within which is a third series of petals,
representing, says M. de Candolle, the inner row of stamens that theory
suggests should exist in the natural condition. Moreover, the carpels in this
variety are five in number instead of three. In Erica Tetralix the corolla may
not unfrequently be found divided to the base into its constituent petals, and
the place of the stamens occupied by a series of petal-like structures entirely
destitute of anther.
In monocotyledonous flowers, especially those with a coloured perianth, the
substitution of segments of the perianth for stamens occurs not
unfrequently. M. Seringe has observed this in the stamens of Lilium
Martagon, and there is in cultivation a variety of the white lily, Lilium
candidum, sometimes called the double white lily, in which the segments of
the perianth, in place of being arranged in two rows, are greatly increased in
number, and disposed in a spiral manner. In these flowers, not only are the
stamens and pistils thus modified, but also the upper leaves of the stem. In
so-called double tulips there is likewise a replacement of stamens by
coloured segments of the perianth, but this happens generally in connection
with an increase in the number of organs. Moquin-Tandon remarks having
seen in a garden in the environs of Montpelier a tulip, the stamens of which
showed all possible stages of transition between the form proper to them
and that of the perianth. The pistil in this case was transformed into several
small leaves. Similar appearances have been observed in Iris, Hyacinths,
Narcissus, Colchicum, and Crocus. M. Fournier[300] describes a flower of
Narcissus Tazetta from within the normal perianth of which sprang a second
one, equally provided with a cup and occupying the space usually filled by
the stamens. Flowers of Narcissus poeticus may also be met with in which
figured a singular form of Viola odorata, known under the name of
"Bruneau," in Switzerland, in which the stamens are absent, and their place
supplied by a second row of petals, within which is a third series of petals,
representing, says M. de Candolle, the inner row of stamens that theory
suggests should exist in the natural condition. Moreover, the carpels in this
variety are five in number instead of three. In Erica Tetralix the corolla may
not unfrequently be found divided to the base into its constituent petals, and
the place of the stamens occupied by a series of petal-like structures entirely
destitute of anther.
In monocotyledonous flowers, especially those with a coloured perianth, the
substitution of segments of the perianth for stamens occurs not
unfrequently. M. Seringe has observed this in the stamens of Lilium
Martagon, and there is in cultivation a variety of the white lily, Lilium
candidum, sometimes called the double white lily, in which the segments of
the perianth, in place of being arranged in two rows, are greatly increased in
number, and disposed in a spiral manner. In these flowers, not only are the
stamens and pistils thus modified, but also the upper leaves of the stem. In
so-called double tulips there is likewise a replacement of stamens by
coloured segments of the perianth, but this happens generally in connection
with an increase in the number of organs. Moquin-Tandon remarks having
seen in a garden in the environs of Montpelier a tulip, the stamens of which
showed all possible stages of transition between the form proper to them
and that of the perianth. The pistil in this case was transformed into several
small leaves. Similar appearances have been observed in Iris, Hyacinths,
Narcissus, Colchicum, and Crocus. M. Fournier[300] describes a flower of
Narcissus Tazetta from within the normal perianth of which sprang a second
one, equally provided with a cup and occupying the space usually filled by
the stamens. Flowers of Narcissus poeticus may also be met with in which
Page 369
the stamens are replaced by six distinct segments exactly resembling those
of the perianth in miniature.[301]
Fig. 154.—Double columbine, Aquilegia—petalody of the filament.
From an examination of these flowers it becomes evident that petalification
is brought about in different flowers in different ways; sometimes it is the
filament which becomes petaloid, sometimes the anther-lobes, while at
other times it is the connective which assumes the appearance of petals.[302]
For instance, in Solanum tuberosum, S. Dulcamara, in Anagallis, in
Fuchsia, and some other plants, the anther-lobes themselves become
petaloid, while the filament remains unchanged.
In gardens two distinct varieties of Columbine are cultivated, the one in
which the filaments are dilated into the form of flat petals almost entirely or
quite destitute of anthers, while in the other the filament is present in its
of the perianth in miniature.[301]
Fig. 154.—Double columbine, Aquilegia—petalody of the filament.
From an examination of these flowers it becomes evident that petalification
is brought about in different flowers in different ways; sometimes it is the
filament which becomes petaloid, sometimes the anther-lobes, while at
other times it is the connective which assumes the appearance of petals.[302]
For instance, in Solanum tuberosum, S. Dulcamara, in Anagallis, in
Fuchsia, and some other plants, the anther-lobes themselves become
petaloid, while the filament remains unchanged.
In gardens two distinct varieties of Columbine are cultivated, the one in
which the filaments are dilated into the form of flat petals almost entirely or
quite destitute of anthers, while in the other the filament is present in its
Page 370
usual form, but the anther is developed in the shape of a tubular hood or
spur.
De Candolle[303] observes that in the Ranunculaceæ the species of Clematis
become double by the expansion of the filament, those of Ranunculus by
the dilatation of the anther, and those of Helleborus by the petal-like
development of both filament and anther. In some cases even on the same
plant all three modifications may be seen, as in Camellias, some of which
may be found with petaloid filaments with anthers on the top, others with
the filaments unchanged, but supporting petaloid anthers, while in others it
is the connective alone which is petal-like. Where the flower naturally
contains a large number of stamens, as in Mallows, Roses, Magnolias, &c.,
petaloid expansion of the filament is most common, though it is by no
means confined to such flowers, the change occurring in Allamanda
cathartica, Jasminum grandiflorum, and many other flowers with few
stamens. A similar change in the anther and connective takes place more
frequently in flowers where the number of stamens is smaller, but there are
of course numerous exceptions to this rule.
In those cases where there is more than one row of stamens, the outermost
are most liable to this change: thus in Saxifraga decipiens, as shown by Ch.
Morren,[304] the outer series of stamens—those opposite to the sepals—
become first affected, and, at a more advanced stage, the inner row also;
and this is the case in most flowers that have their stamens in two rows.
Occasionally it happens that an outer series of stamens is abortive, or
wholly suppressed, while the inner row becomes petalodic; this was the
case in some flowers of Lilium auratum lately exhibited by Messrs. Veitch.
Those flowers in which only a portion of the stamens undergo this change
are called semi-double, while in other cases that will be hereafter
mentioned, not only are the stamens thus rendered petaloid, but their
spur.
De Candolle[303] observes that in the Ranunculaceæ the species of Clematis
become double by the expansion of the filament, those of Ranunculus by
the dilatation of the anther, and those of Helleborus by the petal-like
development of both filament and anther. In some cases even on the same
plant all three modifications may be seen, as in Camellias, some of which
may be found with petaloid filaments with anthers on the top, others with
the filaments unchanged, but supporting petaloid anthers, while in others it
is the connective alone which is petal-like. Where the flower naturally
contains a large number of stamens, as in Mallows, Roses, Magnolias, &c.,
petaloid expansion of the filament is most common, though it is by no
means confined to such flowers, the change occurring in Allamanda
cathartica, Jasminum grandiflorum, and many other flowers with few
stamens. A similar change in the anther and connective takes place more
frequently in flowers where the number of stamens is smaller, but there are
of course numerous exceptions to this rule.
In those cases where there is more than one row of stamens, the outermost
are most liable to this change: thus in Saxifraga decipiens, as shown by Ch.
Morren,[304] the outer series of stamens—those opposite to the sepals—
become first affected, and, at a more advanced stage, the inner row also;
and this is the case in most flowers that have their stamens in two rows.
Occasionally it happens that an outer series of stamens is abortive, or
wholly suppressed, while the inner row becomes petalodic; this was the
case in some flowers of Lilium auratum lately exhibited by Messrs. Veitch.
Those flowers in which only a portion of the stamens undergo this change
are called semi-double, while in other cases that will be hereafter
mentioned, not only are the stamens thus rendered petaloid, but their
Page 371
number is also augmented, as in most double roses, pinks, anemones,
poppies, &c.
In some double flowers, in which the stamens assume more or less
completely the appearance of petals, a singular appearance is afforded by
the presence of four wing-like processes emanating from the central
filaments, two on each side, so that the arrangement may be compared to
two sheets of paper folded in the centre and adherent in that situation,
though perfectly separate elsewhere, except sometimes at the top, where
they form a sort of hood. This change results from an imperfect petalody of
the anther; the two wings on each side of the central vascular cord represent
the front and back walls of an anther lobe, or rather of that portion of the
anther which, under ordinary circumstances, produces pollen. In the
malformed flowers no pollen is formed, at least in the more complete states
of the malformation, but the walls of the anther lobe become preternaturally
enlarged, and petaloid in texture and appearance. This change occurs in
some semi-double rhododendrons and azaleas, in crocuses, and in a species
of violet found at Mentone by Mr. J. T. Moggridge.
There are numerous intermediate forms wherein the wing-like processes
may be traced all the way along the filament till they ultimately lose
themselves in the anther-lobes, with which they become continuous. In
some cases, as in Crocus and Rhododendron, this is shown even more
clearly by the existence of two perfect pollen-sacs or quarter-anthers, the
remaining portions being petaloid and continuous with the dilated filament.
Not unfrequently these semi-petaloid stamens adhere to the fronts of the
petals, and then it appears, at a first glance, as if three organs were stuck
together, one in front of another, while in reality there are but two.[305] (See
antè, p. 35, fig. 12.)
poppies, &c.
In some double flowers, in which the stamens assume more or less
completely the appearance of petals, a singular appearance is afforded by
the presence of four wing-like processes emanating from the central
filaments, two on each side, so that the arrangement may be compared to
two sheets of paper folded in the centre and adherent in that situation,
though perfectly separate elsewhere, except sometimes at the top, where
they form a sort of hood. This change results from an imperfect petalody of
the anther; the two wings on each side of the central vascular cord represent
the front and back walls of an anther lobe, or rather of that portion of the
anther which, under ordinary circumstances, produces pollen. In the
malformed flowers no pollen is formed, at least in the more complete states
of the malformation, but the walls of the anther lobe become preternaturally
enlarged, and petaloid in texture and appearance. This change occurs in
some semi-double rhododendrons and azaleas, in crocuses, and in a species
of violet found at Mentone by Mr. J. T. Moggridge.
There are numerous intermediate forms wherein the wing-like processes
may be traced all the way along the filament till they ultimately lose
themselves in the anther-lobes, with which they become continuous. In
some cases, as in Crocus and Rhododendron, this is shown even more
clearly by the existence of two perfect pollen-sacs or quarter-anthers, the
remaining portions being petaloid and continuous with the dilated filament.
Not unfrequently these semi-petaloid stamens adhere to the fronts of the
petals, and then it appears, at a first glance, as if three organs were stuck
together, one in front of another, while in reality there are but two.[305] (See
antè, p. 35, fig. 12.)
Page 372
Fig. 155.—Four-winged filaments of Rhododendron.
The change in the anther, above alluded to, must not be mistaken for that far
more common one in which only a small portion of the anther becomes
petaloid, forming a sort of lateral wing or appendage to the polliniferous
portion, as happens normally in Pterandra, and is common in some double
fuchsias. In this latter instance there is but a single wing, and the nature of
the case is obvious.
Double flowers of Orchidaceæ generally arise from petalification of the
filaments, with or without other coincident changes. What makes double
flowers in this order the more interesting is the development, in a petaloid
condition, of some or all of those stamens which under ordinary
circumstances are wholly suppressed, so that the morphological structure of
the flower, at first a matter of theory, becomes actually realised. Fig. 156 is
a diagram showing the presence of two additional labella within the
ordinary one in a species of Catasetum, and representing two petaloid
stamens, thus evidently completing the outer staminal whorl, of which there
is usually but a single representative (see Peloria, Multiplication,
Prolification). In some of these double orchids it is, however, necessary not
to confound a petaloid condition of the existing column with the
The change in the anther, above alluded to, must not be mistaken for that far
more common one in which only a small portion of the anther becomes
petaloid, forming a sort of lateral wing or appendage to the polliniferous
portion, as happens normally in Pterandra, and is common in some double
fuchsias. In this latter instance there is but a single wing, and the nature of
the case is obvious.
Double flowers of Orchidaceæ generally arise from petalification of the
filaments, with or without other coincident changes. What makes double
flowers in this order the more interesting is the development, in a petaloid
condition, of some or all of those stamens which under ordinary
circumstances are wholly suppressed, so that the morphological structure of
the flower, at first a matter of theory, becomes actually realised. Fig. 156 is
a diagram showing the presence of two additional labella within the
ordinary one in a species of Catasetum, and representing two petaloid
stamens, thus evidently completing the outer staminal whorl, of which there
is usually but a single representative (see Peloria, Multiplication,
Prolification). In some of these double orchids it is, however, necessary not
to confound a petaloid condition of the existing column with the
Page 373
development of usually suppressed stamens in a petaloid form. Thus, in
Lycaste Skinneri the column is frequently provided with two petal-like
wings, which might readily be supposed to be two stamens of the inner
whorl adherent to the column; a little attention, however, to the relative
position of these adventitious wings is generally sufficient to enable the
observer to ascertain the true nature of the appearance.[306]
Fig. 156.—Diagram of flower of Catasetum, with two labella.
Some forms of duplicate or hose in hose corollas are apparently due, not so
much to the formation of a second corolla within the first, as to the presence
of an inner series of petal-like stamens, which, by their cohesion, form a
second pseudo-corolla within the first. The staminal nature of this pseudo-
corolla is inferred from the occasional presence of anthers on it.[307] In
Datura fastuosa, as well as in Gloxinia, a pseudo-corolla of this kind
sometimes occurs with the addition of a series of petaloid stamens attached
to its outer surface.[308]
Lycaste Skinneri the column is frequently provided with two petal-like
wings, which might readily be supposed to be two stamens of the inner
whorl adherent to the column; a little attention, however, to the relative
position of these adventitious wings is generally sufficient to enable the
observer to ascertain the true nature of the appearance.[306]
Fig. 156.—Diagram of flower of Catasetum, with two labella.
Some forms of duplicate or hose in hose corollas are apparently due, not so
much to the formation of a second corolla within the first, as to the presence
of an inner series of petal-like stamens, which, by their cohesion, form a
second pseudo-corolla within the first. The staminal nature of this pseudo-
corolla is inferred from the occasional presence of anthers on it.[307] In
Datura fastuosa, as well as in Gloxinia, a pseudo-corolla of this kind
sometimes occurs with the addition of a series of petaloid stamens attached
to its outer surface.[308]
Page 374
When the petalody specially affects the anther-lobes, as in Arbutus,
Petunia, Fuchsia, &c., the venation of the petal-like portion is very
frequently laminar, thus tending to show that the anther is in such cases
really a modification of the blade of the leaf; but as, on the other hand, we
often find petal-like filaments bearing pollen-sacs on their sides, it is clear
that we must not attribute the formation of pollen to the blade of the leaf
only, but we must admit that it may be formed in the filament as well.[309]
Fig. 158.—Portion of a double columbine (Aquilegia), showing petalody of the connective.
Petunia, Fuchsia, &c., the venation of the petal-like portion is very
frequently laminar, thus tending to show that the anther is in such cases
really a modification of the blade of the leaf; but as, on the other hand, we
often find petal-like filaments bearing pollen-sacs on their sides, it is clear
that we must not attribute the formation of pollen to the blade of the leaf
only, but we must admit that it may be formed in the filament as well.[309]
Fig. 158.—Portion of a double columbine (Aquilegia), showing petalody of the connective.
Page 375
Fig. 159.—Petaloid stamens, Hibiscus.
Petalody of the connective is of less frequent occurrence than the
corresponding change in the other portions of the stamen. It may be seen in
some forms of double columbine,[310] in which the connective forms a
tubular petal or nectary, and in double petunias and fuchsias. When it
occurs, the true anther-lobes are usually atrophied, and little or no pollen is
formed.
An occurrence of this nature in Tacsonia pinnatistipula, in conjunction with
the partial detachment of the stamens from the gynophore, led Karsten to
establish a genus which he called Poggendorffia.[311]
From the subjoined list of genera in which petalody of the stamens, in some
form or other, has been observed, it will be seen that it happens more often
in plants with numerous distinct organs (Polypetalæ, Polyandria, Polygynia,
&c.) than in other plants with a smaller number of parts, and which are
Petalody of the connective is of less frequent occurrence than the
corresponding change in the other portions of the stamen. It may be seen in
some forms of double columbine,[310] in which the connective forms a
tubular petal or nectary, and in double petunias and fuchsias. When it
occurs, the true anther-lobes are usually atrophied, and little or no pollen is
formed.
An occurrence of this nature in Tacsonia pinnatistipula, in conjunction with
the partial detachment of the stamens from the gynophore, led Karsten to
establish a genus which he called Poggendorffia.[311]
From the subjoined list of genera in which petalody of the stamens, in some
form or other, has been observed, it will be seen that it happens more often
in plants with numerous distinct organs (Polypetalæ, Polyandria, Polygynia,
&c.) than in other plants with a smaller number of parts, and which are
Page 376
more or less adherent one to the other. The tendency to petalification is,
moreover, greater among those plants which have their floral elements
arranged in spiral series, than among those where the verticillate
arrangement exists; and in any given flower, if the stamens are spirally
arranged while the carpels are grouped in whorls, the former will be more
liable to petalody than the latter, and vice versâ. It has been before
remarked, that this condition is far more common in plants whose petals,
&c., have straight veins, like those in the sheath of a leaf, than in those the
venation of which is reticulate, as in the blade of the leaf. It must also be
remembered that in the same genus, even in the same species, different
kinds of doubling occur. Familiar illustrations of this are afforded in the
case of anemones, columbines, fuchsias, and other plants.
The existence of "compound stamens" in some flowers, as pointed out by
Payer, and others, and the researches of Dr. Alexander Dickson, confer
additional importance on the subject of petalody, and necessitate the
examination of double flowers with special reference to these compound
stamens, and to the order of their development.[312] The presence of these
compound stamens affords a satisfactory explanation of the appearance in
some double Malvaceæ, wherein the tufts of adventitious petals are very
liable to be mistaken for buds, produced by axillary prolification in the axils
of the petals, but which are in reality compound and petaloid stamens. At
other times, however, true axillary prolification exists in these flowers; but
then the supplemental florets have always a calyx, which is wanting in the
other instances.
Petalody of the stamens has been met with most frequently in the following
genera:
*Ranunculus!
*Anemone!
*Papaver!
moreover, greater among those plants which have their floral elements
arranged in spiral series, than among those where the verticillate
arrangement exists; and in any given flower, if the stamens are spirally
arranged while the carpels are grouped in whorls, the former will be more
liable to petalody than the latter, and vice versâ. It has been before
remarked, that this condition is far more common in plants whose petals,
&c., have straight veins, like those in the sheath of a leaf, than in those the
venation of which is reticulate, as in the blade of the leaf. It must also be
remembered that in the same genus, even in the same species, different
kinds of doubling occur. Familiar illustrations of this are afforded in the
case of anemones, columbines, fuchsias, and other plants.
The existence of "compound stamens" in some flowers, as pointed out by
Payer, and others, and the researches of Dr. Alexander Dickson, confer
additional importance on the subject of petalody, and necessitate the
examination of double flowers with special reference to these compound
stamens, and to the order of their development.[312] The presence of these
compound stamens affords a satisfactory explanation of the appearance in
some double Malvaceæ, wherein the tufts of adventitious petals are very
liable to be mistaken for buds, produced by axillary prolification in the axils
of the petals, but which are in reality compound and petaloid stamens. At
other times, however, true axillary prolification exists in these flowers; but
then the supplemental florets have always a calyx, which is wanting in the
other instances.
Petalody of the stamens has been met with most frequently in the following
genera:
*Ranunculus!
*Anemone!
*Papaver!
Page 377
*Clematis!
*Hepatica!
*Ficaria!
Thalictrum.
*Caltha!
*Trollius!
*Nigella!
*Aquilegia!
*Delphinium!
*Adonis!
*Pæonia!
*Nelumbium!
*Nymphæa!
*Berberis!
*Papaver!
*Chelidonium!
Sanguinaria.
Podophyllum.
*Mathiola!
*Cheiranthus!
*Iberis!
*Cardamine!
*Hesperis.
*Barbarea!
*Sinapis!
*Brassica!
*Helianthemum!
*Viola!
*Dianthus!
*Saponaria!
*Lychnis!
*Silene!
*Sagina!
*Hibiscus!
*Althæa!
*Malva!
Æsculus!
*Geranium!
*Pelargonium.
*Hepatica!
*Ficaria!
Thalictrum.
*Caltha!
*Trollius!
*Nigella!
*Aquilegia!
*Delphinium!
*Adonis!
*Pæonia!
*Nelumbium!
*Nymphæa!
*Berberis!
*Papaver!
*Chelidonium!
Sanguinaria.
Podophyllum.
*Mathiola!
*Cheiranthus!
*Iberis!
*Cardamine!
*Hesperis.
*Barbarea!
*Sinapis!
*Brassica!
*Helianthemum!
*Viola!
*Dianthus!
*Saponaria!
*Lychnis!
*Silene!
*Sagina!
*Hibiscus!
*Althæa!
*Malva!
Æsculus!
*Geranium!
*Pelargonium.
Page 378
*Tropæolum!
Oxalis!
*Impatiens!
*Camellia!
Thea!
Trifolium!
Medicago!
*Ulex!
Spartianthus.
Clitoria.
Pisum!
Orobus!
Genista!
Spartium!
Cytisus!
Anthyllis.
Coronilla.
Lotus!
*Rosa!
*Kerria!
*Spiræa!
*Fragaria!
*Potentilla!
*Cratægus!
Cydonia.
*Pyrus!
Eriobotrya!
*Amygdalus!
*Prunus!
*Myrtus!
*Punica!
*Philadelphus!
*Deutzia!
*Fuchsia!
Godetia!
Clarkia!
Portulaca!
Ribes!
Saxifraga!
Oxalis!
*Impatiens!
*Camellia!
Thea!
Trifolium!
Medicago!
*Ulex!
Spartianthus.
Clitoria.
Pisum!
Orobus!
Genista!
Spartium!
Cytisus!
Anthyllis.
Coronilla.
Lotus!
*Rosa!
*Kerria!
*Spiræa!
*Fragaria!
*Potentilla!
*Cratægus!
Cydonia.
*Pyrus!
Eriobotrya!
*Amygdalus!
*Prunus!
*Myrtus!
*Punica!
*Philadelphus!
*Deutzia!
*Fuchsia!
Godetia!
Clarkia!
Portulaca!
Ribes!
Saxifraga!
Page 379
Daucus.
Ixora.
Serissa!
Gardenia!
Lonicera!
Sambucus.
Viburnum.
Scabiosa.
*Campanula!
Platycodon!
Calluna!
Azalea!
Rhododendron!
*Arbutus!
*Erica!
*Anagallis!
*Primula!
*Jasminum!
Syringa!
*Vinca!
*Nerium!
Allamanda!
Tabernæmontana.
*Calystegia!
Convolvulus!
Ipomœa.
*Datura!
*Petunia!
Solanum!
Orobanche.
Gentiana.
Mimulus.
*Antirrhinum!
Gratiola!
*Digitalis!
*Linaria!
Veronica!
Calceolaria!
Achimenes.
Ixora.
Serissa!
Gardenia!
Lonicera!
Sambucus.
Viburnum.
Scabiosa.
*Campanula!
Platycodon!
Calluna!
Azalea!
Rhododendron!
*Arbutus!
*Erica!
*Anagallis!
*Primula!
*Jasminum!
Syringa!
*Vinca!
*Nerium!
Allamanda!
Tabernæmontana.
*Calystegia!
Convolvulus!
Ipomœa.
*Datura!
*Petunia!
Solanum!
Orobanche.
Gentiana.
Mimulus.
*Antirrhinum!
Gratiola!
*Digitalis!
*Linaria!
Veronica!
Calceolaria!
Achimenes.
Page 380
Gloxinia!
Clerodendron!
Bignonia.
Cyclamen!
Mirabilis.
Laurus!
Gladiolus!
Crocus!
Iris!
*Galanthus!
Leucojum!
Sternbergia!
Hippeastrum.
*Narcissus!
*Orchis!
Catasetum!
Hydrocharis.
Asphodelus.
*Tulipa!
Scilla.
*Convallaria!
Fritillaria!
*Lilium!
*Hyacinthus!
*Polianthes!
*Hemerocallis!
*Colchicum!
*Sagittaria!
*Tradescantia!
Commelyna!
Tofieldia.
Petalody of the pistils.—Taken by itself, this is much less common than
the corresponding change in the stamens. It generally affects the style and
stigma only, as happens normally in Petalostylis, Iris, &c., but this is by no
means always necessarily the case. In some of the cultivated varieties of
Anemone and Ranunculus all the parts of the flower remain in their normal
state, except the pistils, which latter assume a petaloid appearance.
Clerodendron!
Bignonia.
Cyclamen!
Mirabilis.
Laurus!
Gladiolus!
Crocus!
Iris!
*Galanthus!
Leucojum!
Sternbergia!
Hippeastrum.
*Narcissus!
*Orchis!
Catasetum!
Hydrocharis.
Asphodelus.
*Tulipa!
Scilla.
*Convallaria!
Fritillaria!
*Lilium!
*Hyacinthus!
*Polianthes!
*Hemerocallis!
*Colchicum!
*Sagittaria!
*Tradescantia!
Commelyna!
Tofieldia.
Petalody of the pistils.—Taken by itself, this is much less common than
the corresponding change in the stamens. It generally affects the style and
stigma only, as happens normally in Petalostylis, Iris, &c., but this is by no
means always necessarily the case. In some of the cultivated varieties of
Anemone and Ranunculus all the parts of the flower remain in their normal
state, except the pistils, which latter assume a petaloid appearance.
Page 381
Many of the double flowers owe their peculiar appearance to the
combination of the following appearances—a petal-like form of the
stamens, increase in the number of these organs and similar changes
affecting the pistils, and is applied to several distinct conditions. If in any
given flower all the stamens and all the pistils become wholly petaloid, no
pollen is formed, and of course no seeds can be produced, but this very
rarely happens, as usually some pollen is produced, and some ovules
capable of being fertilised are developed.
In double flowers of Primula sinensis it frequently happens that the capsule
is either partially leafy or partly petal-like; in either case the fruit is open at
the extremity, and often destitute of the style and stigma. It is, however,
doubtful if the ovules can be fertilised in these flowers.
The following list comprises the names of those genera in which this
change has been most frequently observed, independently of corresponding
alterations in the stamens, but it is more usual for both sets of organs to be
similarly affected.
*Ranunculus!
*Anemone!
Nigella.
*Papaver!
*Dianthus!
Saponaria!
Viola!
Camellia!
Alcea.
Hibiscus!
Amygdalus!
Lonicera!
Scabiosa.
Æschynanthus!
Primula!
combination of the following appearances—a petal-like form of the
stamens, increase in the number of these organs and similar changes
affecting the pistils, and is applied to several distinct conditions. If in any
given flower all the stamens and all the pistils become wholly petaloid, no
pollen is formed, and of course no seeds can be produced, but this very
rarely happens, as usually some pollen is produced, and some ovules
capable of being fertilised are developed.
In double flowers of Primula sinensis it frequently happens that the capsule
is either partially leafy or partly petal-like; in either case the fruit is open at
the extremity, and often destitute of the style and stigma. It is, however,
doubtful if the ovules can be fertilised in these flowers.
The following list comprises the names of those genera in which this
change has been most frequently observed, independently of corresponding
alterations in the stamens, but it is more usual for both sets of organs to be
similarly affected.
*Ranunculus!
*Anemone!
Nigella.
*Papaver!
*Dianthus!
Saponaria!
Viola!
Camellia!
Alcea.
Hibiscus!
Amygdalus!
Lonicera!
Scabiosa.
Æschynanthus!
Primula!
Page 382
Petalody of the ovules.—The principal changes which occur in the ovule
have already been alluded to at pp. 262–272; it may here be stated,
however, that the ovules are occasionally represented by small stalked
petal-like structures. This happens with especial frequency among
Cruciferæ.[313]
Petalody of the accessory organs.—A petaloid condition of the disc, of the
scales, or other excrescences from the axis or from the lateral portions of
the flower, is of frequent occurrence, though it is but rarely that the change
is of any great importance in a morphological point of view. C. Morren has
given the name adenopetaly to a case wherein one of the glands at the base
of the petals in Lopezia was replaced by a petal.[314] A similar change may
be seen in the double Oleander.
Staminody of the bracts.—An instance of this has been already alluded to
in Abies excelsa, as observed by Prof. Dickson, and in which some of the
bracts were seen assuming the form and characteristic of the stamens see
ante: p. 192. Signor Licopoli met with a similar substitution of anthers for
bracts in Melianthus major.[315]
Staminody of the sepals and petals.—In the first named this is of very
rare occurrence. M. Gris has recorded an instance in Philadelphus
speciosus[316] which appears to be the only case on record. The
corresponding change in the case of the petals is far more common. De
Candolle cites in illustration of this occurrence flowers of the common
haricot, in which the alæ and carina of the corolla were thus changed.[317]
There is in cultivation a form of Saxifraga granulata wherein the petals are
replaced by stamens, so that there are fifteen stamens. A similar change has
been observed in Capsella bursa-pastoris.
have already been alluded to at pp. 262–272; it may here be stated,
however, that the ovules are occasionally represented by small stalked
petal-like structures. This happens with especial frequency among
Cruciferæ.[313]
Petalody of the accessory organs.—A petaloid condition of the disc, of the
scales, or other excrescences from the axis or from the lateral portions of
the flower, is of frequent occurrence, though it is but rarely that the change
is of any great importance in a morphological point of view. C. Morren has
given the name adenopetaly to a case wherein one of the glands at the base
of the petals in Lopezia was replaced by a petal.[314] A similar change may
be seen in the double Oleander.
Staminody of the bracts.—An instance of this has been already alluded to
in Abies excelsa, as observed by Prof. Dickson, and in which some of the
bracts were seen assuming the form and characteristic of the stamens see
ante: p. 192. Signor Licopoli met with a similar substitution of anthers for
bracts in Melianthus major.[315]
Staminody of the sepals and petals.—In the first named this is of very
rare occurrence. M. Gris has recorded an instance in Philadelphus
speciosus[316] which appears to be the only case on record. The
corresponding change in the case of the petals is far more common. De
Candolle cites in illustration of this occurrence flowers of the common
haricot, in which the alæ and carina of the corolla were thus changed.[317]
There is in cultivation a form of Saxifraga granulata wherein the petals are
replaced by stamens, so that there are fifteen stamens. A similar change has
been observed in Capsella bursa-pastoris.
Page 383
Cramer figures and describes a stamen occupying the place of a petal in
Daucus Carota.[318] Turpin[319] describes a similar occurrence in Monarda
fistulosa, in which the lower lip terminated in an anther, but this may have
been a case of adhesion. Moquin cites from Chamisso, Digitalis purpurea,
and from Jussieu, Asphodelus ramosus, as having presented this change,
and Wiegmann[320] has seen anthers developed on the awns of Avena
chinensis. In semi-double flowers of Ophrys aranifera and Orchis mascula,
the lateral petals are occasionally partially antheroid, and others occur in
which two of the outer series of stamens, which are ordinarily suppressed,
are present, but in a petaloid state. Reichenbach[321] figures an illustration
of this change, and also Moggridge.[322]
Staminody of the pistils.—The existence of this change has been denied
by several authors, nevertheless, it is of sufficiently common occurrence.
Alexander Braun notices the transformation of pistils into stamens in
Chives (Allium Scorodoprasum), and in which three stamens appeared in
the place of as many pistils, and had extrorse anthers, while the six normal
anthers are introrse. In the horse-radish (Armoracia rusticana), two of the
carpels are frequently converted into stamens, while two other organs
absent from the normal flower make their appearance as carpels. Roeper
has observed this phenomenon in Euphorbia palustris,[323] and in Gentiana
campestris.[324] In these examples one of the carpels was apparently absent,
and its place supplied by an anther. Roeper has also mentioned a balsam
with a supernumerary stamen occupying exactly the position of a carpel.
[325]
Agardh has observed a similar thing in a hyacinth, one half of the fruit of
which contained seeds, and the other half, anthers. B. Clarke mentions an
instance in Mathiola incana in which the carpels were disunited, and
antheriferous at the margin.[326]
Daucus Carota.[318] Turpin[319] describes a similar occurrence in Monarda
fistulosa, in which the lower lip terminated in an anther, but this may have
been a case of adhesion. Moquin cites from Chamisso, Digitalis purpurea,
and from Jussieu, Asphodelus ramosus, as having presented this change,
and Wiegmann[320] has seen anthers developed on the awns of Avena
chinensis. In semi-double flowers of Ophrys aranifera and Orchis mascula,
the lateral petals are occasionally partially antheroid, and others occur in
which two of the outer series of stamens, which are ordinarily suppressed,
are present, but in a petaloid state. Reichenbach[321] figures an illustration
of this change, and also Moggridge.[322]
Staminody of the pistils.—The existence of this change has been denied
by several authors, nevertheless, it is of sufficiently common occurrence.
Alexander Braun notices the transformation of pistils into stamens in
Chives (Allium Scorodoprasum), and in which three stamens appeared in
the place of as many pistils, and had extrorse anthers, while the six normal
anthers are introrse. In the horse-radish (Armoracia rusticana), two of the
carpels are frequently converted into stamens, while two other organs
absent from the normal flower make their appearance as carpels. Roeper
has observed this phenomenon in Euphorbia palustris,[323] and in Gentiana
campestris.[324] In these examples one of the carpels was apparently absent,
and its place supplied by an anther. Roeper has also mentioned a balsam
with a supernumerary stamen occupying exactly the position of a carpel.
[325]
Agardh has observed a similar thing in a hyacinth, one half of the fruit of
which contained seeds, and the other half, anthers. B. Clarke mentions an
instance in Mathiola incana in which the carpels were disunited, and
antheriferous at the margin.[326]
Page 384
The passage of pistils to stamens in willows has been frequently remarked,
as in Salix babylonica, silesiaca, cinerea, Caprea and nigricans. One of the
most curious illustrations of this transformation in this genus is given by
Henry and Macquart (Erst. Jahrb. des bot. Vereines am m. et n. Rhein.,
1837). In the flowers in question the series of changes were as follows:—
first, the ovary opened by a slit, and then expanded into a cup; next, anther-
cells were developed on the margin of the cup, with stigmas alternating
with them, the ovules at the same time disappearing; lastly, the margin
became divided, and bore three perfect anthers, which in the more perfect
states were raised on three filaments.
Campanula persicifolia, C. rapunculoides, and C. glomerata have been
observed to present an anther surmounting the pistil.[327] Double tulips
often present this change, and a like appearance has been observed in
Galanthus nivalis, and Narcissus Tazetta.
Moquin mentions the existence of this condition in a female plant of maize,
some of the pistils of which were wholly or partially converted into anther-
like organs. Mohl has recorded an analogous malformation in Chamærops
humilis, and in which the three carpels were normally formed, and only
differed from natural ovaries in this, that along the two edges of the ventral
suture there was a yellow thickening, which a cross section of the ovary
showed to be an anther-lobe filled with pollen.[328]
In Tofieldia calyculata a similar substitution of a stamen for a carpel has
been observed by Klotsch,[329] and Weber[330] gives other instances in
Prunus and Pæonia. Corresponding alterations may be met with in
cultivated tulips, in the cowslip and other plants. In most of the above cases
the transmutation has been perfect, but in quite an equal number of cases a
portion only of the carpel is thus changed, generally the style or the stigma;
thus Baillon describes the stigmas of Ricinus communis as having been in
as in Salix babylonica, silesiaca, cinerea, Caprea and nigricans. One of the
most curious illustrations of this transformation in this genus is given by
Henry and Macquart (Erst. Jahrb. des bot. Vereines am m. et n. Rhein.,
1837). In the flowers in question the series of changes were as follows:—
first, the ovary opened by a slit, and then expanded into a cup; next, anther-
cells were developed on the margin of the cup, with stigmas alternating
with them, the ovules at the same time disappearing; lastly, the margin
became divided, and bore three perfect anthers, which in the more perfect
states were raised on three filaments.
Campanula persicifolia, C. rapunculoides, and C. glomerata have been
observed to present an anther surmounting the pistil.[327] Double tulips
often present this change, and a like appearance has been observed in
Galanthus nivalis, and Narcissus Tazetta.
Moquin mentions the existence of this condition in a female plant of maize,
some of the pistils of which were wholly or partially converted into anther-
like organs. Mohl has recorded an analogous malformation in Chamærops
humilis, and in which the three carpels were normally formed, and only
differed from natural ovaries in this, that along the two edges of the ventral
suture there was a yellow thickening, which a cross section of the ovary
showed to be an anther-lobe filled with pollen.[328]
In Tofieldia calyculata a similar substitution of a stamen for a carpel has
been observed by Klotsch,[329] and Weber[330] gives other instances in
Prunus and Pæonia. Corresponding alterations may be met with in
cultivated tulips, in the cowslip and other plants. In most of the above cases
the transmutation has been perfect, but in quite an equal number of cases a
portion only of the carpel is thus changed, generally the style or the stigma;
thus Baillon describes the stigmas of Ricinus communis as having been in
Page 385
one instance antheriferous.[331] Moggridge figures a flower of Ophrys
insectifera in which the rostellate process was replaced by an anther.[332]
Mohl remarks that the change of pistils into stamens is more common in
monocarpellary pistils than it is in those which are made up of several
carpels. It seems clear that in this transformation the lobes of the anther and
the development of pollen have no relation to the production of ovules.
Staminody of the accessory organs of the flower.—The scales that are
met with in some plants, either as excrescences from the petals, or as
imperfect representatives of stamens or other organs, are occasionally
staminoid; thus the scales of Saponaria officinalis, of Silene, Nerium
Oleander, the rays of Passiflora, the corona of Narcissus, have all been
observed occasionally to bear anthers.[333] In the case of Narcissus the loose
spongy tissue of the corona seems to have the nearest analogy to the anther-
lobes, while the prolonged connective is more like the ordinary segments of
the perianth in texture. The species in which this change may most
frequently be observed are, N. poeticus, N. incomparabilis, and N.
montanus.
M. Bureau found in some flowers of Antirrhinum majus two petal-like
bodies standing up in front of, or opposite to the two petals of the upper lip,
[334] and similar developments in which each of the two adventitious
segments are surmounted by an anther may be met with frequently. It does
not follow because these organs bear anthers that they are morphologically
true stamens. They are really scales, &c., taking on themselves accidentally
the characters proper to stamens.
Pistillody of the perianth.—The passage of the segments of the perianth
into carpels has been observed frequently in Tulipa Gesneriana, the change
in question being generally attended by a partial virescence. M. Gay is said
insectifera in which the rostellate process was replaced by an anther.[332]
Mohl remarks that the change of pistils into stamens is more common in
monocarpellary pistils than it is in those which are made up of several
carpels. It seems clear that in this transformation the lobes of the anther and
the development of pollen have no relation to the production of ovules.
Staminody of the accessory organs of the flower.—The scales that are
met with in some plants, either as excrescences from the petals, or as
imperfect representatives of stamens or other organs, are occasionally
staminoid; thus the scales of Saponaria officinalis, of Silene, Nerium
Oleander, the rays of Passiflora, the corona of Narcissus, have all been
observed occasionally to bear anthers.[333] In the case of Narcissus the loose
spongy tissue of the corona seems to have the nearest analogy to the anther-
lobes, while the prolonged connective is more like the ordinary segments of
the perianth in texture. The species in which this change may most
frequently be observed are, N. poeticus, N. incomparabilis, and N.
montanus.
M. Bureau found in some flowers of Antirrhinum majus two petal-like
bodies standing up in front of, or opposite to the two petals of the upper lip,
[334] and similar developments in which each of the two adventitious
segments are surmounted by an anther may be met with frequently. It does
not follow because these organs bear anthers that they are morphologically
true stamens. They are really scales, &c., taking on themselves accidentally
the characters proper to stamens.
Pistillody of the perianth.—The passage of the segments of the perianth
into carpels has been observed frequently in Tulipa Gesneriana, the change
in question being generally attended by a partial virescence. M. Gay is said
Page 386
by Moquin to have observed a flower of Crocus nudiflorus in which the
segments of the perianth were cleft and fringed at the same time, so that
they presented the appearance of the stigmas.
Fig. 160.—Flower of tulip, allowing vertical attachment of a leaf, and also the existence of ovules on
the margins of the segments of the perianth. Some of the parts are removed.
Pistillody of the sepals.—In some double flowers of the garden pea
communicated by Mr. Laxton, among other peculiarities was a
supernumerary 5–6-leaved calyx, some of the segments of which were of a
carpellary nature, and bore imperfect ovules on their margins, while at their
extremities they were drawn out into styles.[335]
Pistillody of the stamens.—This change whereby the stamens assume
more or less the appearance of pistils is more commonly met with than is
the metamorphosis of the envelopes of the flower into carpels. In some
cases the whole of the stamen appears to be changed, while in others it is
segments of the perianth were cleft and fringed at the same time, so that
they presented the appearance of the stigmas.
Fig. 160.—Flower of tulip, allowing vertical attachment of a leaf, and also the existence of ovules on
the margins of the segments of the perianth. Some of the parts are removed.
Pistillody of the sepals.—In some double flowers of the garden pea
communicated by Mr. Laxton, among other peculiarities was a
supernumerary 5–6-leaved calyx, some of the segments of which were of a
carpellary nature, and bore imperfect ovules on their margins, while at their
extremities they were drawn out into styles.[335]
Pistillody of the stamens.—This change whereby the stamens assume
more or less the appearance of pistils is more commonly met with than is
the metamorphosis of the envelopes of the flower into carpels. In some
cases the whole of the stamen appears to be changed, while in others it is
Page 387
the filament alone that is altered, the anther being deficient, or rudimentary;
while, in a third class of cases, the filament is unaffected, and the anther
undergoes the change in question. In those instances in which the filament
appears to be the portion most implicated, it becomes dilated so as to
resemble a leaf-sheath rather than a leaf-stalk, as it does usually.
One of the most curious cases of this kind is that recorded in the 'Botanical
Magazine,' (tab. 5160, f. 4) as having occurred in Begonia frigida already
alluded to, and in which, in the centre of a male flower, were four free
ovoid ovaries alternating with as many stamens. In the normal flowers of
this plant, as is well known, the male flowers have several stamens, while in
the female flowers the ovary is strictly inferior, so that, in the singular
flower just described, the perianth was inferior instead of being superior, as
it is usually. It should be added also that the perianth in these malformed
flowers was precisely like that which occurs ordinarily in the male flowers.
Fig. 161.—Supernumerary carpels in the orange, arising from substitution of pistils for stamens.
while, in a third class of cases, the filament is unaffected, and the anther
undergoes the change in question. In those instances in which the filament
appears to be the portion most implicated, it becomes dilated so as to
resemble a leaf-sheath rather than a leaf-stalk, as it does usually.
One of the most curious cases of this kind is that recorded in the 'Botanical
Magazine,' (tab. 5160, f. 4) as having occurred in Begonia frigida already
alluded to, and in which, in the centre of a male flower, were four free
ovoid ovaries alternating with as many stamens. In the normal flowers of
this plant, as is well known, the male flowers have several stamens, while in
the female flowers the ovary is strictly inferior, so that, in the singular
flower just described, the perianth was inferior instead of being superior, as
it is usually. It should be added also that the perianth in these malformed
flowers was precisely like that which occurs ordinarily in the male flowers.
Fig. 161.—Supernumerary carpels in the orange, arising from substitution of pistils for stamens.
Page 388
In some varieties of the orange, called by the French "bigarades cornues,"
the thalamus of the flower, which is usually short, and terminated by a
glandular ring-like disc, is prolonged into a little stalk or gynophore,
bearing a ring of supernumerary carpels. These carpels are isolated one
from another, and are formed by the transformation of the filaments of the
stamens.[336]
The additional carpels in the case of the apple of St. Valéry, in which the
petals are of a green colour, like the sepals, are by some attributed to the
transformation of the stamens into carpels. These adventitious carpels
frequently contain imperfect ovules and form a whorl above the normal
ones. (See Pyrus dioica of Willdenow.)[337] A similar change occasionally
happens in the stamens of Magnolia fuscata, while in double tulips this
phenomenon is very frequent, and among them may be found all stages of
transition between stamens and pistils, and many of the parts combining the
characters of both.[338] Dunal and Campdera have described flowers of
Rumex crispus, with seven pistils, occupying the place of as many stamens.
the thalamus of the flower, which is usually short, and terminated by a
glandular ring-like disc, is prolonged into a little stalk or gynophore,
bearing a ring of supernumerary carpels. These carpels are isolated one
from another, and are formed by the transformation of the filaments of the
stamens.[336]
The additional carpels in the case of the apple of St. Valéry, in which the
petals are of a green colour, like the sepals, are by some attributed to the
transformation of the stamens into carpels. These adventitious carpels
frequently contain imperfect ovules and form a whorl above the normal
ones. (See Pyrus dioica of Willdenow.)[337] A similar change occasionally
happens in the stamens of Magnolia fuscata, while in double tulips this
phenomenon is very frequent, and among them may be found all stages of
transition between stamens and pistils, and many of the parts combining the
characters of both.[338] Dunal and Campdera have described flowers of
Rumex crispus, with seven pistils, occupying the place of as many stamens.
Page 389
Fig. 162.—Substitution of carpels for stamens in Papaver.
In Papaver bracteatum a considerable number of the stamens sometimes
become developed into pistils, especially those which are nearest to the
centre of the flower, and in these flowers the filaments are said to become
the ovaries, while the anthers are curled so as to resemble stigmas. A similar
change is not infrequent Papaver somniferum. Goeppert, who found
numerous instances of the kind in a field near Breslau, says the peculiarity
was reproduced by seed for two years in succession.[339] Wigand ('Flora,'
1856, p. 717) has noticed among other changes the pistil of Gentiana
Amarella bearing two sessile anthers. Polemonium cæruleum is another
plant very subject to this change. Brongniart[340] describes a flower of this
species in which the stamens were represented by a circle of carpels united
to each other so as to form a sheath around the central ovary. By artificial
fertilization M. Brongniart obtained fertile seeds from the central normal
ovary as well as from the surrounding metamorphosed stamens.
In Papaver bracteatum a considerable number of the stamens sometimes
become developed into pistils, especially those which are nearest to the
centre of the flower, and in these flowers the filaments are said to become
the ovaries, while the anthers are curled so as to resemble stigmas. A similar
change is not infrequent Papaver somniferum. Goeppert, who found
numerous instances of the kind in a field near Breslau, says the peculiarity
was reproduced by seed for two years in succession.[339] Wigand ('Flora,'
1856, p. 717) has noticed among other changes the pistil of Gentiana
Amarella bearing two sessile anthers. Polemonium cæruleum is another
plant very subject to this change. Brongniart[340] describes a flower of this
species in which the stamens were represented by a circle of carpels united
to each other so as to form a sheath around the central ovary. By artificial
fertilization M. Brongniart obtained fertile seeds from the central normal
ovary as well as from the surrounding metamorphosed stamens.
Page 390
Cheiranthus Cheiri has long been known as one of the plants most subject
to this anomaly. De Candolle even mentions it in his 'Prodromus' as a
distinct variety, under the name of gynantherus. Brongniart (loc. cit.) thus
refers to the Cheiranthus:—"Sometimes these six carpellary leaves are
perfectly free, and in this case they spread open, presenting two rows of
ovules along their inner edges, or these edges maybe soldered together,
forming a kind of follicle like that of the columbine; at other times, these
staminal pistils are fused into two lateral bundles of three in each bundle, or
into a single cylinder which encircles the true pistil. In a third set of cases
these outer carpels are only four in number, two lateral and two antero-
posterior, all fused in such a manner as to form around the normal pistil a
prism-shaped sheath, with four sides presenting four parietal placentæ,
corresponding to the lines of junction of the staminal carpels."
In the accompanying figures (fig. 163, a-d) the nature of this change is
illustrated. In some of the specimens it is easy to see that the two shorter
stamens undergo the change into carpels later and less perfectly than the
four longer ones, and not infrequently the outer pair are altogether absent.
In most of the flowers of this variety the petals are smaller and less
perfectly developed than usual.[341]
to this anomaly. De Candolle even mentions it in his 'Prodromus' as a
distinct variety, under the name of gynantherus. Brongniart (loc. cit.) thus
refers to the Cheiranthus:—"Sometimes these six carpellary leaves are
perfectly free, and in this case they spread open, presenting two rows of
ovules along their inner edges, or these edges maybe soldered together,
forming a kind of follicle like that of the columbine; at other times, these
staminal pistils are fused into two lateral bundles of three in each bundle, or
into a single cylinder which encircles the true pistil. In a third set of cases
these outer carpels are only four in number, two lateral and two antero-
posterior, all fused in such a manner as to form around the normal pistil a
prism-shaped sheath, with four sides presenting four parietal placentæ,
corresponding to the lines of junction of the staminal carpels."
In the accompanying figures (fig. 163, a-d) the nature of this change is
illustrated. In some of the specimens it is easy to see that the two shorter
stamens undergo the change into carpels later and less perfectly than the
four longer ones, and not infrequently the outer pair are altogether absent.
In most of the flowers of this variety the petals are smaller and less
perfectly developed than usual.[341]
Page 391
Fig. 163.—Cheiranthus Cheiri, var. gynantherus. a. Sepals and petals removed to show carpellodic
stamens. b. The same laid open. c. Transverse section. d. Plan of flower with four carpel-like
stamens, &c.
In Lilium tigrinum, some specimens of which were gathered by Mr. J.
Salter, in addition to various degrees of synanthy and other changes, some
of the stamens were developed in the form of carpels, adherent by their
edges so as to form an imperfect tube or sheath around the normal pistil.
Fig. 164 shows one of the intermediate organs from these flowers, in which
half the structure seems devoted to the formation of ovules, while the other
half bears a one-celled anther. Lindley[342] has also described a case of this
kind in a species of Amaryllis.
stamens. b. The same laid open. c. Transverse section. d. Plan of flower with four carpel-like
stamens, &c.
In Lilium tigrinum, some specimens of which were gathered by Mr. J.
Salter, in addition to various degrees of synanthy and other changes, some
of the stamens were developed in the form of carpels, adherent by their
edges so as to form an imperfect tube or sheath around the normal pistil.
Fig. 164 shows one of the intermediate organs from these flowers, in which
half the structure seems devoted to the formation of ovules, while the other
half bears a one-celled anther. Lindley[342] has also described a case of this
kind in a species of Amaryllis.
Page 392
Fig. 164.—Structure half anther, half carpel, Lilium.
In Saxifraga crassifolia it sometimes happens that mixed with the stamens,
and originating with them, are a number of distinct and perfectly formed
carpels, wholly separated from the normal carpels, in the centre of the
flower. In this particular instance there is usually no intermediate condition
between the stamen and the pistil. Guillemin[343] also describes a
transformation of the stamens into carpels in Euphorbia esula.
When the anther is involved it may be only partially so, or almost the whole
organ may be transformed. As instances of very partial change may be cited
the passage of the connective into a stigma in Thalictrum minus, or the
passage of the points of the anthers into imperfect styles in some species of
bamboo.[344]
In Rosa arvensis similar transformations have been observed of a slightly
more complex character than those just mentioned, and passing into more
In Saxifraga crassifolia it sometimes happens that mixed with the stamens,
and originating with them, are a number of distinct and perfectly formed
carpels, wholly separated from the normal carpels, in the centre of the
flower. In this particular instance there is usually no intermediate condition
between the stamen and the pistil. Guillemin[343] also describes a
transformation of the stamens into carpels in Euphorbia esula.
When the anther is involved it may be only partially so, or almost the whole
organ may be transformed. As instances of very partial change may be cited
the passage of the connective into a stigma in Thalictrum minus, or the
passage of the points of the anthers into imperfect styles in some species of
bamboo.[344]
In Rosa arvensis similar transformations have been observed of a slightly
more complex character than those just mentioned, and passing into more
Page 393
important changes, especially to the formation of pollen within ovules,
formed on the edges of an open carpellodic anther (see p. 186).
Mr. Berkeley has recorded an analogous case in a gourd in which the
stamens bore numerous ovules (p. 200), and Baillon describes another
gourd in which certain fleshy appendages surrounding the andrœcium were
provided with ovules.[345]
Payer, in his 'Organogénie,' p. 38, mentions a stamen of Dionæa bearing not
only an anther, but likewise an ovule.
Sempervivum tectorum and S. montanum, have long been noticed as being
very prone to present this change. Mohl[346] remarks that, in the
transformation of the stamens to the pistil in the common houseleek, the
filament of the stamen generally preserves its form, the anthers alone
undergoing change. At other times, however, the transformation takes place
at the same time, both in the filament and in the anther. When the stamens
are numerous some of them remain in their normal state, while others, and
especially the inner ones, undergo a change. Sometimes all the stamens are
changed simultaneously, while at other times some of these organs may be
found in which the anther is partially filled with ovules, and partially with
pollen.
In the accompanying figures (fig. 165, a-h) a series of intermediate stages is
shown between the ordinary stamen of Sempervivum tectorum and the
ordinary carpel, from which it will be seen that the filament is little, if at all,
affected, and that in those cases where there is a combination of the
attributes of the stamen and of the pistil in the same organ the pollen is
formed in the upper or inner surface of the leaf-organ, while the ovules arise
from the opposite surface from the free edge, (b, c, d, e, f, g).
formed on the edges of an open carpellodic anther (see p. 186).
Mr. Berkeley has recorded an analogous case in a gourd in which the
stamens bore numerous ovules (p. 200), and Baillon describes another
gourd in which certain fleshy appendages surrounding the andrœcium were
provided with ovules.[345]
Payer, in his 'Organogénie,' p. 38, mentions a stamen of Dionæa bearing not
only an anther, but likewise an ovule.
Sempervivum tectorum and S. montanum, have long been noticed as being
very prone to present this change. Mohl[346] remarks that, in the
transformation of the stamens to the pistil in the common houseleek, the
filament of the stamen generally preserves its form, the anthers alone
undergoing change. At other times, however, the transformation takes place
at the same time, both in the filament and in the anther. When the stamens
are numerous some of them remain in their normal state, while others, and
especially the inner ones, undergo a change. Sometimes all the stamens are
changed simultaneously, while at other times some of these organs may be
found in which the anther is partially filled with ovules, and partially with
pollen.
In the accompanying figures (fig. 165, a-h) a series of intermediate stages is
shown between the ordinary stamen of Sempervivum tectorum and the
ordinary carpel, from which it will be seen that the filament is little, if at all,
affected, and that in those cases where there is a combination of the
attributes of the stamen and of the pistil in the same organ the pollen is
formed in the upper or inner surface of the leaf-organ, while the ovules arise
from the opposite surface from the free edge, (b, c, d, e, f, g).
Page 394
In a drawing made by the Rev. G. E. Smith of a malformed flower of
Primula acaulis, and which the writer has had the opportunity of
examining, the stamens are represented as detached from the corolla, and
their anthers replaced by open carpels, with ovules arising, not only from
their edges, but also from their surfaces, while the apex of the carpellary
leaf was drawn out into a long style, terminated by a flattened spathulate
stigma.
Delphinium elatum is one of the plants in which this change has been most
frequently noticed.[347]
Fig. 165.—Sempervivum tecotorum. a. Normal stamen. h. Normal carpel. b, c, e, f, g. Structure partly
staminal, partly carpellary. d. Transverse section through c, showing pollen internally, ovules
externally.
In willows the change of pistils into staminal organs has been frequently
observed. In Salix babylonica Prof. Schnizlein has described various
Primula acaulis, and which the writer has had the opportunity of
examining, the stamens are represented as detached from the corolla, and
their anthers replaced by open carpels, with ovules arising, not only from
their edges, but also from their surfaces, while the apex of the carpellary
leaf was drawn out into a long style, terminated by a flattened spathulate
stigma.
Delphinium elatum is one of the plants in which this change has been most
frequently noticed.[347]
Fig. 165.—Sempervivum tecotorum. a. Normal stamen. h. Normal carpel. b, c, e, f, g. Structure partly
staminal, partly carpellary. d. Transverse section through c, showing pollen internally, ovules
externally.
In willows the change of pistils into staminal organs has been frequently
observed. In Salix babylonica Prof. Schnizlein has described various
Page 395
transition stages between the carpels and the stamens, and in one instance,
in addition to this change, a perfect cup-shaped perianth was present, as
happens normally in Populus[348]. Mr. Lowe also records the conversion of
stamens into ovaries in Salix Andersoniana, and this by every conceivable
intermediate gradation.[349]
The following list will serve to show what plants are most subject to this
anomaly. It is difficult to draw any accurate inference from this
enumeration, but attention may be called to the frequency of this occurrence
in certain plants, such as the Sempervivum, the wallflower, the poppy, and
the heath. Why these plants should specially be subject to these changes
cannot be at present stated.
By the student of animal physiology such a change as above described—
equivalent to the substitution of an ovary or a uterus for a testis—would be
looked on as next to impossible; the simpler and less specialised structure
of plants renders such a change in them far more easy of comprehension.
Thalictrum minus.
Delphinium elatum.
Magnolia fuscata.
Bocconia cordata.
*Papaver bracteatum!
*somniferum!
nudicaule.
Dionæa muscipula!
Barbarea vulgaris.
*Cheiranthus Cheiri!
Cochlearia Armoracia.
Tropæolum majus.
Citrus Aurantium.
*Sempervivum tectorum!
montanum.
Begonia frigida!
Cucumis, sp.
in addition to this change, a perfect cup-shaped perianth was present, as
happens normally in Populus[348]. Mr. Lowe also records the conversion of
stamens into ovaries in Salix Andersoniana, and this by every conceivable
intermediate gradation.[349]
The following list will serve to show what plants are most subject to this
anomaly. It is difficult to draw any accurate inference from this
enumeration, but attention may be called to the frequency of this occurrence
in certain plants, such as the Sempervivum, the wallflower, the poppy, and
the heath. Why these plants should specially be subject to these changes
cannot be at present stated.
By the student of animal physiology such a change as above described—
equivalent to the substitution of an ovary or a uterus for a testis—would be
looked on as next to impossible; the simpler and less specialised structure
of plants renders such a change in them far more easy of comprehension.
Thalictrum minus.
Delphinium elatum.
Magnolia fuscata.
Bocconia cordata.
*Papaver bracteatum!
*somniferum!
nudicaule.
Dionæa muscipula!
Barbarea vulgaris.
*Cheiranthus Cheiri!
Cochlearia Armoracia.
Tropæolum majus.
Citrus Aurantium.
*Sempervivum tectorum!
montanum.
Begonia frigida!
Cucumis, sp.
Page 396
Cucurbita Pepo.
Pyrus Malus.
Rosa arvensis!
Saxifraga crassifolia!
Myrtus, sp.
Campanula rapunculoides.
Polemonium cæruleum.
Gentiana Amarella.
*Erica Tetralix.
Stachys germanica.
Primula acaulis.
Rumex crispus.
*Salix, sp. plur.!
Euphorbia esula.
Glochidion.
Asphodelus ramosus.
Amaryllis.
Lilium tigrinum!
longiflorum.
*Tulipa Gesneriana!
var. cult. plurim.!
Hemerocallis.
Zea Mays.
Bambusa, sp.
Pistillody of the ovule.—An instance of this extraordinary transformation
in the carnation, as observed by the Rev. Mr. Berkeley, is given at p. 268.
FOOTNOTES:
[295] 'Neue Denkschrift. Schweiz. Gesellsch.,' band v, p. 9.
[296] 'Bull. Acad. Belg.,' xix, part 2, p. 93.
[297] Schlechtendal, 'Linnæa,' ix, p. 737.
[298] Misbilld., 'Cult. Gewachs.,' p. 32.
[299] Linn., 'Phil. Botan.,' § 120.
[300] 'Bull. Soc. Bot. France,' 1859, vol. vi, p. 199.
Pyrus Malus.
Rosa arvensis!
Saxifraga crassifolia!
Myrtus, sp.
Campanula rapunculoides.
Polemonium cæruleum.
Gentiana Amarella.
*Erica Tetralix.
Stachys germanica.
Primula acaulis.
Rumex crispus.
*Salix, sp. plur.!
Euphorbia esula.
Glochidion.
Asphodelus ramosus.
Amaryllis.
Lilium tigrinum!
longiflorum.
*Tulipa Gesneriana!
var. cult. plurim.!
Hemerocallis.
Zea Mays.
Bambusa, sp.
Pistillody of the ovule.—An instance of this extraordinary transformation
in the carnation, as observed by the Rev. Mr. Berkeley, is given at p. 268.
FOOTNOTES:
[295] 'Neue Denkschrift. Schweiz. Gesellsch.,' band v, p. 9.
[296] 'Bull. Acad. Belg.,' xix, part 2, p. 93.
[297] Schlechtendal, 'Linnæa,' ix, p. 737.
[298] Misbilld., 'Cult. Gewachs.,' p. 32.
[299] Linn., 'Phil. Botan.,' § 120.
[300] 'Bull. Soc. Bot. France,' 1859, vol. vi, p. 199.
Page 397
[301] Seemann's 'Journal of Botany,' vol. iii, p. 105; also Morren, 'Bull. Acad. Belg.,' vol. xx,
part 2, p. 264.
[302] Morren, 'Bull. Belg.,' xviii, p. 503.
[303] 'Organ. Vég.,' t. i, p. 513.
[304] 'Bull. Acad. Roy. Belg.,' tome xvii; and Lobelia, p. 65.
[305] Masters, "On Double Flowers," 'Rep. Internat. Bot. Congress,' London, 1866. p. 127.
[306] See also C. Morren, "Sur les vraies fleurs doubles chez les Orchidées," 'Bull. Acad.
Roy. Belg.,' vol. xix, part ii, 1852. p. 171.
[307] C. Morren, 'Bull. Acad. Belg.,' vol. xx, 1853, part ii, p. 284 (Syringa).
[308] 'Rep. Bot. Congress,' London, 1866, p. 135, t. vii, f. 14.
[309] Although it is generally admitted that the filament of the stamen corresponds to the
stalk of the leaf, and the anther to the leaf-blade, yet there are some points on which
uncertainty still rests. One of these is as to the sutures of the anther. Do these chinks through
which the pollen escapes correspond (as would at first sight seem probable) to the margins of
the antheral leaf, or do they answer to the lines that separate the two pollen-cavities on each
half of the anther one from the other? Professor Oliver, 'Trans. Linn. Soc.,' vol. xxiii, 1862, p.
423, in alluding to the views held by others on this subject, concludes, from an examination of
some geranium flowers in which the stamens were more or less petaloid, that Bischoff's notion
as to the sutures of the anther is correct, viz., that they are the equivalents of the septa of
untransformed tissue between the pollen-sacs. Some double fuchsias ('Gard. Chron.,' 1863, p.
989) add confirmation to this opinion. In these flowers the petals were present as usual, but the
stamens were more or less petaloid, the filaments were unchanged, but the anthers existed in
the form of a petal-like cup from the centre of which projected two imperfect pollen-lobes (the
other two lobes being petaloid). Now, in this case, the margins of the anther were coherent to
form the cup, and the pollen was emitted along a line separating the polliniferous from the
petaloid portion of the anther. This view is also borne out by the double-flowered Arbutus
Unedo, and also by what occurs in some double violets, wherein the anther exists in the guise
of a broad lancet-shaped expansion, from the surface of which project four plates (fig. 157),
representing apparently the walls of the pollen-sacs, but destitute of pollen; the chink left
between these plates corresponds thus to the suture of the normal anther.
part 2, p. 264.
[302] Morren, 'Bull. Belg.,' xviii, p. 503.
[303] 'Organ. Vég.,' t. i, p. 513.
[304] 'Bull. Acad. Roy. Belg.,' tome xvii; and Lobelia, p. 65.
[305] Masters, "On Double Flowers," 'Rep. Internat. Bot. Congress,' London, 1866. p. 127.
[306] See also C. Morren, "Sur les vraies fleurs doubles chez les Orchidées," 'Bull. Acad.
Roy. Belg.,' vol. xix, part ii, 1852. p. 171.
[307] C. Morren, 'Bull. Acad. Belg.,' vol. xx, 1853, part ii, p. 284 (Syringa).
[308] 'Rep. Bot. Congress,' London, 1866, p. 135, t. vii, f. 14.
[309] Although it is generally admitted that the filament of the stamen corresponds to the
stalk of the leaf, and the anther to the leaf-blade, yet there are some points on which
uncertainty still rests. One of these is as to the sutures of the anther. Do these chinks through
which the pollen escapes correspond (as would at first sight seem probable) to the margins of
the antheral leaf, or do they answer to the lines that separate the two pollen-cavities on each
half of the anther one from the other? Professor Oliver, 'Trans. Linn. Soc.,' vol. xxiii, 1862, p.
423, in alluding to the views held by others on this subject, concludes, from an examination of
some geranium flowers in which the stamens were more or less petaloid, that Bischoff's notion
as to the sutures of the anther is correct, viz., that they are the equivalents of the septa of
untransformed tissue between the pollen-sacs. Some double fuchsias ('Gard. Chron.,' 1863, p.
989) add confirmation to this opinion. In these flowers the petals were present as usual, but the
stamens were more or less petaloid, the filaments were unchanged, but the anthers existed in
the form of a petal-like cup from the centre of which projected two imperfect pollen-lobes (the
other two lobes being petaloid). Now, in this case, the margins of the anther were coherent to
form the cup, and the pollen was emitted along a line separating the polliniferous from the
petaloid portion of the anther. This view is also borne out by the double-flowered Arbutus
Unedo, and also by what occurs in some double violets, wherein the anther exists in the guise
of a broad lancet-shaped expansion, from the surface of which project four plates (fig. 157),
representing apparently the walls of the pollen-sacs, but destitute of pollen; the chink left
between these plates corresponds thus to the suture of the normal anther.
Page 398
Fig. 157.—Petaloid stamen of Viola, with four projecting plates.
The inner or upper portion of the anther-leaf is that which is most intimately concerned in the
formation of pollen; it comparatively rarely (query ever) happens that the back or lower
surface of the antheral leaf is specially devoted to the formation of pollen. On the other hand,
in cases like those of the common houseleek, where we meet with petaloid organs combining
the attributes of anthers and of carpels, we find the inner layers devoted to the production of
pollen, the outer to the formation of ovules.
That the pollen-lobes are not to be taken as halves of a staminal leaf, but rather as specialised
portions of it, not necessarily occupying half its surface, is shown also in the case of double-
flowered Malvaceæ, in which the stamens are frequently partly petal-like, partly divided into
numerous separate filaments, each bearing a one-, or it may be even a two-lobed anther. This
circumstance is confirmatory of the opinion held by Payer, Duchartre, Dickson, and other
organogenists, as to the compound nature of the stamens in these plants. The stamens are here
analogues not of a simple entire leaf, but of a lobed, digitate, or compound leaf, each
subdivision bearing its separate anther. On this subject the reader may consult M. Müller's
paper on the anther of Jatropha Pohliana, &c., referred to at page 255.
[310] See C. Morren, "On Spur-shaped Nectarines," &c., 'Ann. Nat. Hist.,' March, 1841, p. 1.
tab. 11.
[311] Karsten, 'Flor. Columb. Spec.,' tab. xxix.
The inner or upper portion of the anther-leaf is that which is most intimately concerned in the
formation of pollen; it comparatively rarely (query ever) happens that the back or lower
surface of the antheral leaf is specially devoted to the formation of pollen. On the other hand,
in cases like those of the common houseleek, where we meet with petaloid organs combining
the attributes of anthers and of carpels, we find the inner layers devoted to the production of
pollen, the outer to the formation of ovules.
That the pollen-lobes are not to be taken as halves of a staminal leaf, but rather as specialised
portions of it, not necessarily occupying half its surface, is shown also in the case of double-
flowered Malvaceæ, in which the stamens are frequently partly petal-like, partly divided into
numerous separate filaments, each bearing a one-, or it may be even a two-lobed anther. This
circumstance is confirmatory of the opinion held by Payer, Duchartre, Dickson, and other
organogenists, as to the compound nature of the stamens in these plants. The stamens are here
analogues not of a simple entire leaf, but of a lobed, digitate, or compound leaf, each
subdivision bearing its separate anther. On this subject the reader may consult M. Müller's
paper on the anther of Jatropha Pohliana, &c., referred to at page 255.
[310] See C. Morren, "On Spur-shaped Nectarines," &c., 'Ann. Nat. Hist.,' March, 1841, p. 1.
tab. 11.
[311] Karsten, 'Flor. Columb. Spec.,' tab. xxix.
Page 399
[312] See Dickson, "On Diplostemonous Flowers," 'Trans. Bot. Soc. Edin.,' vol. viii, p. 100;
and on the Andrœcium of Mentzelia, &c., in Seemann's 'Journal of Botany,' vol. iii, p. 209, and
vol. iv (1866) p. 273 (Potentilla, &c.).
[313] See Baillon, 'Adansonia,' iii, p. 351, tab. 12, Sinapis.
[314] 'Bull. Acad. Belg.,' xvii, part i, p. 516, c. tab., and 'Lobelia,' p. 83.
[315] Cited in 'Bull. Soc. Bot. France,' xiv, p. 253 ('Rev. Bibl.').
[316] 'Bull. Soc. Bot. Fr.,' 1858, p. 331.
[317] 'Mem. Legum.,' p. 44.
[318] 'Bildungsabweich, 'Pflanz. Fam.,' tab. 8, f. 12.
[319] 'Atlas de Göthe' p. 55, t. 4, f. 18.
[320] Wiegmann, 'Bot. Zeit.,' 1831, p. 5, tab. i.
[321] 'Ic. Flor. Germ.,' xiii, tab. 112, cccclxiv, f. 2.
[322] Seemann's 'Journal of Botany,' 1867, p. 317, t. 72, A (Ophrys).
[323] 'Enum. Euphorb.' p. 53.
[324] 'Linnæa.' i, p. 457.
[325] 'De Balsam,' p. 17.
[326] B. Clarke, 'Arrangement of Phænog. Plants,' p. 23.
[327] See 'Engelmann,' p. 26, tab. 3, f. 10, 11, 14.
[328] 'Ann. Sc. Nat.,' ser. 2, t. viii, 1837, p. 58.
[329] 'Bot. Zeit.,' 4, 1846, 889.
[330] 'Verhandl. Nat. Hist. Ver. Preuss. Rheinl. und Westph.,' 1858, 1860, p. 381. Cramer
also, 'Bildungsabweich,' p. 90, cites a case in Pæonia where the carpel was open and petaloid,
and bore an anther on one margin, and four ovules on the other.
[331] 'Euphorbiaceæ,' p. 205.
[332] Seemann's 'Journ. Bot.,' iv, p. 168, tab. 47, f. 1.
and on the Andrœcium of Mentzelia, &c., in Seemann's 'Journal of Botany,' vol. iii, p. 209, and
vol. iv (1866) p. 273 (Potentilla, &c.).
[313] See Baillon, 'Adansonia,' iii, p. 351, tab. 12, Sinapis.
[314] 'Bull. Acad. Belg.,' xvii, part i, p. 516, c. tab., and 'Lobelia,' p. 83.
[315] Cited in 'Bull. Soc. Bot. France,' xiv, p. 253 ('Rev. Bibl.').
[316] 'Bull. Soc. Bot. Fr.,' 1858, p. 331.
[317] 'Mem. Legum.,' p. 44.
[318] 'Bildungsabweich, 'Pflanz. Fam.,' tab. 8, f. 12.
[319] 'Atlas de Göthe' p. 55, t. 4, f. 18.
[320] Wiegmann, 'Bot. Zeit.,' 1831, p. 5, tab. i.
[321] 'Ic. Flor. Germ.,' xiii, tab. 112, cccclxiv, f. 2.
[322] Seemann's 'Journal of Botany,' 1867, p. 317, t. 72, A (Ophrys).
[323] 'Enum. Euphorb.' p. 53.
[324] 'Linnæa.' i, p. 457.
[325] 'De Balsam,' p. 17.
[326] B. Clarke, 'Arrangement of Phænog. Plants,' p. 23.
[327] See 'Engelmann,' p. 26, tab. 3, f. 10, 11, 14.
[328] 'Ann. Sc. Nat.,' ser. 2, t. viii, 1837, p. 58.
[329] 'Bot. Zeit.,' 4, 1846, 889.
[330] 'Verhandl. Nat. Hist. Ver. Preuss. Rheinl. und Westph.,' 1858, 1860, p. 381. Cramer
also, 'Bildungsabweich,' p. 90, cites a case in Pæonia where the carpel was open and petaloid,
and bore an anther on one margin, and four ovules on the other.
[331] 'Euphorbiaceæ,' p. 205.
[332] Seemann's 'Journ. Bot.,' iv, p. 168, tab. 47, f. 1.
Page 400
[333] Moquin-Tandon, l. c., 220, Passiflora. Masters, 'Journ. Linn. Soc.,' 1857, p. 159,
Saponaria. Seemann's 'Journ. Botany,' vol. iii, p. 107, Narcissus.
[334] 'Bull. Soc. Bot. Fr.,' 1857, p. 452.
[335] 'Gardeners' Chronicle,' 1866, p. 897.
[336] Maout, 'Leçons Element.,' vol. ii, p. 488.
[337] Poiteau and Turpin, 'Arb. Fruit,' t. 37, and Trécul, 'Bull Soc. Bot. France,' vol. i. p. 307.
[338] Clos, 'Mem. Acad. Toulouse,' 5 ser., vol. iii.
[339] 'Bot. Zeit.,' 1850, t. viii, pp. 514, 664. 'Flora,' (B. Z.) 1832, t. xv, p. 252; also cited in
'Ann. des Serres et des jardins,' vi, pp. 241–5. See also Schlechtendal, 'Bot. Zeit.,' 1845, t. 3, p.
6.
[340] 'Bull. Soc. Bot. France,' t. viii, p. 453.
[341] See also Allmann, 'Rep. Brit. Assoc.,' July, 1851.
[342] 'Theory of Horticulture,' ed. 2, p. 82.
[343] 'Mém. Soc. Hist. Nat. Paris.' i, 16.
[344] Gen. Munro, 'Trans. Linn. Soc.,' xxvii, p. 7.
[345] 'Bull. Soc. Bot. Fr.,' 1857, p. 21.
[346] 'Ann. Scienc. Nat.,' t. viii, 1837, p. 50, and 'Bot. Zeit.' (R.), 1836, t. xix, p. 513, &c. See
also MM. Sourd Dussiples and G. Bergeron, 'Bull. Soc. Bot. France,' viii, p. 349; Von
Schmidel, 'Icon. plant. et Anal. part.' 1782, p. 210, fig. 54.
[347] Godron, 'Bull. Soc. Bot. Fr.,' xiii, p. 82, Rev. Bibl.
[348] Cited in Henfrey, 'Bot. Gazette,' iii, p. 12.
[349] 'Ann. Nat. Hist.,' September, 1856, p. 56. See also Kirschleger, 'Flora (Bot. Zeit.),' xxiv,
1841, p. 340, Salix alba. Henschel, 'Flora (Bot. Zeit.),' 1832, t. xv, p. 253, S. cinerea.
Hartmann, 'Flora (Bot. Zeit.),' xxiv, p. 199, S. nigricans. Meyer, C. A., 'Bull. Phys. Math.,' t. x,
S. alba.
Saponaria. Seemann's 'Journ. Botany,' vol. iii, p. 107, Narcissus.
[334] 'Bull. Soc. Bot. Fr.,' 1857, p. 452.
[335] 'Gardeners' Chronicle,' 1866, p. 897.
[336] Maout, 'Leçons Element.,' vol. ii, p. 488.
[337] Poiteau and Turpin, 'Arb. Fruit,' t. 37, and Trécul, 'Bull Soc. Bot. France,' vol. i. p. 307.
[338] Clos, 'Mem. Acad. Toulouse,' 5 ser., vol. iii.
[339] 'Bot. Zeit.,' 1850, t. viii, pp. 514, 664. 'Flora,' (B. Z.) 1832, t. xv, p. 252; also cited in
'Ann. des Serres et des jardins,' vi, pp. 241–5. See also Schlechtendal, 'Bot. Zeit.,' 1845, t. 3, p.
6.
[340] 'Bull. Soc. Bot. France,' t. viii, p. 453.
[341] See also Allmann, 'Rep. Brit. Assoc.,' July, 1851.
[342] 'Theory of Horticulture,' ed. 2, p. 82.
[343] 'Mém. Soc. Hist. Nat. Paris.' i, 16.
[344] Gen. Munro, 'Trans. Linn. Soc.,' xxvii, p. 7.
[345] 'Bull. Soc. Bot. Fr.,' 1857, p. 21.
[346] 'Ann. Scienc. Nat.,' t. viii, 1837, p. 50, and 'Bot. Zeit.' (R.), 1836, t. xix, p. 513, &c. See
also MM. Sourd Dussiples and G. Bergeron, 'Bull. Soc. Bot. France,' viii, p. 349; Von
Schmidel, 'Icon. plant. et Anal. part.' 1782, p. 210, fig. 54.
[347] Godron, 'Bull. Soc. Bot. Fr.,' xiii, p. 82, Rev. Bibl.
[348] Cited in Henfrey, 'Bot. Gazette,' iii, p. 12.
[349] 'Ann. Nat. Hist.,' September, 1856, p. 56. See also Kirschleger, 'Flora (Bot. Zeit.),' xxiv,
1841, p. 340, Salix alba. Henschel, 'Flora (Bot. Zeit.),' 1832, t. xv, p. 253, S. cinerea.
Hartmann, 'Flora (Bot. Zeit.),' xxiv, p. 199, S. nigricans. Meyer, C. A., 'Bull. Phys. Math.,' t. x,
S. alba.
Page 401
PART IV.
HETEROMORPHY.
There are certain malformations that have little in common beyond this, that
they cannot readily be allocated in either of the great groups proposed by
writers on teratology. There are also deformities which, unlike the majority
of deviations from the ordinary structure, are absolute and not relative.
While the latter are due to an exaggeration, or to an imperfection of
development, or, it may be, to a partial perversion in organization, the
former differ from the normal standard, not merely in degree, but
absolutely. This is often the case when disease or injury affects the plant;
for instance, in the case of galls arising from insect-puncture the structure is
rather a new growth altogether, than dependent on mere hypertrophy of the
original tissues. These absolute deformities arising from the causes just
mentioned belong rather to pathology than to teratology strictly so called;
but, under the head of deformities, may be mentioned sundry deviations not
elsewhere alluded to.
CHAPTER I.
DEFORMITIES.
The special meaning here attached to the term deformity is sufficiently
explained in the preceding paragraph; it remains to give a few illustrations,
and to refer to other headings, such as Heterotaxy, Hypertrophy, Atrophy,
&c., for malformations capable of more rigid classification than those here
alluded to.
HETEROMORPHY.
There are certain malformations that have little in common beyond this, that
they cannot readily be allocated in either of the great groups proposed by
writers on teratology. There are also deformities which, unlike the majority
of deviations from the ordinary structure, are absolute and not relative.
While the latter are due to an exaggeration, or to an imperfection of
development, or, it may be, to a partial perversion in organization, the
former differ from the normal standard, not merely in degree, but
absolutely. This is often the case when disease or injury affects the plant;
for instance, in the case of galls arising from insect-puncture the structure is
rather a new growth altogether, than dependent on mere hypertrophy of the
original tissues. These absolute deformities arising from the causes just
mentioned belong rather to pathology than to teratology strictly so called;
but, under the head of deformities, may be mentioned sundry deviations not
elsewhere alluded to.
CHAPTER I.
DEFORMITIES.
The special meaning here attached to the term deformity is sufficiently
explained in the preceding paragraph; it remains to give a few illustrations,
and to refer to other headings, such as Heterotaxy, Hypertrophy, Atrophy,
&c., for malformations capable of more rigid classification than those here
alluded to.
Page 402
Fig. 166.—Portion of the under surface of a cabbage-leaf, with horn-like excrescences projecting
from it.
Formation of tubes.—The production of ascidia or pitchers from the
cohesion of the margins of one or more leaves has been already alluded to
(see pp. 21, 30), but there is another class of cases in which the tubular
formation is due, not so much to the union of the margins of a leaf as to the
disproportionate growth of some portions as contrasted with others, whence
arises either a depressed cavity, as in the case of a leaf, or an expanded and
excavated structure, when the stem or some portion of it is affected.
The fruit of the rose, the apple, the fig, and many others, is now generally
admitted to be composed externally of the dilated end of the flower-stalk in
which the true carpels become imbedded. Between such cases and that of a
peltate leaf with a depressed centre, such as often occurs, to some extent, in
Nelumbium, there is but little difference.
from it.
Formation of tubes.—The production of ascidia or pitchers from the
cohesion of the margins of one or more leaves has been already alluded to
(see pp. 21, 30), but there is another class of cases in which the tubular
formation is due, not so much to the union of the margins of a leaf as to the
disproportionate growth of some portions as contrasted with others, whence
arises either a depressed cavity, as in the case of a leaf, or an expanded and
excavated structure, when the stem or some portion of it is affected.
The fruit of the rose, the apple, the fig, and many others, is now generally
admitted to be composed externally of the dilated end of the flower-stalk in
which the true carpels become imbedded. Between such cases and that of a
peltate leaf with a depressed centre, such as often occurs, to some extent, in
Nelumbium, there is but little difference.
Page 403
In cabbages and lettuces there not unfrequently occurs a production of leaf-
like processes projecting from the primary blade at a right angle (see
Enation). Sometimes these are developed in a tubular form, so as to form a
series of little horn-like tubes, or shallow troughs, as in Aristolochia sipho.
At other times the nerves or ribs of the leaf project beyond the blade, and
bear at their extremities structures similar to those just described.
Fig. 167.—Lettuce leaf, bearing on the back a stalked cup, arising from the dilatation of the stalk (?).
In a variety of Codiæum variegatum a similar formation may be seen to a
minor extent. Even the common Scolopendrium vulgare occasionally
produces small pitchers of this character, as in the varieties named perafero-
corautum, Moore, and peraferum, Woll.[350]
In carnations leaves may sometimes be seen from both surfaces, from
which project long, sharp-pointed tubular spurs at irregular intervals. A very
like processes projecting from the primary blade at a right angle (see
Enation). Sometimes these are developed in a tubular form, so as to form a
series of little horn-like tubes, or shallow troughs, as in Aristolochia sipho.
At other times the nerves or ribs of the leaf project beyond the blade, and
bear at their extremities structures similar to those just described.
Fig. 167.—Lettuce leaf, bearing on the back a stalked cup, arising from the dilatation of the stalk (?).
In a variety of Codiæum variegatum a similar formation may be seen to a
minor extent. Even the common Scolopendrium vulgare occasionally
produces small pitchers of this character, as in the varieties named perafero-
corautum, Moore, and peraferum, Woll.[350]
In carnations leaves may sometimes be seen from both surfaces, from
which project long, sharp-pointed tubular spurs at irregular intervals. A very
Page 404
singular illustration of this is figured by Trattinick,[351], in which the leaves,
epicalyx, sepals, and petals, were all provided with tubular spurs.
In Cephalotus follicularis rudimentary or imperfect pitchers may be
frequently met with, in which the stalk of the leaf is tubular and bears at its
extremity a very small rudimentary leaf-blade. It is not in all cases easy to
trace the origin and true nature of the ascidium, as the venation is
sometimes obscure. If there be a single well-marked midrib the probability
is that the case is one of cohesion of the margins of the leaf; but if the veins
are all of about equal size, and radiate from a common stalk, the pouch-like
formation is probably due to dilatation and hollowing of the petiole. Again,
when the result of a union of the margins of the leaf, the pitcher is generally
less regular than when formed from the hollowed end of a leaf-stalk.
Further information is especially needed as to the mode of development and
formation of these tubular organs, so as to ascertain clearly when they are
the result of a true cupping process, and when of cohesion of the margins of
one or more leaves. (See Cohesion, p. 31. For bibliographical references
consult also A. Braun, 'Flora v. Bot. Zeit.,' 1835, t. xviii, p. 41,
Aristolochia.)
Tubular formations in the flower.—A similar formation of tubes happens
in some double flowers; for instance, it is not infrequent in double flowers
of Primula sinensis, in which tubular petal-like structures are attached to
the inner surface of the corolla; sometimes these petaloid tubes replace the
stamens, while at other times they appear to have no relation to those
organs. In the particular flowers now alluded to the tubular form seems due
to a dilatation, and not to a cohesion of the margins. (See Cohesion, p. 23.)
These tubular petals resemble in form and colour almost precisely the
normal corolla in miniature, but are not surrounded by a calyx, nor do they
contain stamens, while the less perfect forms show clearly their origin from
a single tube-like organ.
epicalyx, sepals, and petals, were all provided with tubular spurs.
In Cephalotus follicularis rudimentary or imperfect pitchers may be
frequently met with, in which the stalk of the leaf is tubular and bears at its
extremity a very small rudimentary leaf-blade. It is not in all cases easy to
trace the origin and true nature of the ascidium, as the venation is
sometimes obscure. If there be a single well-marked midrib the probability
is that the case is one of cohesion of the margins of the leaf; but if the veins
are all of about equal size, and radiate from a common stalk, the pouch-like
formation is probably due to dilatation and hollowing of the petiole. Again,
when the result of a union of the margins of the leaf, the pitcher is generally
less regular than when formed from the hollowed end of a leaf-stalk.
Further information is especially needed as to the mode of development and
formation of these tubular organs, so as to ascertain clearly when they are
the result of a true cupping process, and when of cohesion of the margins of
one or more leaves. (See Cohesion, p. 31. For bibliographical references
consult also A. Braun, 'Flora v. Bot. Zeit.,' 1835, t. xviii, p. 41,
Aristolochia.)
Tubular formations in the flower.—A similar formation of tubes happens
in some double flowers; for instance, it is not infrequent in double flowers
of Primula sinensis, in which tubular petal-like structures are attached to
the inner surface of the corolla; sometimes these petaloid tubes replace the
stamens, while at other times they appear to have no relation to those
organs. In the particular flowers now alluded to the tubular form seems due
to a dilatation, and not to a cohesion of the margins. (See Cohesion, p. 23.)
These tubular petals resemble in form and colour almost precisely the
normal corolla in miniature, but are not surrounded by a calyx, nor do they
contain stamens, while the less perfect forms show clearly their origin from
a single tube-like organ.
Page 405
Fig. 168.—Corolla of Primula sinensis turned back to show a tubular petal springing from it. One
only is shown for the sake of clearness; they are generally numerous.
The formation of spurs or spur-like tubes in a quasi-regular manner has
been spoken of under the head of Irregular Peloria, p. 228, but we
occasionally meet with tubular processes which seem to occur in an
irregular manner, and to have no reference to the symmetrical plan of the
flower, and which are due probably to the same causes as those which
induce hypertrophy. Such spurs have frequently been seen on the corolla of
Digitalis purpurea, Antirrhinum majus,[352] Tulipa Gesneriana, and
occasionally on the sepals of Fuchsia. They are very frequent in some
seasons in the corolla of certain calceolarias (C. floribunda). By Morren this
production of adventitious spurs was called "Ceratomanie."
only is shown for the sake of clearness; they are generally numerous.
The formation of spurs or spur-like tubes in a quasi-regular manner has
been spoken of under the head of Irregular Peloria, p. 228, but we
occasionally meet with tubular processes which seem to occur in an
irregular manner, and to have no reference to the symmetrical plan of the
flower, and which are due probably to the same causes as those which
induce hypertrophy. Such spurs have frequently been seen on the corolla of
Digitalis purpurea, Antirrhinum majus,[352] Tulipa Gesneriana, and
occasionally on the sepals of Fuchsia. They are very frequent in some
seasons in the corolla of certain calceolarias (C. floribunda). By Morren this
production of adventitious spurs was called "Ceratomanie."
Page 406
Fig. 169.—Corolla of Calceolaria, showing irregular tubular spurs projecting from the lower lip.
Similar processes may sometimes be seen in the capsules of Linaria
vulgaris, as also in the fruits of some of the Solanums, quite without
reference to the arrangement of the carpels, so that their production seems
to be purely irregular.
Morren, as previously remarked, gave the name "Solenaidie" to tubular
deformities affecting the stamens, a term which has not been generally
adopted; the deformity in question is by no means of uncommon occurrence
in some double or partially pelorised flowers, as Antirrhinum, Linaria, &c.
A similar formation of conical out-growths may frequently be met with in
the fruits quite irrespectively of any disjunction of the carpels.
Contortion.—An irregular twisting or bending of the stem or branches is
by no means of uncommon occurrence, the inducing causes being often
some restriction to growth in certain directions, or the undue or
Similar processes may sometimes be seen in the capsules of Linaria
vulgaris, as also in the fruits of some of the Solanums, quite without
reference to the arrangement of the carpels, so that their production seems
to be purely irregular.
Morren, as previously remarked, gave the name "Solenaidie" to tubular
deformities affecting the stamens, a term which has not been generally
adopted; the deformity in question is by no means of uncommon occurrence
in some double or partially pelorised flowers, as Antirrhinum, Linaria, &c.
A similar formation of conical out-growths may frequently be met with in
the fruits quite irrespectively of any disjunction of the carpels.
Contortion.—An irregular twisting or bending of the stem or branches is
by no means of uncommon occurrence, the inducing causes being often
some restriction to growth in certain directions, or the undue or
Page 407
disproportionate growth in one direction, as contrasted with that in another.
Hence it may arise from insect-puncture, parasitic growth, or any obstacle
to the natural development. Frequently it exists in conjunction with
fasciation, the ends of the branches being curved round like a shepherd's
crook, from the growth on one side being so much greater than on the other.
Sometimes it is a mere exaggeration of a normal condition; thus, in what
are termed flexuose stems the stem twists alternately to one side or another,
frequently in association with an oblique form of the leaf. This state is
sometimes present to an extreme degree, as in some varieties of shrubs
(Cratægus, Robinia, &c.) cultivated for their singularly tortuous branches.
Fig. 170.—Portion of the culm of a Juncus, bent irregularly.
Hence it may arise from insect-puncture, parasitic growth, or any obstacle
to the natural development. Frequently it exists in conjunction with
fasciation, the ends of the branches being curved round like a shepherd's
crook, from the growth on one side being so much greater than on the other.
Sometimes it is a mere exaggeration of a normal condition; thus, in what
are termed flexuose stems the stem twists alternately to one side or another,
frequently in association with an oblique form of the leaf. This state is
sometimes present to an extreme degree, as in some varieties of shrubs
(Cratægus, Robinia, &c.) cultivated for their singularly tortuous branches.
Fig. 170.—Portion of the culm of a Juncus, bent irregularly.
Page 408
Fig. 171.—Portion of a branch of Cratægus oxyacantha, var. tortuosa.
Such cases as those just mentioned, however, are but slightly irregular
compared to others in which the deformity exists to such an extent that the
traces of the ordinary mode of growth are almost obliterated. M. Moquin-
Tandon[353] alludes to a case of this kind in a species of pine (Pinus), in
which a branch ended in four unequal divisions, which were strongly
curved from without inwards, then became united in pairs, these latter in
their turn blending into a single mass.
In the case of some beeches growing in the forest of Verzy, near Rheims,
the trunks of the trees are contorted in every direction, and, at a height of
from fifteen to twenty feet, a number of branches are also given off, also
much contorted, and occasionally intergrafted, so that it seems as if a heavy
weight had been placed on the trees and literally flattened them. Similar
Such cases as those just mentioned, however, are but slightly irregular
compared to others in which the deformity exists to such an extent that the
traces of the ordinary mode of growth are almost obliterated. M. Moquin-
Tandon[353] alludes to a case of this kind in a species of pine (Pinus), in
which a branch ended in four unequal divisions, which were strongly
curved from without inwards, then became united in pairs, these latter in
their turn blending into a single mass.
In the case of some beeches growing in the forest of Verzy, near Rheims,
the trunks of the trees are contorted in every direction, and, at a height of
from fifteen to twenty feet, a number of branches are also given off, also
much contorted, and occasionally intergrafted, so that it seems as if a heavy
weight had been placed on the trees and literally flattened them. Similar
Page 409
malformations may occasionally be met with in the branches of the oak, and
commonly in the weeping ash.
M. Fournier[354] mentions the stems of Ruscus aculeatus rolled in a circle,
others twisted spirally.
The phenomenon is not confined to woody plants, but has been met with in
chicory, in Antirrhinum, and other herbaceous species.
It is very difficult in some cases to separate these instances of irregular
torsion from those in which the twisting takes place in a more or less
regular spiral direction. In the former case the fibres of the plant are only
indirectly involved, but in the latter the fibres themselves are coiled spirally
from right to left, or vice versâ (spiral torsion), while not unfrequently both
conditions may be met with at the same time.
The leaves also are subject to similar deformities, of which a notable
illustration has been recorded in the case of the date palm, Phœnix
dactylifera, originally observed by Goethe, and figured and described by
Jaeger;[355] the leaves are folded and twisted in every direction, in
consequence of the fibrous band or cord which surrounds the leaves, and
which generally breaks as the leaflets increase in size, remaining from some
cause or other unbroken, and thus serving to restrain the growth.
A similar irregularity of growth occurs, not unfrequently, in the case of
crocus leaves, when in the course of their growth, as they push their way
through the soil, their progress becomes checked either by a stone or even
by frost.
Spiral torsion.—Growth in a spiral direction, and the arrangement of the
various organs of the plant in a spiral manner, are among the most common
of natural phenomena in plants.[356] Fibres are coiled spirally in the minute
commonly in the weeping ash.
M. Fournier[354] mentions the stems of Ruscus aculeatus rolled in a circle,
others twisted spirally.
The phenomenon is not confined to woody plants, but has been met with in
chicory, in Antirrhinum, and other herbaceous species.
It is very difficult in some cases to separate these instances of irregular
torsion from those in which the twisting takes place in a more or less
regular spiral direction. In the former case the fibres of the plant are only
indirectly involved, but in the latter the fibres themselves are coiled spirally
from right to left, or vice versâ (spiral torsion), while not unfrequently both
conditions may be met with at the same time.
The leaves also are subject to similar deformities, of which a notable
illustration has been recorded in the case of the date palm, Phœnix
dactylifera, originally observed by Goethe, and figured and described by
Jaeger;[355] the leaves are folded and twisted in every direction, in
consequence of the fibrous band or cord which surrounds the leaves, and
which generally breaks as the leaflets increase in size, remaining from some
cause or other unbroken, and thus serving to restrain the growth.
A similar irregularity of growth occurs, not unfrequently, in the case of
crocus leaves, when in the course of their growth, as they push their way
through the soil, their progress becomes checked either by a stone or even
by frost.
Spiral torsion.—Growth in a spiral direction, and the arrangement of the
various organs of the plant in a spiral manner, are among the most common
of natural phenomena in plants.[356] Fibres are coiled spirally in the minute
Page 410
vessels of flowering plants, and are not wholly wanting even among fungi.
The leaf-organs are very generally spirally arranged; the leaf-stalks are
often so twisted as to bring leaves on one plane which otherwise would
occupy several. In the leaf itself we have a spiral twist taking place
constantly in Alstrœmeria, in Avena, and other plants. A similar tendency is
manifested in the flower-stalks, as in Cyclamen and Vallisneria, and the
whole inflorescence, as in Spiranthes. Even the bark and wood of trees is
often disposed spirally. This is very noticeable in some firs, and in the bark
of the sweet chestnut (Castanea), of Thuja occidentalis, and other trees. The
knaurs or excrescences which are sometimes found on the roots or stems of
trees afford other illustrations of this universal tendency. These bodies
consist of a number of embryo buds, which, from some cause or other, are
incapable of lengthening. On examination every rudimentary or
undeveloped bud may be seen to be surrounded by densely crowded fibres
arranged spirally.
The axes of nearly all twining plants are themselves twisted, and twisted in
a direction corresponding to the spontaneous revolving movement exhibited
by these plants, as in the hop, the convolvulus, passion flower, &c., the
degree of twisting being dependent to a great extent on the roughness of the
surface around which the stem twines[357].
Considered as an exceptional occurrence, it occurs frequently in certain
plants, and, when it affects the stem or branches, necessarily causes some
changes in the arrangement of the parts attached to them; thus, spiral torsion
of the axial organs is generally accompanied by displacement of the leaves,
whorled leaves becoming alternate, and opposite or whorled leaves
becoming arranged on one side of the stem only. Frequently also this
condition is associated with fasciation, or, at least, with a distended or
dilated state. An illustration of this in Asparagus has been figured at p. 14.
The leaf-organs are very generally spirally arranged; the leaf-stalks are
often so twisted as to bring leaves on one plane which otherwise would
occupy several. In the leaf itself we have a spiral twist taking place
constantly in Alstrœmeria, in Avena, and other plants. A similar tendency is
manifested in the flower-stalks, as in Cyclamen and Vallisneria, and the
whole inflorescence, as in Spiranthes. Even the bark and wood of trees is
often disposed spirally. This is very noticeable in some firs, and in the bark
of the sweet chestnut (Castanea), of Thuja occidentalis, and other trees. The
knaurs or excrescences which are sometimes found on the roots or stems of
trees afford other illustrations of this universal tendency. These bodies
consist of a number of embryo buds, which, from some cause or other, are
incapable of lengthening. On examination every rudimentary or
undeveloped bud may be seen to be surrounded by densely crowded fibres
arranged spirally.
The axes of nearly all twining plants are themselves twisted, and twisted in
a direction corresponding to the spontaneous revolving movement exhibited
by these plants, as in the hop, the convolvulus, passion flower, &c., the
degree of twisting being dependent to a great extent on the roughness of the
surface around which the stem twines[357].
Considered as an exceptional occurrence, it occurs frequently in certain
plants, and, when it affects the stem or branches, necessarily causes some
changes in the arrangement of the parts attached to them; thus, spiral torsion
of the axial organs is generally accompanied by displacement of the leaves,
whorled leaves becoming alternate, and opposite or whorled leaves
becoming arranged on one side of the stem only. Frequently also this
condition is associated with fasciation, or, at least, with a distended or
dilated state. An illustration of this in Asparagus has been figured at p. 14.
Page 411
Very often the leaves are produced in a spiral line round the stem, as in a
specimen of Dracocephalum speciosum described and figured by C.
Morren. The leaves of this plant are naturally rectiserial and decussate, but,
in the twisted stem the leaves were curviserial, and arranged according to
the 5/13 plan. Now, referring to the ordinary notation of alternate leaves, we
shall have the first leaf covered by the fifth, with two turns of the spiral;
since decussate leaves result from two conjugate lines, the formula will be
necessarily 2/5. The fraction 5/13 hence comes regularly into the 2/5 series
(2/5, 3/8, 5/13). Thus, the leaves in assuming a new phyllotaxy, take one
quite analogous to the normal one.
One of the most curious instances that have fallen under the writer's own
observation occurred in the stem of Dipsacus fullonum. (See 'Proceedings
of the Linnean Society,' March 6, 1855, vol. ii, p. 370). The stem was
distended, and hollow, and twisted on itself; its fibres, moreover, were
arranged in an oblique or spiral direction; the branches or leaf-stalks, which
usually are arranged in an opposite and decussate manner, were, in this
case, disposed in a linear series, one over the other, following the line of
curvature of the stem. When the course of the fibres was traced from the
base of one of the stalks, upward around the stem, a spiral was found to be
completed at the base of the second stalk, above that which was made the
starting point. Now, if opposite leaves depend on the shortened condition of
the internode between the two leaves, then, in the teazel-stem just
described, each turn of the spiral would represent a lengthened internode;
and, if the fibres of this specimen could be untwisted, and made to assume
the vertical direction, and, at the same time, the internodes were shortened,
the result would be the opposition of the branches and the decussation of
the pairs; this explanation is borne out by the similar twisting which takes
place so frequently in the species of Galium and other Rubiaceæ.
specimen of Dracocephalum speciosum described and figured by C.
Morren. The leaves of this plant are naturally rectiserial and decussate, but,
in the twisted stem the leaves were curviserial, and arranged according to
the 5/13 plan. Now, referring to the ordinary notation of alternate leaves, we
shall have the first leaf covered by the fifth, with two turns of the spiral;
since decussate leaves result from two conjugate lines, the formula will be
necessarily 2/5. The fraction 5/13 hence comes regularly into the 2/5 series
(2/5, 3/8, 5/13). Thus, the leaves in assuming a new phyllotaxy, take one
quite analogous to the normal one.
One of the most curious instances that have fallen under the writer's own
observation occurred in the stem of Dipsacus fullonum. (See 'Proceedings
of the Linnean Society,' March 6, 1855, vol. ii, p. 370). The stem was
distended, and hollow, and twisted on itself; its fibres, moreover, were
arranged in an oblique or spiral direction; the branches or leaf-stalks, which
usually are arranged in an opposite and decussate manner, were, in this
case, disposed in a linear series, one over the other, following the line of
curvature of the stem. When the course of the fibres was traced from the
base of one of the stalks, upward around the stem, a spiral was found to be
completed at the base of the second stalk, above that which was made the
starting point. Now, if opposite leaves depend on the shortened condition of
the internode between the two leaves, then, in the teazel-stem just
described, each turn of the spiral would represent a lengthened internode;
and, if the fibres of this specimen could be untwisted, and made to assume
the vertical direction, and, at the same time, the internodes were shortened,
the result would be the opposition of the branches and the decussation of
the pairs; this explanation is borne out by the similar twisting which takes
place so frequently in the species of Galium and other Rubiaceæ.
Page 412
Fig. 172.—Twisted stem of Dipsacus fullonum.
G. Franc[358] was one of the first to notice this twisting in Galium, and M.
Duchartre,[359] in mentioning a similar instance, gives the following
explanation of the appearance which will be found to apply to most of these
cases. In the normal stem of Galium Mollugo the branches are opposite in
each verticil and crossed in the two successive ones. The stem is four-
angled, each angle having a nerve. Each of these nerves, springing from the
origin of a branch in one whorl, terminates in the interval which separates
the point of origin of the two branches in the whorl next above it. In the
deformed stem one of the nerves corresponds to the insertion of a branch,
its neighbour is in the adjoining vacant space; hence it results that four
nerves correspond to two branches and to two consecutive interspaces, and
hence the analogy between a single normal internode provided with its two
branches and its four nerves. What confirms this inference is that the nerve,
which begins at the point of origin of a branch, after making one spiral turn
G. Franc[358] was one of the first to notice this twisting in Galium, and M.
Duchartre,[359] in mentioning a similar instance, gives the following
explanation of the appearance which will be found to apply to most of these
cases. In the normal stem of Galium Mollugo the branches are opposite in
each verticil and crossed in the two successive ones. The stem is four-
angled, each angle having a nerve. Each of these nerves, springing from the
origin of a branch in one whorl, terminates in the interval which separates
the point of origin of the two branches in the whorl next above it. In the
deformed stem one of the nerves corresponds to the insertion of a branch,
its neighbour is in the adjoining vacant space; hence it results that four
nerves correspond to two branches and to two consecutive interspaces, and
hence the analogy between a single normal internode provided with its two
branches and its four nerves. What confirms this inference is that the nerve,
which begins at the point of origin of a branch, after making one spiral turn
Page 413
round the stem, terminates in the interval that separates the two following
branches, just as in a branch of the normal stem it ends in the upper whorl
between the two next branches. The torsion, then, in this Galium caused the
separation of the two opposite branches of the same verticil, and placed
them one above another, and this being reproduced in all the whorls, all the
branches come to be arranged on the same longitudinal line. The leaves are
susceptible of the same explanation; they are inserted in groups of three or
four in one arc round the origin of each branch. In the malformation each
series or group of four leaves, with its central branch, is equivalent to half a
whorl of the natural plant with its axillary branch. In other words, the
malformation consists in a torsion of the stem, which separates each whorl
into two distinct halves; these half-whorls, with their axillary branches, are
placed on a single longitudinal series one above another. This case is quoted
at some length, as it is an admirable example of a very common form of
malformation in these plants.
In some parts of Holland where madder is cultivated a similar deformation
is particularly frequent. The leaves, however, are not always grouped in the
way in which they were described by M. Duchartre, but more commonly
form a single continuous line; when arranged in leaf-whorls it generally
happens that some of the leaves are turned downwards, while others are
erect. It has been said that this condition occurs particularly frequently in
plants growing in damp places. It is certainly true that spiral torsion of the
stem is specially frequent in the species of Equisetum, most of which grow
in such spots. In these plants either the whole of the upper part of the stem
is thus twisted, or a portion only: thus Reinsch[360] cites a case in Equisetum
Telmateia, where the upper and lower portions of the stem were normal,
while the intermediate portion was twisted spirally. In this instance the
whorl next beneath the spiral had twenty-eight branchlets, and that
immediately above it thirty. Along the course of the spire there were two
hundred and three; dividing this latter number by the mean of the two
branches, just as in a branch of the normal stem it ends in the upper whorl
between the two next branches. The torsion, then, in this Galium caused the
separation of the two opposite branches of the same verticil, and placed
them one above another, and this being reproduced in all the whorls, all the
branches come to be arranged on the same longitudinal line. The leaves are
susceptible of the same explanation; they are inserted in groups of three or
four in one arc round the origin of each branch. In the malformation each
series or group of four leaves, with its central branch, is equivalent to half a
whorl of the natural plant with its axillary branch. In other words, the
malformation consists in a torsion of the stem, which separates each whorl
into two distinct halves; these half-whorls, with their axillary branches, are
placed on a single longitudinal series one above another. This case is quoted
at some length, as it is an admirable example of a very common form of
malformation in these plants.
In some parts of Holland where madder is cultivated a similar deformation
is particularly frequent. The leaves, however, are not always grouped in the
way in which they were described by M. Duchartre, but more commonly
form a single continuous line; when arranged in leaf-whorls it generally
happens that some of the leaves are turned downwards, while others are
erect. It has been said that this condition occurs particularly frequently in
plants growing in damp places. It is certainly true that spiral torsion of the
stem is specially frequent in the species of Equisetum, most of which grow
in such spots. In these plants either the whole of the upper part of the stem
is thus twisted, or a portion only: thus Reinsch[360] cites a case in Equisetum
Telmateia, where the upper and lower portions of the stem were normal,
while the intermediate portion was twisted spirally. In this instance the
whorl next beneath the spiral had twenty-eight branchlets, and that
immediately above it thirty. Along the course of the spire there were two
hundred and three; dividing this latter number by the mean of the two
Page 414
preceding, it was seen that the spire included the constituents of seven
ordinary verticils.
Fig. 173.—Stem of Galium spirally twisted. From a specimen communicated by Mr. Darwin.
Here also may be mentioned a curious bamboo, the stem of which is
preserved in the British Museum, and in which the internodes, on the
exterior, and the corresponding diaphragms and cavities within are spiral or
oblique in direction.
The root is also subject to the same malformation, the inducing cause being
usually some obstruction to downward growth, as when a plant has been
grown in a small pot, and becomes, as gardeners say, pot-bound.
ordinary verticils.
Fig. 173.—Stem of Galium spirally twisted. From a specimen communicated by Mr. Darwin.
Here also may be mentioned a curious bamboo, the stem of which is
preserved in the British Museum, and in which the internodes, on the
exterior, and the corresponding diaphragms and cavities within are spiral or
oblique in direction.
The root is also subject to the same malformation, the inducing cause being
usually some obstruction to downward growth, as when a plant has been
grown in a small pot, and becomes, as gardeners say, pot-bound.
Page 415
Fig. 174.—Showing "pot-bound" root twisted spirally (from the 'Gard. Chron.,' 1849).
The axial portion of the flower, the thalamus, is also occasionally twisted in
a spiral direction, the lateral parts of the flower being in consequence
displaced. Morren spoke of this displacement of the floral organs as
"speiranthie."[361]
Morren draws a distinction between spiral-torsion or spiralism and the less
regular torsion spoken of in the preceding section; in the former case not
only is the axis twisted, but its constituent fibres also. The condition in
question in some cases seems to be inherited in the seedling plants.
The following is a list of the plants in which spiral torsion of the stem or
branches has been most frequently observed. (See also under Fasciation and
Contortion.)
Hesperis matronalis.
Dianthus barbatus.
The axial portion of the flower, the thalamus, is also occasionally twisted in
a spiral direction, the lateral parts of the flower being in consequence
displaced. Morren spoke of this displacement of the floral organs as
"speiranthie."[361]
Morren draws a distinction between spiral-torsion or spiralism and the less
regular torsion spoken of in the preceding section; in the former case not
only is the axis twisted, but its constituent fibres also. The condition in
question in some cases seems to be inherited in the seedling plants.
The following is a list of the plants in which spiral torsion of the stem or
branches has been most frequently observed. (See also under Fasciation and
Contortion.)
Hesperis matronalis.
Dianthus barbatus.
Page 416
Pyrus Malus.
torminalis.
Cercis siliquastrum!
Punica Granatum.
Robinia pseudacacia!
Rubia tinctorum.
Dipsacus fullonum!
pilosus.
Gmelini.
Scabiosa arvensis.
*Valeriana officinalis!
dioica!
Galium aparine!
*Mollugo!
verum!
Hippuris vulgaris!
Veronica spicata.
longifolia.
Hyssopus officinalis.
Thymus Serpyllum.
Lamium purpureum!
Dracocephalum speciosum.
Mentha aquatica.
Mentha viridis.
Fraxinus vulgaris!
Sambucus nigra.
Zinnia.
Phylica.
Beta.
Rumex, sp.
Ulmus campestris.
Casuarina rigida.
Abies excelsa!
Lilium Martagon!
candidum.
*Asparagus officinalis!
Sagittaria sagittifolia.
Epipactis palustris.
Triticum repens!
torminalis.
Cercis siliquastrum!
Punica Granatum.
Robinia pseudacacia!
Rubia tinctorum.
Dipsacus fullonum!
pilosus.
Gmelini.
Scabiosa arvensis.
*Valeriana officinalis!
dioica!
Galium aparine!
*Mollugo!
verum!
Hippuris vulgaris!
Veronica spicata.
longifolia.
Hyssopus officinalis.
Thymus Serpyllum.
Lamium purpureum!
Dracocephalum speciosum.
Mentha aquatica.
Mentha viridis.
Fraxinus vulgaris!
Sambucus nigra.
Zinnia.
Phylica.
Beta.
Rumex, sp.
Ulmus campestris.
Casuarina rigida.
Abies excelsa!
Lilium Martagon!
candidum.
*Asparagus officinalis!
Sagittaria sagittifolia.
Epipactis palustris.
Triticum repens!
Page 417
Lolium perenne!
Phleum pratense.
Juncus conglomeratus!
Scirpus lacustris.
Equisetum Telmateia.
limosum.
fluviatile.
arvense!
Among the more important papers relating to this subject may be
mentioned:
Phleum pratense.
Juncus conglomeratus!
Scirpus lacustris.
Equisetum Telmateia.
limosum.
fluviatile.
arvense!
Among the more important papers relating to this subject may be
mentioned:
Page 418
Moquin-Tandon, 'El. Ter. Veg.,' p. 181. Kros, 'De Spira in plantis conspicua.' Morren, 'Bull.
Acad. Roy. Belg.,' 1851, tom. xviii, part i, p. 27. Milde, 'Nov. Act. Acad. Leop. Carol. Nat.
Cur., 1839. Ibid., vol. xxvi, part ii, p. 429, Equisetum. Irmisch, 'Flora,' 1858, t. ii,
Equisetum. Vrolik, 'Nouv. Mem. Instit. Amsterdam,' Lilium. Schlechtendal, 'Bot. Zeit.,' xiv,
p. 69, et v, p. 66. De Candolle, 'Organ. Veget., t. i, p. 155, tab. xxxvi, Mentha, &c. Alph. de
Candolle, 'Neue Denkschr. Allg. Schweiz. Gesellschft.,' band v, tab. vi, Valeriana.
Duchartre, 'Ann. Sc. Nat.,' ser. 3, vol. i, p. 292. 'Gardeners' Chronicle,' July 5, 1856, p. 452,
c. ic. xylogr., spiral branches from Guatemala—tree not known.
Spiral twisting of the leaf is scarcely of so common occurrence as the
corresponding condition in the stem. In Alstrœmeria it occurs normally, as
also in some grasses. In the variety annularis of Salix babylonica the leaf is
constantly coiled round spirally. A similar contortion occurs in a variety of
Codiæum variegatum lately introduced from the islands of the South Seas
by Mr. J. G. Veitch.
Fern fronds are occasionally found twisted in the same manner, e.g.
Scolopendrium vulgare var. spirale.[362]
Adventitious tendrils.—Under ordinary circumstances tendrils may be
described as modifications of the leaf, the stipule, the branch, or of the
flower stalk, so that it is not a matter of surprise to find tendrils occasionally
springing from the sepals or petals, as indeed happens normally in
Hodgsonia, Strophanthus, &c.
M. Decaisne[363] found a flower of the melon in which one of the segments
of the calyx was prolonged into a tendril, and Kirschleger records a similar
instance in the cucumber, while Mr. Holland ('Science Gossip,' 1865, p.
105) mentions a case in which one of the prickles on the fruit of a cucumber
had grown out into a tendril.
In Cobæa scandens the foliar nature of the tendril is shown by the
occasional presence of a small leaflet on one of the branches of the tendril,
Acad. Roy. Belg.,' 1851, tom. xviii, part i, p. 27. Milde, 'Nov. Act. Acad. Leop. Carol. Nat.
Cur., 1839. Ibid., vol. xxvi, part ii, p. 429, Equisetum. Irmisch, 'Flora,' 1858, t. ii,
Equisetum. Vrolik, 'Nouv. Mem. Instit. Amsterdam,' Lilium. Schlechtendal, 'Bot. Zeit.,' xiv,
p. 69, et v, p. 66. De Candolle, 'Organ. Veget., t. i, p. 155, tab. xxxvi, Mentha, &c. Alph. de
Candolle, 'Neue Denkschr. Allg. Schweiz. Gesellschft.,' band v, tab. vi, Valeriana.
Duchartre, 'Ann. Sc. Nat.,' ser. 3, vol. i, p. 292. 'Gardeners' Chronicle,' July 5, 1856, p. 452,
c. ic. xylogr., spiral branches from Guatemala—tree not known.
Spiral twisting of the leaf is scarcely of so common occurrence as the
corresponding condition in the stem. In Alstrœmeria it occurs normally, as
also in some grasses. In the variety annularis of Salix babylonica the leaf is
constantly coiled round spirally. A similar contortion occurs in a variety of
Codiæum variegatum lately introduced from the islands of the South Seas
by Mr. J. G. Veitch.
Fern fronds are occasionally found twisted in the same manner, e.g.
Scolopendrium vulgare var. spirale.[362]
Adventitious tendrils.—Under ordinary circumstances tendrils may be
described as modifications of the leaf, the stipule, the branch, or of the
flower stalk, so that it is not a matter of surprise to find tendrils occasionally
springing from the sepals or petals, as indeed happens normally in
Hodgsonia, Strophanthus, &c.
M. Decaisne[363] found a flower of the melon in which one of the segments
of the calyx was prolonged into a tendril, and Kirschleger records a similar
instance in the cucumber, while Mr. Holland ('Science Gossip,' 1865, p.
105) mentions a case in which one of the prickles on the fruit of a cucumber
had grown out into a tendril.
In Cobæa scandens the foliar nature of the tendril is shown by the
occasional presence of a small leaflet on one of the branches of the tendril,
Page 419
and a similar appearance may frequently be seen in Eccremocarpus scaber.
On the other hand, in the vine, the axial nature of the tendril is revealed by
the not infrequent presence of flowers or berries on them, as also in
Modecca and some Passifloraceæ.
Darwin, speaking of the tendrils of Bignonia capreolata, says it is a highly
remarkable fact that a leaf should be metamorphosed into a branched organ,
which turns from the light, and which can, by its extremities, either crawl
like a root into crevices, or seize hold of minute projecting points, these
extremities subsequently forming cellular masses, which envelope by their
growth the first fibres and secrete an adhesive cement.
Interrupted growth.—This term is here used in the same sense as in
ordinary descriptive botany, as when an "interruptedly pinnate" leaf is
spoken of. A similar alternation may be observed occasionally as a
teratological occurrence, though it is not easy to account for it.
On the other hand, in the vine, the axial nature of the tendril is revealed by
the not infrequent presence of flowers or berries on them, as also in
Modecca and some Passifloraceæ.
Darwin, speaking of the tendrils of Bignonia capreolata, says it is a highly
remarkable fact that a leaf should be metamorphosed into a branched organ,
which turns from the light, and which can, by its extremities, either crawl
like a root into crevices, or seize hold of minute projecting points, these
extremities subsequently forming cellular masses, which envelope by their
growth the first fibres and secrete an adhesive cement.
Interrupted growth.—This term is here used in the same sense as in
ordinary descriptive botany, as when an "interruptedly pinnate" leaf is
spoken of. A similar alternation may be observed occasionally as a
teratological occurrence, though it is not easy to account for it.
Page 420
Fig. 175.—Interrupted growth of Radish (from the 'American Agriculturist.')
Page 421
Fig. 176.—Interrupted growth in Apple.
Fig. 175 shows an instance of the kind in a radish, and fig. 176 a similar
deformity in the case of an apple, the dilatation of the flower-stalk below
the ordinary fruit producing an appearance as if there were two fruits one
above another.
In leaves this peculiar irregularity of development is more common.
In some varieties of Codiæum variegatum the leaves resemble those of
Nepenthes, as the basal portion is broad, and terminates in a projecting
midrib destitute of cellular covering, and this again terminates in a small
pouch or pitcher. Somewhat similar variations may be found in ferns,
especially Scolopendrium vulgare.
Instead of the pouch there is formed sometimes in the plant last mentioned a
supplementary four-lobed lamina, the four lobes being in two different
planes, and diverging from the midrib, so that the section would resemble
[symbol: Sideways X], the point of intersection of the x representing the
position of the midrib. This four-winged lamina is thus very similar to the
four-winged filaments described and figured at p. 289, and to the leaf-like
anther of Jatropha described by M. Müller, p. 255.
Cornute leaves (Folia cornuta).—The condition to which this term applies
is that in which the midrib, after running for a certain distance, generally
nearly to the point of the leaf, suddenly projects, often in a plane different
from that of the leaf, and thus forms a small spine-like out-growth. Should
this happen to be terminated by a second laminar portion, an interrupted
leaf would be formed. In Scolopendrium vulgare and other ferns this
condition has been noticed, as also in some of the varieties of Codiæum
variegatum already referred to.
Fig. 175 shows an instance of the kind in a radish, and fig. 176 a similar
deformity in the case of an apple, the dilatation of the flower-stalk below
the ordinary fruit producing an appearance as if there were two fruits one
above another.
In leaves this peculiar irregularity of development is more common.
In some varieties of Codiæum variegatum the leaves resemble those of
Nepenthes, as the basal portion is broad, and terminates in a projecting
midrib destitute of cellular covering, and this again terminates in a small
pouch or pitcher. Somewhat similar variations may be found in ferns,
especially Scolopendrium vulgare.
Instead of the pouch there is formed sometimes in the plant last mentioned a
supplementary four-lobed lamina, the four lobes being in two different
planes, and diverging from the midrib, so that the section would resemble
[symbol: Sideways X], the point of intersection of the x representing the
position of the midrib. This four-winged lamina is thus very similar to the
four-winged filaments described and figured at p. 289, and to the leaf-like
anther of Jatropha described by M. Müller, p. 255.
Cornute leaves (Folia cornuta).—The condition to which this term applies
is that in which the midrib, after running for a certain distance, generally
nearly to the point of the leaf, suddenly projects, often in a plane different
from that of the leaf, and thus forms a small spine-like out-growth. Should
this happen to be terminated by a second laminar portion, an interrupted
leaf would be formed. In Scolopendrium vulgare and other ferns this
condition has been noticed, as also in some of the varieties of Codiæum
variegatum already referred to.
Page 422
Flattening.—There are some plants whose stem or branches, instead of
assuming the ordinary cylindrical form, are compressed or flattened; such
are some species of Epiphyllum, Coccoloba, Bauhinia, &c. The same thing
occurs in the leaf-like branches of Ruscus, the flower-stalks of Xylophylla,
Phyllanthus, Pterisanthes. Martins proposes to apply the word 'cladodium'
to such expansions, just as the term phyllodium is applied to the similar
dilatation of the leaf-stalks. If we exclude instances of fasciation, i.e. where
several branches are fused together and flattened, we must admit that this
flattening does not occur very often as a teratological appearance.
Mr. Rennie figures and describes a root of a tree which had become greatly
flattened in its passage between the stones at the bottom of a stream, and
had become, as it were, moulded to the stones with which it came into
contact.[364]
The spadix of Arum, as also of the cocoa-nut palm, has been observed
flattened out, apparently without increase in the number of organs.
When the blade of the leaf is suppressed it often happens that the stalk of
the leaf is flattened, as it were, by compensation, and the petiole has then
much the appearance of a flat ribbon (phyllode). This happens constantly in
certain species of Acacia, Oxalis, &c., and has been attributed, but
doubtless erroneously, to the fusion of the leaflets in an early state of
development and in the position of rest.[365]
In some water plants, as Sagittaria, Alisma, Potamogeton, &c., the leaf-
stalks are apt to get flattened out into ribbon-like bodies; and Olivier has
figured and described a Cyclamen, called by him C. linearifolium, in which,
owing to the suppression of the lamina, the petiole had become dilated into
a ribbon-like expansion—déformation rubanée of Moquin.
FOOTNOTES:
assuming the ordinary cylindrical form, are compressed or flattened; such
are some species of Epiphyllum, Coccoloba, Bauhinia, &c. The same thing
occurs in the leaf-like branches of Ruscus, the flower-stalks of Xylophylla,
Phyllanthus, Pterisanthes. Martins proposes to apply the word 'cladodium'
to such expansions, just as the term phyllodium is applied to the similar
dilatation of the leaf-stalks. If we exclude instances of fasciation, i.e. where
several branches are fused together and flattened, we must admit that this
flattening does not occur very often as a teratological appearance.
Mr. Rennie figures and describes a root of a tree which had become greatly
flattened in its passage between the stones at the bottom of a stream, and
had become, as it were, moulded to the stones with which it came into
contact.[364]
The spadix of Arum, as also of the cocoa-nut palm, has been observed
flattened out, apparently without increase in the number of organs.
When the blade of the leaf is suppressed it often happens that the stalk of
the leaf is flattened, as it were, by compensation, and the petiole has then
much the appearance of a flat ribbon (phyllode). This happens constantly in
certain species of Acacia, Oxalis, &c., and has been attributed, but
doubtless erroneously, to the fusion of the leaflets in an early state of
development and in the position of rest.[365]
In some water plants, as Sagittaria, Alisma, Potamogeton, &c., the leaf-
stalks are apt to get flattened out into ribbon-like bodies; and Olivier has
figured and described a Cyclamen, called by him C. linearifolium, in which,
owing to the suppression of the lamina, the petiole had become dilated into
a ribbon-like expansion—déformation rubanée of Moquin.
FOOTNOTES:
Page 423
[350] Moore, 'Nature Printed Ferns,' 8vo edition, vol. ii, p. 154, et p. 173.
[351] 'Flora (B. Z.),' 1821, vol. iv, p. 717, c. tab.
[352] Chavannes, 'Mon. Antirrh.'
[353] 'Bull. Soc. Bot. France,' t. vii, 1860, p. 877.
[354] Ibid., t. iv, 1857, p. 759.
[355] Jaeger, "De monstrosa folii Phœnicis dactyliferæ conformatione a Goetheo olim
observata," 'Act. Acad. Leop. Car. Nat. Cur.,' vol. xvii, suppl., p. 293, c. tab. color. iv.
[356] See Goethe, 'Ueber die spiral Tendenz.'
[357] See Darwin "On Climbing Plants," 'Journ. Linn. Soc. Botany,' vol. ix, p. 5.
[358] 'Ephem. Nat. Cur.,' dec. 2, ann. 1, 1683, p. 68, fig. 14.
[359] 'Ann. des Scienc. Nat.,' third series, vol. i, 1844, p. 292.
[360] 'Flora' Feb. 4, 1858, p. 69, tab. ii, f. 3, and also 'Flora,' 1860, p. 737, tab. vii, f. 9.
[361] 'Bull. Acad, Belg.,' t. xvii, p. 196, "Lobelia," p. 53, c. tab.
[362] Moore, 'Nature-printed Ferns,' 8vo edition, vol. ii, p. 183.
[363] 'Bull. Soc. Bot. Fr.,' 1860, vol. vii, p. 461. See also Naudin, 'Ann. Sc. Nat.,' 4 ser., t. iv,
p. 5. Clos, 'Bull. Soc. Bot. Fr.,' t. iii, p. 546.
[364] London's 'Magazine Nat. Hist.,' vol. ii, p. 463.
[365] C. Morren, 'Bull. Acad. Belg.,' 1852, t. xix, part iii, p. 444.
CHAPTER II.
POLYMORPHY.
Usually the several organs of the same individual plant do not differ to any
great extent one from another. One adult leaf has nearly the same
appearance and dimensions as another; one flower resembles very closely
[351] 'Flora (B. Z.),' 1821, vol. iv, p. 717, c. tab.
[352] Chavannes, 'Mon. Antirrh.'
[353] 'Bull. Soc. Bot. France,' t. vii, 1860, p. 877.
[354] Ibid., t. iv, 1857, p. 759.
[355] Jaeger, "De monstrosa folii Phœnicis dactyliferæ conformatione a Goetheo olim
observata," 'Act. Acad. Leop. Car. Nat. Cur.,' vol. xvii, suppl., p. 293, c. tab. color. iv.
[356] See Goethe, 'Ueber die spiral Tendenz.'
[357] See Darwin "On Climbing Plants," 'Journ. Linn. Soc. Botany,' vol. ix, p. 5.
[358] 'Ephem. Nat. Cur.,' dec. 2, ann. 1, 1683, p. 68, fig. 14.
[359] 'Ann. des Scienc. Nat.,' third series, vol. i, 1844, p. 292.
[360] 'Flora' Feb. 4, 1858, p. 69, tab. ii, f. 3, and also 'Flora,' 1860, p. 737, tab. vii, f. 9.
[361] 'Bull. Acad, Belg.,' t. xvii, p. 196, "Lobelia," p. 53, c. tab.
[362] Moore, 'Nature-printed Ferns,' 8vo edition, vol. ii, p. 183.
[363] 'Bull. Soc. Bot. Fr.,' 1860, vol. vii, p. 461. See also Naudin, 'Ann. Sc. Nat.,' 4 ser., t. iv,
p. 5. Clos, 'Bull. Soc. Bot. Fr.,' t. iii, p. 546.
[364] London's 'Magazine Nat. Hist.,' vol. ii, p. 463.
[365] C. Morren, 'Bull. Acad. Belg.,' 1852, t. xix, part iii, p. 444.
CHAPTER II.
POLYMORPHY.
Usually the several organs of the same individual plant do not differ to any
great extent one from another. One adult leaf has nearly the same
appearance and dimensions as another; one flower resembles very closely
Page 424
another flower of the same age and so on. Nevertheless it occasionally
happens that there is a very considerable difference in form in the same
organs, not only at different times, but it may also be at the same time.
Descriptive botanists recognise this occurrence in the case of leaves, and
apply the epithet heterophyllous to plants possessed of these variable foliar
characters. In the case of the flower, where similar diversity of form
occasionally exists, the term dimorphism is used.
As these phenomena appear constantly in particular plants, they are hardly
to be looked on, under such circumstances, as abnormal, but where they
occur in plants not usually polymorphic, they may be considered as coming
within the scope of teratology.
Heterophylly.—As a general rule, the leaves or leaf-organs in each portion
of a plant, from the rhizome or underground axis, where it exists, to the
carpellary leaf, have their own special configuration, subject only to slight
variations, dependent upon age, conditions of growth, &c. The cotyledons
are very uniform in shape in each plant, and are scarcely ever subject to
variation. The leaves near the base of the stem, the root-leaves as they are
not unfrequently called, sometimes differ in form from the stem-leaves;
these again differ from the bracts or leaves in proximity to the flower. The
floral envelopes themselves, as well as the bud-scales, all have their own
allotted form in particular plants, a form by which they may, in most cases,
be readily recognised. Hence, then, in the majority of plants there is
naturally very considerable difference in the form of the leaf-organs,
according to the place they occupy and the functions they have to fulfil; but,
in addition to this, it not unfrequently happens that the leaf-organs in the
same portion of the stem are subject to great variation in form. This is the
condition to which the term heterophylly properly applies. The variation in
form is usually dependent on a greater or less degree of lobing of the
margin of the leaf; thus, in the yellow jasmine, almost every intermediate
happens that there is a very considerable difference in form in the same
organs, not only at different times, but it may also be at the same time.
Descriptive botanists recognise this occurrence in the case of leaves, and
apply the epithet heterophyllous to plants possessed of these variable foliar
characters. In the case of the flower, where similar diversity of form
occasionally exists, the term dimorphism is used.
As these phenomena appear constantly in particular plants, they are hardly
to be looked on, under such circumstances, as abnormal, but where they
occur in plants not usually polymorphic, they may be considered as coming
within the scope of teratology.
Heterophylly.—As a general rule, the leaves or leaf-organs in each portion
of a plant, from the rhizome or underground axis, where it exists, to the
carpellary leaf, have their own special configuration, subject only to slight
variations, dependent upon age, conditions of growth, &c. The cotyledons
are very uniform in shape in each plant, and are scarcely ever subject to
variation. The leaves near the base of the stem, the root-leaves as they are
not unfrequently called, sometimes differ in form from the stem-leaves;
these again differ from the bracts or leaves in proximity to the flower. The
floral envelopes themselves, as well as the bud-scales, all have their own
allotted form in particular plants, a form by which they may, in most cases,
be readily recognised. Hence, then, in the majority of plants there is
naturally very considerable difference in the form of the leaf-organs,
according to the place they occupy and the functions they have to fulfil; but,
in addition to this, it not unfrequently happens that the leaf-organs in the
same portion of the stem are subject to great variation in form. This is the
condition to which the term heterophylly properly applies. The variation in
form is usually dependent on a greater or less degree of lobing of the
margin of the leaf; thus, in the yellow jasmine, almost every intermediate
Page 425
stage may be traced from an ovate entire leaf to one very deeply and
irregularly stalked. Broussonettia papyrifera, and Laurus Sassafras, and the
species of Panax, may be mentioned as presenting this condition.
Sometimes in the last-named genus, as also in Pteridophyllum, every
gradation between simple and compound leaves may be traced. The horse-
radish (Cochlearia Armoracia) may also be instanced as a common
illustration of polymorphism in the leaves. In ferns it is likewise of frequent
occurrence, markedly so in Scolopendrium D'Urvillei, in which plant every
gradation from a simple oblong frond to an exceedingly divided one may be
found springing from the same rhizome at the same time.
Fig. 177.—Syringa persica laciniata, showing polymorphous leaves.
A similar protean state, but little less remarkable, occurs in many of our
British ferns, notably in Scolopendrium vulgare, of which Mr. Moore
enumerates no fewer than 155 varieties,[366] many of the forms occurring on
irregularly stalked. Broussonettia papyrifera, and Laurus Sassafras, and the
species of Panax, may be mentioned as presenting this condition.
Sometimes in the last-named genus, as also in Pteridophyllum, every
gradation between simple and compound leaves may be traced. The horse-
radish (Cochlearia Armoracia) may also be instanced as a common
illustration of polymorphism in the leaves. In ferns it is likewise of frequent
occurrence, markedly so in Scolopendrium D'Urvillei, in which plant every
gradation from a simple oblong frond to an exceedingly divided one may be
found springing from the same rhizome at the same time.
Fig. 177.—Syringa persica laciniata, showing polymorphous leaves.
A similar protean state, but little less remarkable, occurs in many of our
British ferns, notably in Scolopendrium vulgare, of which Mr. Moore
enumerates no fewer than 155 varieties,[366] many of the forms occurring on
Page 426
the same plant at the same time. Cultivators have availed themselves of this
tendency to produce multiform foliage, not only for the purposes of
decoration or curiosity, as in the many cut-leaved or crisped-leaved
varieties, but also for more material uses, as, for instance, the many
varieties of cabbages, of lettuces, &c. Most of these variations are
mentioned under the head of the particular morphological change of which
they are illustrations.
The effect of a change in the conditions of growth in producing diversity in
the form of the leaf may be here alluded to. Ficus stipulata, a plant used to
cover the walls of plant-stoves in this country, and growing naturally on
walls in India, like ivy, produces leaves of very different form, size, and
texture, when grown as a standard, from what it does when adhering to a
wall. Marcgraavia umbellata furnishes another example of a similar nature,
as indeed, to a less extent, does the common ivy.
Allusion has been already made to the occasional persistence of forms in
adult life, which are commonly confined to a young state, as in the case of
some conifers which present on the same plant, at the same time, two
different forms of leaves. Mention has also been made of the presence of
adventitious buds on leaves and in other situations. The leaves that spring
from these buds are usually of the same form as the other leaves of the
plant, but now and then they differ. Of this a remarkable illustration is
afforded by a fern, Pteris quadriaurita, in which the fronds emerging from
an adventitious bud are very different from the ordinary fronds.
tendency to produce multiform foliage, not only for the purposes of
decoration or curiosity, as in the many cut-leaved or crisped-leaved
varieties, but also for more material uses, as, for instance, the many
varieties of cabbages, of lettuces, &c. Most of these variations are
mentioned under the head of the particular morphological change of which
they are illustrations.
The effect of a change in the conditions of growth in producing diversity in
the form of the leaf may be here alluded to. Ficus stipulata, a plant used to
cover the walls of plant-stoves in this country, and growing naturally on
walls in India, like ivy, produces leaves of very different form, size, and
texture, when grown as a standard, from what it does when adhering to a
wall. Marcgraavia umbellata furnishes another example of a similar nature,
as indeed, to a less extent, does the common ivy.
Allusion has been already made to the occasional persistence of forms in
adult life, which are commonly confined to a young state, as in the case of
some conifers which present on the same plant, at the same time, two
different forms of leaves. Mention has also been made of the presence of
adventitious buds on leaves and in other situations. The leaves that spring
from these buds are usually of the same form as the other leaves of the
plant, but now and then they differ. Of this a remarkable illustration is
afforded by a fern, Pteris quadriaurita, in which the fronds emerging from
an adventitious bud are very different from the ordinary fronds.
Page 427
Fig. 178.—Portion of a frond of Pteris quadriaurita, with an adventitious bud, the form of the
constituent foliage of which is very different from that of the parent frond.
Dimorphism.—This term, applied specially to the varied form which the
flowers or some of their constituent elements assume on the same plant, is
an analogous phenomenon to what has been above spoken of as
heterophylly, and, like it, it cannot, except under special circumstances, be
considered as of teratological importance. A few illustrative cases, however,
may here be cited.
Sir George Mackenzie describes a variety of the potato[367] (Solanum
tuberosum), which produces first double and sterile flowers, and
subsequently single fertile ones; the other portions of the plant do not differ
much.
Stackhousia juncea, according to Clarke, has mixed with its perfect flowers
a number of apetalous blossoms destitute of anthers.[368]
constituent foliage of which is very different from that of the parent frond.
Dimorphism.—This term, applied specially to the varied form which the
flowers or some of their constituent elements assume on the same plant, is
an analogous phenomenon to what has been above spoken of as
heterophylly, and, like it, it cannot, except under special circumstances, be
considered as of teratological importance. A few illustrative cases, however,
may here be cited.
Sir George Mackenzie describes a variety of the potato[367] (Solanum
tuberosum), which produces first double and sterile flowers, and
subsequently single fertile ones; the other portions of the plant do not differ
much.
Stackhousia juncea, according to Clarke, has mixed with its perfect flowers
a number of apetalous blossoms destitute of anthers.[368]
Page 428
This peculiarity is well exemplified in the tribe Gaudichaudieæ of the order
Malpighiaceæ. A. de Jussieu, in his monograph, speaks of these flowers as
being very small, green, destitute of petals, or nearly so, with a single,
generally imperfect anther; the carpels also are more or less imperfect, but
not sufficiently so to prevent some seeds from being formed. A similar
production of imperfect flowers has been noticed in many other orders, e.g.
Violaceæ, Campanulaceæ, &c. In some cases these supplementary
blossoms are more fertile and prolific in good seeds than are the normally
constructed flowers. M. Durieu de Maisonneuve alludes to a case where
flowers of this description are produced below the surface of the ground.
The plant in question is Scrophularia arguta, and it appears that towards the
end of the summer the lowest branches springing from the stem bend
downwards, and penetrate the soil; the branches immediately above the
lowest ones also bend downwards, but do not always enter the earth. These
branches bear fertile flowers: those which are completely below the soil are
completely destitute of petals; those which are on the surface have a four-
lobed corolla whose divisions are nearly equal, like those of Veronica.[369]
To Sprengel, and specially to Darwin, physiologists are indebted for the
demonstration of the relation of di- and trimorphic flowers to fertilisation.
In certain genera of orchids, such as Catasetum, &c., flowers of such
different form are produced that botanists, without hesitation, considered
them as belonging to different genera, until the fact of their occasional
production on the same plant showed that they were not of even specific
importance. It was reserved for Mr. Darwin to show experimentally that
these very different flowers are really sexual forms of one and the same
species, ordinarily occurring on different plants, i.e. diœcious, but
occasionally formed on the same spike. The same excellent observer has
demonstrated that the di- and trimorphic forms of Primula, of Linum,
Lythrum, and other plants—forms differing mainly in the relative length of
the stamens and styles, are also connected with striking differences in the
Malpighiaceæ. A. de Jussieu, in his monograph, speaks of these flowers as
being very small, green, destitute of petals, or nearly so, with a single,
generally imperfect anther; the carpels also are more or less imperfect, but
not sufficiently so to prevent some seeds from being formed. A similar
production of imperfect flowers has been noticed in many other orders, e.g.
Violaceæ, Campanulaceæ, &c. In some cases these supplementary
blossoms are more fertile and prolific in good seeds than are the normally
constructed flowers. M. Durieu de Maisonneuve alludes to a case where
flowers of this description are produced below the surface of the ground.
The plant in question is Scrophularia arguta, and it appears that towards the
end of the summer the lowest branches springing from the stem bend
downwards, and penetrate the soil; the branches immediately above the
lowest ones also bend downwards, but do not always enter the earth. These
branches bear fertile flowers: those which are completely below the soil are
completely destitute of petals; those which are on the surface have a four-
lobed corolla whose divisions are nearly equal, like those of Veronica.[369]
To Sprengel, and specially to Darwin, physiologists are indebted for the
demonstration of the relation of di- and trimorphic flowers to fertilisation.
In certain genera of orchids, such as Catasetum, &c., flowers of such
different form are produced that botanists, without hesitation, considered
them as belonging to different genera, until the fact of their occasional
production on the same plant showed that they were not of even specific
importance. It was reserved for Mr. Darwin to show experimentally that
these very different flowers are really sexual forms of one and the same
species, ordinarily occurring on different plants, i.e. diœcious, but
occasionally formed on the same spike. The same excellent observer has
demonstrated that the di- and trimorphic forms of Primula, of Linum,
Lythrum, and other plants—forms differing mainly in the relative length of
the stamens and styles, are also connected with striking differences in the
Page 429
number of perfect seeds produced. The most perfect degree of fertility is
obtained when the stigma of one form is fertilised by the pollen taken from
stamens of a corresponding height. On the other hand, when the union is, as
Mr. Darwin states, illegitimate, that is, when the pollen is taken from
stamens not corresponding in length to the style, more or less complete
sterility ensues in the progeny, sometimes even utter infertility, such as
happens when two distinct species are crossed, so that, in point of fact, the
offspring of these illegitimate unions correspond almost precisely to
hybrids.[370]
Mere variations of form arising from hybridisation or other causes hardly
fall within the limits of this work, though it is quite impossible to say where
variations end and malformations begin. There are, however, two or three
cases cited by Mr. Darwin[371] from Gallesio and Risso to which it is
desirable to allude. Gallesio impregnated an orange with pollen from a
lemon, and the fruit borne on the mother tree had a raised stripe of peel like
that of a lemon both in colour and taste, but the pulp was like that of an
orange, and included only imperfect seeds. Risso describes a variety of the
common orange which produces "rounded-oval leaves, spotted with yellow,
borne on petioles, with heart-shaped wings; when these leaves fall off they
are succeeded by longer and narrower leaves, with undulated margins, of a
pale green colour, embroidered with yellow, borne on foot-stalks without
wings. The fruit whilst young is pear-shaped, yellow, longitudinally striated
and sweet; but, as it ripens, it becomes spherical, of a reddish-yellow, and
bitter."
Sports or bud variations.—These curious departures from the normal
form can only be mentioned incidentally in this place, as they pertain more
to variation than to malformation.
obtained when the stigma of one form is fertilised by the pollen taken from
stamens of a corresponding height. On the other hand, when the union is, as
Mr. Darwin states, illegitimate, that is, when the pollen is taken from
stamens not corresponding in length to the style, more or less complete
sterility ensues in the progeny, sometimes even utter infertility, such as
happens when two distinct species are crossed, so that, in point of fact, the
offspring of these illegitimate unions correspond almost precisely to
hybrids.[370]
Mere variations of form arising from hybridisation or other causes hardly
fall within the limits of this work, though it is quite impossible to say where
variations end and malformations begin. There are, however, two or three
cases cited by Mr. Darwin[371] from Gallesio and Risso to which it is
desirable to allude. Gallesio impregnated an orange with pollen from a
lemon, and the fruit borne on the mother tree had a raised stripe of peel like
that of a lemon both in colour and taste, but the pulp was like that of an
orange, and included only imperfect seeds. Risso describes a variety of the
common orange which produces "rounded-oval leaves, spotted with yellow,
borne on petioles, with heart-shaped wings; when these leaves fall off they
are succeeded by longer and narrower leaves, with undulated margins, of a
pale green colour, embroidered with yellow, borne on foot-stalks without
wings. The fruit whilst young is pear-shaped, yellow, longitudinally striated
and sweet; but, as it ripens, it becomes spherical, of a reddish-yellow, and
bitter."
Sports or bud variations.—These curious departures from the normal
form can only be mentioned incidentally in this place, as they pertain more
to variation than to malformation.
Page 430
The occasional production of shoots bearing leaves, flowers, or fruits of a
different character from those found on the normal plant, is a fact of which
gardeners have largely availed themselves in the cultivation of new
varieties. The productions in question have been attributed to various
causes, such as cross-breeding, grafting, budding, dissociation of hybrid
characters, or reversion to some ancestral form, all of which explanations
may be true in certain cases, but none of them supply the clue to the reason
why one particular branch should be so affected, and the rest not; or why
the same plant, at the same time, as often happens in Pelargoniums, should
produce two, three, or more "sports" of a different character.
These bud variations may be perpetuated by grafts or by cuttings,
sometimes even by seed. With reference to cuttings a curious circumstance
has been observed, viz., that if taken from the lower part of the stem, near
the root, the peculiarity is not transmitted, but the young plant reverts to the
characters of the typical form (Carrière). This circumstance, however, is not
of universal occurrence.
For further particulars on this interesting subject the reader is referred to
Darwin's 'Variation of Animals and Plants,' i, p. 373, where numerous
references are given, and wherein certain well-known and highly
remarkable instances, such as the Cytisus Adami, the trifacial orange, &c.,
are discussed.
FOOTNOTES:
[366] 'Nature-printed Ferns,' 8vo edition, vol. ii, p. 197.
[367] 'Gard. Chron.,' 1845. p. 790.
[368] 'A New Arrangement of Phænog. Plants,' p. 36.
[369] 'Bull. Soc. Bot. France,' 1856, t. iii, p. 569.
different character from those found on the normal plant, is a fact of which
gardeners have largely availed themselves in the cultivation of new
varieties. The productions in question have been attributed to various
causes, such as cross-breeding, grafting, budding, dissociation of hybrid
characters, or reversion to some ancestral form, all of which explanations
may be true in certain cases, but none of them supply the clue to the reason
why one particular branch should be so affected, and the rest not; or why
the same plant, at the same time, as often happens in Pelargoniums, should
produce two, three, or more "sports" of a different character.
These bud variations may be perpetuated by grafts or by cuttings,
sometimes even by seed. With reference to cuttings a curious circumstance
has been observed, viz., that if taken from the lower part of the stem, near
the root, the peculiarity is not transmitted, but the young plant reverts to the
characters of the typical form (Carrière). This circumstance, however, is not
of universal occurrence.
For further particulars on this interesting subject the reader is referred to
Darwin's 'Variation of Animals and Plants,' i, p. 373, where numerous
references are given, and wherein certain well-known and highly
remarkable instances, such as the Cytisus Adami, the trifacial orange, &c.,
are discussed.
FOOTNOTES:
[366] 'Nature-printed Ferns,' 8vo edition, vol. ii, p. 197.
[367] 'Gard. Chron.,' 1845. p. 790.
[368] 'A New Arrangement of Phænog. Plants,' p. 36.
[369] 'Bull. Soc. Bot. France,' 1856, t. iii, p. 569.
Page 431
[370] The reader will find an abstract of Mr. Darwin's views in his work on the 'Variation of
Animals and Plants,' vol. ii, p. 181.
[371] Loc. cit., i, 336.
CHAPTER III.
ALTERATIONS OF COLOUR.[372]
Changes in the colour of the several organs of plants are more often either
pathological or the result of variation than of malformation properly so
called.
Alterations in colour arise from a diminished or an increased amount of
colouring matter, or from an unusual distribution of the solid or fluid
matters on which the colour depends. The superposition of cells containing
colouring material of different tints produces naturally a very different set
of hues from those which are manifested when the colours are not blended.
Referring the reader to the ordinary text-books on vegetable physiology and
chemistry for details as to the nature and disposition of colouring materials
in plants under natural circumstances, it will only be necessary to cite a few
instances of deviation from the general colour of plants or their organs.
Albinism.—This change is due to the deficient formation of green
colouring matter or chlorophyll, and is more a pathological condition than a
deformity.
It seems necessary to draw a distinction between this state and ordinary
blanching or etiolation. In the former case chlorophyll seems never to be
formed in the affected parts, even if they be exposed to light, while an
etiolated organ, when placed under favorable circumstances, speedily
Animals and Plants,' vol. ii, p. 181.
[371] Loc. cit., i, 336.
CHAPTER III.
ALTERATIONS OF COLOUR.[372]
Changes in the colour of the several organs of plants are more often either
pathological or the result of variation than of malformation properly so
called.
Alterations in colour arise from a diminished or an increased amount of
colouring matter, or from an unusual distribution of the solid or fluid
matters on which the colour depends. The superposition of cells containing
colouring material of different tints produces naturally a very different set
of hues from those which are manifested when the colours are not blended.
Referring the reader to the ordinary text-books on vegetable physiology and
chemistry for details as to the nature and disposition of colouring materials
in plants under natural circumstances, it will only be necessary to cite a few
instances of deviation from the general colour of plants or their organs.
Albinism.—This change is due to the deficient formation of green
colouring matter or chlorophyll, and is more a pathological condition than a
deformity.
It seems necessary to draw a distinction between this state and ordinary
blanching or etiolation. In the former case chlorophyll seems never to be
formed in the affected parts, even if they be exposed to light, while an
etiolated organ, when placed under favorable circumstances, speedily
Page 432
assumes a green colour. In Richardia æthiopica one or more leaves become
occasionally as white as the spathe is usually.
Virescence.—Engelmann[373] pointed out that, so far as flowers were
concerned, there are two ways in which they assume a green colour, either
by a simple development of chlorophyll in place of the colouring matter
proper to the flower, or by an actual development of leaf-like organs in the
room of the petals—frondescence. Morren[374] judiciously proposed to keep
these two conditions separate, calling the one virescence, the other
frondescence (see p. 241).
Many of the cases recorded as reversions of the parts of the flower to leaves
are simply instances of virescence; indeed, it is not in all cases easy to
distinguish between the two states. The examination of the arrangement of
the veins is often of assistance in determining this point; for instance, if,
under ordinary circumstances, the venation of the petal be such as is
characteristic of the sheath of the leaf, while in the green-coloured flower of
the same species the venation is more like that which belongs to the blade
of the leaf, the inference would, of course, be that the green colour was due
to frondescence or phyllody.
The persistence or duration of petals is often increased when they are
subject to this change; instead of falling off speedily they become persistent
when so affected.
Some flowers are more liable to virescence than others. The common
honeysuckle, Lonicera Periclymenum, is one of these, and it is noticeable in
this plant that the calyx remains unaffected—a circumstance which Morren
says shows the distinctness of virescence from frondescence; for, in this
instance, we have the most foliaceous portion of the flower remaining
unchanged, while the corolla and other organs, usually less leaf-like in their
occasionally as white as the spathe is usually.
Virescence.—Engelmann[373] pointed out that, so far as flowers were
concerned, there are two ways in which they assume a green colour, either
by a simple development of chlorophyll in place of the colouring matter
proper to the flower, or by an actual development of leaf-like organs in the
room of the petals—frondescence. Morren[374] judiciously proposed to keep
these two conditions separate, calling the one virescence, the other
frondescence (see p. 241).
Many of the cases recorded as reversions of the parts of the flower to leaves
are simply instances of virescence; indeed, it is not in all cases easy to
distinguish between the two states. The examination of the arrangement of
the veins is often of assistance in determining this point; for instance, if,
under ordinary circumstances, the venation of the petal be such as is
characteristic of the sheath of the leaf, while in the green-coloured flower of
the same species the venation is more like that which belongs to the blade
of the leaf, the inference would, of course, be that the green colour was due
to frondescence or phyllody.
The persistence or duration of petals is often increased when they are
subject to this change; instead of falling off speedily they become persistent
when so affected.
Some flowers are more liable to virescence than others. The common
honeysuckle, Lonicera Periclymenum, is one of these, and it is noticeable in
this plant that the calyx remains unaffected—a circumstance which Morren
says shows the distinctness of virescence from frondescence; for, in this
instance, we have the most foliaceous portion of the flower remaining
unchanged, while the corolla and other organs, usually less leaf-like in their
Page 433
nature, assume a green colour; but this may rather be attributed to the axial
nature of the so-called adherent calyx. The stamens in these green-flowered
honeysuckles are usually green also, but with abortive anthers, and the pistil
also is in a rudimentary condition. Umbelliferæ are not unfrequently subject
to this change, e.g., Torilis Anthriscus, Daucus Carota, Heracleum
Sphondylium, Carum carui, &c. Primulaceæ, again, are frequently subject
to virescence. Among Compositæ the following species are recorded as
having had green flowers—Cirsium tricephalodes, Senecio vulgaris,
Calendula officinalis, Pyrethrum Parthenium, Carduus crispus, Hypochæris
radicata, Hieracium prealtum, Cirsium arvense, Coreopsis Drummondi.
[375] In Ranunculaceæ virescence has been observed in Delphinium elatum,
crassicaule and Ajacis, Anemone hortensis and nemorosa, Aquilegia
vulgaris, Ranunculus Philonotis.
Many of these cases, and others that might be cited, are probably instances
of frondescence or phyllody (see p. 241).
Chromatism.—This term is here intended to apply specially to those cases
in which any organ of a plant assumes a colour approximating to that of the
petals, or in which the normal green is replaced by tints of some other
colour. To a certain extent the change in question is the same as that spoken
of under the head of petalody (see p. 283), but there are cases in which,
while the ordinary situation and form are those of leaves, the coloration is
that of the petals. Such was the case in the Gesnera mentioned by Morren
(see p. 88), and in which a leaf occupied the position of an inflorescence,
and became brightly coloured. In tulips the presence of a highly coloured
leaf on the flower-stalk, below the flower, is not uncommon. So also the
bracts or leaves below the perianth in Anemone coronaria and hortensis not
unfrequently assume the coloration usually confined to the parts of the
perianth. A similar illustration has presented itself, as this sheet is passing
through the press, in which two of the leaflets of the compound leaf of a
nature of the so-called adherent calyx. The stamens in these green-flowered
honeysuckles are usually green also, but with abortive anthers, and the pistil
also is in a rudimentary condition. Umbelliferæ are not unfrequently subject
to this change, e.g., Torilis Anthriscus, Daucus Carota, Heracleum
Sphondylium, Carum carui, &c. Primulaceæ, again, are frequently subject
to virescence. Among Compositæ the following species are recorded as
having had green flowers—Cirsium tricephalodes, Senecio vulgaris,
Calendula officinalis, Pyrethrum Parthenium, Carduus crispus, Hypochæris
radicata, Hieracium prealtum, Cirsium arvense, Coreopsis Drummondi.
[375] In Ranunculaceæ virescence has been observed in Delphinium elatum,
crassicaule and Ajacis, Anemone hortensis and nemorosa, Aquilegia
vulgaris, Ranunculus Philonotis.
Many of these cases, and others that might be cited, are probably instances
of frondescence or phyllody (see p. 241).
Chromatism.—This term is here intended to apply specially to those cases
in which any organ of a plant assumes a colour approximating to that of the
petals, or in which the normal green is replaced by tints of some other
colour. To a certain extent the change in question is the same as that spoken
of under the head of petalody (see p. 283), but there are cases in which,
while the ordinary situation and form are those of leaves, the coloration is
that of the petals. Such was the case in the Gesnera mentioned by Morren
(see p. 88), and in which a leaf occupied the position of an inflorescence,
and became brightly coloured. In tulips the presence of a highly coloured
leaf on the flower-stalk, below the flower, is not uncommon. So also the
bracts or leaves below the perianth in Anemone coronaria and hortensis not
unfrequently assume the coloration usually confined to the parts of the
perianth. A similar illustration has presented itself, as this sheet is passing
through the press, in which two of the leaflets of the compound leaf of a
Page 434
rose were brightly coloured like the petals, the others being of their ordinary
green colour.
The occurrence of coloured bracts, as in Poinsettia, Bougainvillea, &c., is
very common under natural conditions, and need not here be further alluded
to.
Increased intensity of colour often accompanies teratological changes; an
instance has just been alluded to in the Gesnera; the feather hyacinth,
Muscari comosum, furnishes another illustration, the adventitious pedicels
being brightly coloured.
In fasciated stems, also, of herbaceous plants, it not unfrequently happens
that the upper portions of the stem are brightly coloured.
The occurrence of flowers or fruits of different colours on the same plant, or
even in the same cluster, is a phenomenon which does not come within the
scope of the present book; the reader may, however, be referred to the
excellent summary on this subject published by Mr. Darwin in his work on
the 'Variation of Animals and Plants under Domestication.'
FOOTNOTES:
[372] These deviations are treated of under the head of alterations of form, because they are
not, in a teratological point of view, of sufficient importance to demand a specific heading,
while they appeal to the sight in the same way as the deviations from the customary forms of
organs.
[373] 'De Antholys,' p. 32, § 38.
[374] 'Bull. Acad. Belg.,' xvii, part 2, p. 131, c. tab.
[375] See Cramer, 'Bildungsabweich,' pp. 17, 55, 82, 65. See also Lucas, 'Verhandl. des Bot.
Vereins. Brandenb.,' heft 1, 2, Anchusa. Christ, 'Flora,' 1867. pp. 376, tab. 5, 6, Stachys.
green colour.
The occurrence of coloured bracts, as in Poinsettia, Bougainvillea, &c., is
very common under natural conditions, and need not here be further alluded
to.
Increased intensity of colour often accompanies teratological changes; an
instance has just been alluded to in the Gesnera; the feather hyacinth,
Muscari comosum, furnishes another illustration, the adventitious pedicels
being brightly coloured.
In fasciated stems, also, of herbaceous plants, it not unfrequently happens
that the upper portions of the stem are brightly coloured.
The occurrence of flowers or fruits of different colours on the same plant, or
even in the same cluster, is a phenomenon which does not come within the
scope of the present book; the reader may, however, be referred to the
excellent summary on this subject published by Mr. Darwin in his work on
the 'Variation of Animals and Plants under Domestication.'
FOOTNOTES:
[372] These deviations are treated of under the head of alterations of form, because they are
not, in a teratological point of view, of sufficient importance to demand a specific heading,
while they appeal to the sight in the same way as the deviations from the customary forms of
organs.
[373] 'De Antholys,' p. 32, § 38.
[374] 'Bull. Acad. Belg.,' xvii, part 2, p. 131, c. tab.
[375] See Cramer, 'Bildungsabweich,' pp. 17, 55, 82, 65. See also Lucas, 'Verhandl. des Bot.
Vereins. Brandenb.,' heft 1, 2, Anchusa. Christ, 'Flora,' 1867. pp. 376, tab. 5, 6, Stachys.
Page 435
BOOK III.
DEVIATIONS FROM THE ORDINARY
NUMBER OF ORGANS.
To a certain extent the number of the organs of a plant is of even greater
consequence for purposes of classification than either their form or their
arrangement; for instance, the number of cotyledons in the embryo is made
the chief basis of separation between the two great groups of flowering
plants, the monocotyledons and the dicotyledons. In the one group,
moreover, the parts of the flower are arranged in groups or whorls of five;
in the other the arrangement is ternary. In mosses the teeth of the peristome
are arranged in fours, or in some multiple of that number. So far as the
larger groups are concerned, and also in cases where the actual number of
parts is small, the numerical relations above described are very constant; on
the other hand, in the minor subdivisions, and especially where the absolute
number of parts is large, considerable variation may occur, so that
descriptive botanists frequently make use of the term indefinite, and apply it
to cases where the number of parts is large and variable, or, at any rate, not
easy to be estimated.
Considered teratologically, the changes, as regards the number of organs,
are readily grouped into those consequent on a decreased and into those
resulting from an increased development. The alteration may be absolute or
DEVIATIONS FROM THE ORDINARY
NUMBER OF ORGANS.
To a certain extent the number of the organs of a plant is of even greater
consequence for purposes of classification than either their form or their
arrangement; for instance, the number of cotyledons in the embryo is made
the chief basis of separation between the two great groups of flowering
plants, the monocotyledons and the dicotyledons. In the one group,
moreover, the parts of the flower are arranged in groups or whorls of five;
in the other the arrangement is ternary. In mosses the teeth of the peristome
are arranged in fours, or in some multiple of that number. So far as the
larger groups are concerned, and also in cases where the actual number of
parts is small, the numerical relations above described are very constant; on
the other hand, in the minor subdivisions, and especially where the absolute
number of parts is large, considerable variation may occur, so that
descriptive botanists frequently make use of the term indefinite, and apply it
to cases where the number of parts is large and variable, or, at any rate, not
easy to be estimated.
Considered teratologically, the changes, as regards the number of organs,
are readily grouped into those consequent on a decreased and into those
resulting from an increased development. The alteration may be absolute or
Page 436
relative. There may be an actual deficiency in the number of parts or an
increase in their number, but in either case the change may be simply a
restoration of the primitive number, a species of peloria, in fact. An
increased number of parts, moreover, may depend not so much on the
formation of additional parts as on the subdivision of one.
It seems also desirable to treat separately those cases in which there is an
increased number of buds either leaf-buds or flower-buds, as the case may
be, as happens in what is termed prolification. This formation of buds
occurring, as it does, often in unwonted situations is treated of under the
head of alterations of arrangement, the mere increase in number being
considered of subordinate importance as contrasted with the altered
disposition (see p. 100).
PART 1.
INCREASED NUMBER OF ORGANS.
An augmentation in the number of parts may arise from several causes, and
may sometimes be more apparent than real. True multiplication exists
simply as a result of over-development; the affected organs are repeated
sometimes over and over again each in their proper relative position, and
without any transmutation of form.
Metamorphy, on the other hand, often gives rise to the impression that parts
are increased in number, when it may be that the stamens and pistils, one or
both, are not so much increased in number as altered in appearance. The
double anemones and ranunculus of gardens, amongst many other
analogous illustrations, may be mentioned. In these flowers, owing to the
petalody of the stamens and pistils, one or both, an impression of
increase in their number, but in either case the change may be simply a
restoration of the primitive number, a species of peloria, in fact. An
increased number of parts, moreover, may depend not so much on the
formation of additional parts as on the subdivision of one.
It seems also desirable to treat separately those cases in which there is an
increased number of buds either leaf-buds or flower-buds, as the case may
be, as happens in what is termed prolification. This formation of buds
occurring, as it does, often in unwonted situations is treated of under the
head of alterations of arrangement, the mere increase in number being
considered of subordinate importance as contrasted with the altered
disposition (see p. 100).
PART 1.
INCREASED NUMBER OF ORGANS.
An augmentation in the number of parts may arise from several causes, and
may sometimes be more apparent than real. True multiplication exists
simply as a result of over-development; the affected organs are repeated
sometimes over and over again each in their proper relative position, and
without any transmutation of form.
Metamorphy, on the other hand, often gives rise to the impression that parts
are increased in number, when it may be that the stamens and pistils, one or
both, are not so much increased in number as altered in appearance. The
double anemones and ranunculus of gardens, amongst many other
analogous illustrations, may be mentioned. In these flowers, owing to the
petalody of the stamens and pistils, one or both, an impression of
Page 437
exaggerated number is produced, which is by no means necessarily a true
one. Fission or lateral subdivision also gives rise to an apparent increase in
number; thus, some so-called double flowers, the elements of which
appeared to be increased in numbers, owe the appearance merely to the
laciniation or subdivision of their petals.
The French botanists, following Dunal and Moquin, attribute an increase in
the number of whorls in the corolla, and other parts of the flower, to a
process which they call chorisis, and they consider the augmentation to be
due to the splitting of one petal, for instance, into several;—somewhat in
the same manner as one may separate successive layers of talc one from the
other.
English botanists, on the other hand, have been slow to admit any such
process, because, in most instances, no alteration in the law of alternation
takes place in these double flowers, and in those few cases where the law is
apparently infringed, the deviation is explained by the probable suppression
of parts, which were they present would restore the natural arrangement of
the flower; and, that this is no imaginary or purely theoretical explanation,
is shown by some of the Primulaceæ, wherein a second row of stamens is
occasionally present in the adult condition, and renders the floral symmetry
perfect.
The double daffodil, where there are from forty to fifty petaloid organs
instead of fifteen, and wherein each piece exhibits a more or less perfect
coronal lobe at the junction of the claw and the limb, has been cited as an
objection to chorisis, though it is difficult to see on what grounds.
In Delphinium, as shown by Braun,[376] the stamens and carpels are
members of a continuous spiral series, and in the double balsam an extra
corolline whorl is produced, without the suppression of the stamens, in the
one. Fission or lateral subdivision also gives rise to an apparent increase in
number; thus, some so-called double flowers, the elements of which
appeared to be increased in numbers, owe the appearance merely to the
laciniation or subdivision of their petals.
The French botanists, following Dunal and Moquin, attribute an increase in
the number of whorls in the corolla, and other parts of the flower, to a
process which they call chorisis, and they consider the augmentation to be
due to the splitting of one petal, for instance, into several;—somewhat in
the same manner as one may separate successive layers of talc one from the
other.
English botanists, on the other hand, have been slow to admit any such
process, because, in most instances, no alteration in the law of alternation
takes place in these double flowers, and in those few cases where the law is
apparently infringed, the deviation is explained by the probable suppression
of parts, which were they present would restore the natural arrangement of
the flower; and, that this is no imaginary or purely theoretical explanation,
is shown by some of the Primulaceæ, wherein a second row of stamens is
occasionally present in the adult condition, and renders the floral symmetry
perfect.
The double daffodil, where there are from forty to fifty petaloid organs
instead of fifteen, and wherein each piece exhibits a more or less perfect
coronal lobe at the junction of the claw and the limb, has been cited as an
objection to chorisis, though it is difficult to see on what grounds.
In Delphinium, as shown by Braun,[376] the stamens and carpels are
members of a continuous spiral series, and in the double balsam an extra
corolline whorl is produced, without the suppression of the stamens, in the
Page 438
following manner: the ordinary stamens are replaced by petals, the carpels
by stamens, while an additional whorl of carpels is produced at the summit
of the axis. In this instance, therefore, the doubling is distinctly referrible to
an absolute increase in the number of whorls, and not to chorisis.[377]
On the other hand, it must be admitted that there are many cases which are
not to be explained in any other way than that suggested by the French
botanists before alluded to. Probably, the main difficulty in the way of
accepting the doctrine of chorisis is the unfortunate selection of the word
used to designate the process; this naturally suggests a splitting of an organ
already perfectly formed into two or more portions, either in the same plane
as the original organs, "parallel chorisis;" or at right angles to it "collateral
chorisis." Indeed, before so much attention had been paid to the way in
which the floral organs are developed, it was thought that an actual splitting
and dilamination did really take place; Dunal and Moquin both assert as
much. The truth would rather seem to be that, in the so-called parallel
chorisis at least, the process is one of hypertrophy and over-development
rather than of splitting. The adventitious petal or scale is an excrescence or
an outgrowth from the primary organ, and formed subsequently to it.
In the case of "compound stamens" the original stamens are first developed
each from its own cellular "mamelon," or growing point; and, after a time,
other secondary growing points emerge from the primary one, and in this
way the stamens are increased in number, without reference, necessarily, to
the so-called law of alternation. Outgrowths from leaves, multiplying the
laminar surface, are alluded to under the head of hypertrophy, and it is
probable that some of the cases of duplication of the flower, or of the
formation of adventitious segments outside the ordinary corolla as alluded
to in succeeding paragraphs (see Pleiotaxy of the corolla), are due to a
similar process.[378]
by stamens, while an additional whorl of carpels is produced at the summit
of the axis. In this instance, therefore, the doubling is distinctly referrible to
an absolute increase in the number of whorls, and not to chorisis.[377]
On the other hand, it must be admitted that there are many cases which are
not to be explained in any other way than that suggested by the French
botanists before alluded to. Probably, the main difficulty in the way of
accepting the doctrine of chorisis is the unfortunate selection of the word
used to designate the process; this naturally suggests a splitting of an organ
already perfectly formed into two or more portions, either in the same plane
as the original organs, "parallel chorisis;" or at right angles to it "collateral
chorisis." Indeed, before so much attention had been paid to the way in
which the floral organs are developed, it was thought that an actual splitting
and dilamination did really take place; Dunal and Moquin both assert as
much. The truth would rather seem to be that, in the so-called parallel
chorisis at least, the process is one of hypertrophy and over-development
rather than of splitting. The adventitious petal or scale is an excrescence or
an outgrowth from the primary organ, and formed subsequently to it.
In the case of "compound stamens" the original stamens are first developed
each from its own cellular "mamelon," or growing point; and, after a time,
other secondary growing points emerge from the primary one, and in this
way the stamens are increased in number, without reference, necessarily, to
the so-called law of alternation. Outgrowths from leaves, multiplying the
laminar surface, are alluded to under the head of hypertrophy, and it is
probable that some of the cases of duplication of the flower, or of the
formation of adventitious segments outside the ordinary corolla as alluded
to in succeeding paragraphs (see Pleiotaxy of the corolla), are due to a
similar process.[378]
Page 439
The formation of parts in unwonted numbers may be merely a reversion to
what is supposed to have been the original form, and in this way there may
be a restoration of parts that are usually undeveloped or suppressed. There
can be little or no doubt that there are in reality six stamens in Orchidaceæ,
of which one only, under ordinary circumstances, is developed. When the
numerical symmetry is restored, as it sometimes is, it is obvious that the
augmentation that occurs is of a different character from that arising from a
repetition or renewed development of organs. When the increased number
arises from multiplication proper, or from repetition, the ordinary laws of
alternation are not interfered with, but if from chorisis or "dédoublement," it
may happen that the normal arrangement is disturbed.
Without studying the mode of development, it is not in all cases possible to
tell under which of the above categories any particular instance should be
placed; hence, in the following sections, except where otherwise stated, the
cases are grouped according to the appearance presented in the adult
condition, rather than to the way in which the changes from the typical
condition are brought about. With reference to the foliar organs it is
necessary to distinguish those cases in which there is, from any cause, an
augmentation in the number of component parts of a whorl, from those in
which the increase takes place in the numbers of the whorls themselves.
FOOTNOTES:
[376] Braun, 'Pringsheim Jahrbuch f. Wiss. Bot.,' 1858, 1, p. 307, tab. 22, 23.
[377] Henfrey, 'Jour. Linn. Soc. Bot.,' vol. iii, p. 159.
[378] On the subject of chorisis or dédoublement the reader may profitably consult Moquin-
Tandon, 'Ess. sur les Dédoublements,' and the same author in 'Ann. Sc. Nat.,' t. xxvii, p. 236.
and 'El. Ter. Veget.,' p. 337. Dunal, 'Consid. Org. Fleur.,' Montpell., 1829, p. 32, note 3. A. de
St. Hilaire in 'Ann. Sc. Nat.,' ser. 3, t. iii, p. 355, adnot. Lindley, 'Elements of Botany,' p. 76.
Asa Gray. 'Botanical Text Book.'
what is supposed to have been the original form, and in this way there may
be a restoration of parts that are usually undeveloped or suppressed. There
can be little or no doubt that there are in reality six stamens in Orchidaceæ,
of which one only, under ordinary circumstances, is developed. When the
numerical symmetry is restored, as it sometimes is, it is obvious that the
augmentation that occurs is of a different character from that arising from a
repetition or renewed development of organs. When the increased number
arises from multiplication proper, or from repetition, the ordinary laws of
alternation are not interfered with, but if from chorisis or "dédoublement," it
may happen that the normal arrangement is disturbed.
Without studying the mode of development, it is not in all cases possible to
tell under which of the above categories any particular instance should be
placed; hence, in the following sections, except where otherwise stated, the
cases are grouped according to the appearance presented in the adult
condition, rather than to the way in which the changes from the typical
condition are brought about. With reference to the foliar organs it is
necessary to distinguish those cases in which there is, from any cause, an
augmentation in the number of component parts of a whorl, from those in
which the increase takes place in the numbers of the whorls themselves.
FOOTNOTES:
[376] Braun, 'Pringsheim Jahrbuch f. Wiss. Bot.,' 1858, 1, p. 307, tab. 22, 23.
[377] Henfrey, 'Jour. Linn. Soc. Bot.,' vol. iii, p. 159.
[378] On the subject of chorisis or dédoublement the reader may profitably consult Moquin-
Tandon, 'Ess. sur les Dédoublements,' and the same author in 'Ann. Sc. Nat.,' t. xxvii, p. 236.
and 'El. Ter. Veget.,' p. 337. Dunal, 'Consid. Org. Fleur.,' Montpell., 1829, p. 32, note 3. A. de
St. Hilaire in 'Ann. Sc. Nat.,' ser. 3, t. iii, p. 355, adnot. Lindley, 'Elements of Botany,' p. 76.
Asa Gray. 'Botanical Text Book.'
Page 440
CHAPTER I.
MULTIPLICATION OF AXILE ORGANS, INFLORESCENCE, ETC.
By Linné an undue number of branches was designated as "plica," from the
analogy with the disease of the hair known as plica polonica: "Plicata
dicitur planta, cum arbor vel ramus excrescit minimis intertextis ramulis,
tanquam plica polonica ex pilis, ceu instar nidi Picæ, quod vulgo a genio
ortum arbitratur; frequens apud nos in Betula, præsertim Norlandiæ, in
Carpino Scaniæ, nec infrequens in Pinu."[379]
By some of the older authors this condition was called polyclady. In some
cases, it would seem to be due to fungi as in the witches' brooms
(hexenbesen) of the German forests; in other instances, it is a result of
mutilation as after the operation of pollarding.
Moquin-Tandon[380] mentions a case in a grafted ash in the botanic garden
of Toulouse, where below the graft there was a large swelling, from which
proceeded more than a thousand densely-packed, interlacing branches.
This must have been similar to the condition so commonly met with in the
birch, and frequently in the hornbeam and the thorn, and which has
prompted so many a schoolboy to climb the tree in quest of the apparent
nest. It is probable that some of the large "gnaurs" or "burrs," met with in
elms, &c., also in certain varieties of apples, are clusters of adventitious
buds, some of which might, and sometimes do, lengthen out into branches.
An increased number of branches also necessarily arises when the flower-
buds are replaced by leaf-buds.
MULTIPLICATION OF AXILE ORGANS, INFLORESCENCE, ETC.
By Linné an undue number of branches was designated as "plica," from the
analogy with the disease of the hair known as plica polonica: "Plicata
dicitur planta, cum arbor vel ramus excrescit minimis intertextis ramulis,
tanquam plica polonica ex pilis, ceu instar nidi Picæ, quod vulgo a genio
ortum arbitratur; frequens apud nos in Betula, præsertim Norlandiæ, in
Carpino Scaniæ, nec infrequens in Pinu."[379]
By some of the older authors this condition was called polyclady. In some
cases, it would seem to be due to fungi as in the witches' brooms
(hexenbesen) of the German forests; in other instances, it is a result of
mutilation as after the operation of pollarding.
Moquin-Tandon[380] mentions a case in a grafted ash in the botanic garden
of Toulouse, where below the graft there was a large swelling, from which
proceeded more than a thousand densely-packed, interlacing branches.
This must have been similar to the condition so commonly met with in the
birch, and frequently in the hornbeam and the thorn, and which has
prompted so many a schoolboy to climb the tree in quest of the apparent
nest. It is probable that some of the large "gnaurs" or "burrs," met with in
elms, &c., also in certain varieties of apples, are clusters of adventitious
buds, some of which might, and sometimes do, lengthen out into branches.
An increased number of branches also necessarily arises when the flower-
buds are replaced by leaf-buds.
Page 441
Fig. 179.—Flower stalks of Bellevalia comosa, nat. size, after Morren.
Occasionally, a great increase in the number of pedicels, or flower-stalks,
may be met with in conjunction with a decreased number of flowers, as in
the wig-plant (Rhus Cotinus), or the feather-hyacinth (Bellevalia comosa).
In these cases the supernumerary pedicels are often brightly coloured. To
this condition Morren gave the name mischomany, from μἱσχος, a pedicel, a
term which has not generally been adopted.[381]
Occasionally, a great increase in the number of pedicels, or flower-stalks,
may be met with in conjunction with a decreased number of flowers, as in
the wig-plant (Rhus Cotinus), or the feather-hyacinth (Bellevalia comosa).
In these cases the supernumerary pedicels are often brightly coloured. To
this condition Morren gave the name mischomany, from μἱσχος, a pedicel, a
term which has not generally been adopted.[381]
Page 442
Fig. 180.—Tuft of branches at the end of the inflorescence of Bellevalia comosa, enlarged after
Morren.
M. Fournier[382] describes a case in the butcher's broom (Ruscus aculeatus),
wherein from the axil of the minute leaf subtending the flower a secondary
flattened branch proceeded.
Duchartre[383] cites the case of a hyacinth which, in addition to the usual
scape, had a second smaller one by its side terminated by a solitary flower;
indeed, such an occurrence is not uncommon.
Some tulips occasionally present three or four, or more, flowers on one
inflorescence, but whether from a branching of the primary scape, or from
the premature development of some of the axillary bulbils into flowering
stems which become adherent to the primary flower-stalk, cannot, in all
cases, be determined. Certainly, in some cases examined by me the latter
was the case.[384]
Morren.
M. Fournier[382] describes a case in the butcher's broom (Ruscus aculeatus),
wherein from the axil of the minute leaf subtending the flower a secondary
flattened branch proceeded.
Duchartre[383] cites the case of a hyacinth which, in addition to the usual
scape, had a second smaller one by its side terminated by a solitary flower;
indeed, such an occurrence is not uncommon.
Some tulips occasionally present three or four, or more, flowers on one
inflorescence, but whether from a branching of the primary scape, or from
the premature development of some of the axillary bulbils into flowering
stems which become adherent to the primary flower-stalk, cannot, in all
cases, be determined. Certainly, in some cases examined by me the latter
was the case.[384]
Page 443
Under this head, too, may be included those cases wherein an ordinarily
spicate inflorescence becomes paniculate owing to the branching of the axis
and the formation of an unwonted number of secondary buds. Instances of
this kind may be met with in willows, hazels, alders, and other amentaceous
plants. In the case of the hazel the unusual development of male catkins
sometimes coincides with an alteration in their position, instead of being
placed near the axil of a leaf; they become terminal. Jaeger figures and
describes a bunch of Pinus sylvestris bearing in one case seventy minute
cones, and in another fifty-nine. These cones preserved the same spiral
arrangement among themselves which is proper to the leaves. These latter,
indeed, replaced the strobili above.[385]
Fig. 181.—Increased number of male catkins in the hazel Corylus avellana.
M. Reichardt describes an analogous case in the same species, and
attributes the inordinate number of cones to a fungus (Peridermium pini). In
spicate inflorescence becomes paniculate owing to the branching of the axis
and the formation of an unwonted number of secondary buds. Instances of
this kind may be met with in willows, hazels, alders, and other amentaceous
plants. In the case of the hazel the unusual development of male catkins
sometimes coincides with an alteration in their position, instead of being
placed near the axil of a leaf; they become terminal. Jaeger figures and
describes a bunch of Pinus sylvestris bearing in one case seventy minute
cones, and in another fifty-nine. These cones preserved the same spiral
arrangement among themselves which is proper to the leaves. These latter,
indeed, replaced the strobili above.[385]
Fig. 181.—Increased number of male catkins in the hazel Corylus avellana.
M. Reichardt describes an analogous case in the same species, and
attributes the inordinate number of cones to a fungus (Peridermium pini). In
Page 444
this case there were no less than 227 cones, but each one half the size of the
ordinary cones.[386]
Of a similar character is the many-headed pineapple. Among grasses such a
branching of the inflorescence is exceedingly common,—which is the more
readily understood as the normal inflorescence is in so many cases
paniculate. Cultivators have, in some instances, availed themselves of this
peculiarity, as in the Egyptian wheat or corn of abundance (Triticum
compositum), certain varieties of Maize, etc. Similar exuberant growths
occur in Orchidaceæ, in Cyperaceæ, e.g. Carex, in Restiaceæ, and indeed
they may be found in any plant with a similar form of inflorescence. In all
these cases the branching begins at the lower part of the spike, and extends
from below upwards in an indefinite manner, even although the primary
inflorescence be definite.
Among the Equisetaceæ a similar plurality of spikes occurs often as a result
of mutilation.[387] The deviation in question might in some instances be
turned to good account, as in the Triticum before mentioned or as in the
broccoli shown at fig. 182, though it must be added that the apparent
advantages are often counterpoised by some undesirable qualities or by
some circumstance which prevents us availing ourselves of the new
condition.
Multiplication of Bulbs.—This occurrence has been briefly alluded to
previously (see p. 84). The most curious cases are those in which one bulb
is placed on the top of another as happened in some bulbs of Leucoium
æstivum described by M. Gay.[388] Irmisch described a similar phenomenon
in L. vernum; and Mr. Moggridge has communicated drawings of a similar
formation in the same species grown in the neighbourhood of Mentone.
ordinary cones.[386]
Of a similar character is the many-headed pineapple. Among grasses such a
branching of the inflorescence is exceedingly common,—which is the more
readily understood as the normal inflorescence is in so many cases
paniculate. Cultivators have, in some instances, availed themselves of this
peculiarity, as in the Egyptian wheat or corn of abundance (Triticum
compositum), certain varieties of Maize, etc. Similar exuberant growths
occur in Orchidaceæ, in Cyperaceæ, e.g. Carex, in Restiaceæ, and indeed
they may be found in any plant with a similar form of inflorescence. In all
these cases the branching begins at the lower part of the spike, and extends
from below upwards in an indefinite manner, even although the primary
inflorescence be definite.
Among the Equisetaceæ a similar plurality of spikes occurs often as a result
of mutilation.[387] The deviation in question might in some instances be
turned to good account, as in the Triticum before mentioned or as in the
broccoli shown at fig. 182, though it must be added that the apparent
advantages are often counterpoised by some undesirable qualities or by
some circumstance which prevents us availing ourselves of the new
condition.
Multiplication of Bulbs.—This occurrence has been briefly alluded to
previously (see p. 84). The most curious cases are those in which one bulb
is placed on the top of another as happened in some bulbs of Leucoium
æstivum described by M. Gay.[388] Irmisch described a similar phenomenon
in L. vernum; and Mr. Moggridge has communicated drawings of a similar
formation in the same species grown in the neighbourhood of Mentone.
Page 445
From the instances cited it is clear that branching of the inflorescence
occurs most frequently in those plants naturally characterised by a dense
compact mode of growth, whether that be definite or indefinite, as in spikes,
umbels, capitula, &c.; so that compound spikes, umbels, &c., are formed in
the place of simple ones (see also prolification of the inflorescence, p. 102).
Fig. 182.—Broccoli, with six perfect heads on one stalk ('Gard. Chron.,' 1856, Oct. 25).
Increased number of florets in the individual spikelets of grasses is also
met with under some circumstances. I have seen this in Hordeum and
Lolium, and an instance is figured in Avena by Dr. Wiegmann.[389] M.
Duval Jouve[390] records a similar occurrence in Catabrosa aquatica, the
spikelets of which contained from two to seven flowers.[391]
FOOTNOTES:
[379] 'Phil Bot.,' § 274.
occurs most frequently in those plants naturally characterised by a dense
compact mode of growth, whether that be definite or indefinite, as in spikes,
umbels, capitula, &c.; so that compound spikes, umbels, &c., are formed in
the place of simple ones (see also prolification of the inflorescence, p. 102).
Fig. 182.—Broccoli, with six perfect heads on one stalk ('Gard. Chron.,' 1856, Oct. 25).
Increased number of florets in the individual spikelets of grasses is also
met with under some circumstances. I have seen this in Hordeum and
Lolium, and an instance is figured in Avena by Dr. Wiegmann.[389] M.
Duval Jouve[390] records a similar occurrence in Catabrosa aquatica, the
spikelets of which contained from two to seven flowers.[391]
FOOTNOTES:
[379] 'Phil Bot.,' § 274.
Page 446
[380] 'El. Ter. Veget.,' p. 392.
[381] 'Bull. Acad. Belg.,' xvii, part ii, p. 38.
[382] 'Bull. Soc. Bot. Fr.,' vol. iv, 1857. p. 760.
[383] Ibid., vol. viii, 1861, p. 159.
[384] See 'Gard. Chron.,' July, 1866, p. 656, and Clusius, 'Plant. Rar.,' lib. 2, p. 143, Tulipa
serotina πολυκλαδης, minor, &c. Hort. Eysttett. Plant. Vern.,' fol. 12.
[385] 'Jaeger de Pini sylvestris monstrositate,' Stuttgardt, 1828.
[386] Cited in 'Bull. Soc. Bot. Fr.,' xiv, p. 265.
[387] Duval Jouve, 'Hist. Nat. Equiset. Fr.,' tab. 8, also Milde, 'Nov. Act. Acad. Nat. Cur.,' t.
xxvi, part 2. For branched inflorescence of orchids, see 'Reichenbach Proc. Lond. Bot.
Congress,' 1866, p. 121.
[388] 'Bull. Soc. Bot. Fr.,' vi, 266, vii, 457. Irmisch, 'Knollen und Zwiebelgew.,' tab. 7, figs.
10, 11.
[389] 'Flora,' 1831, p. 5, tab. i; see also Hanstein, 'Flora,' 1857, p. 513. Schlechtendal, 'Bot.
Zeit.,' xviii, p. 381.
[390] 'Bull. Soc. Bot. Fr.,' ix, p. 8.
[391] It will be seen, from what has been just said, that in some of the cases where the axile
organs, branches, &c., appear to be multiplied, the increased number is due to subdivision
rather than to renewed formation (see Fission). Of this last description is an instance which
came under the writer's notice after the section relating to that subject was in print, and which
may therefore here be alluded to. The instance is that of the subdivision of the leaf-like organs
of Sciadopitys verticillata. In one instance the pseudo leaf divided, and from the division
proceeded a little axis, bearing at its summit a verticil of pseudo leaves. This division and
formation of new axes and verticils affords ample confirmation of the opinion thrown out by
Professor Alexander Dickson, that the apparent leaves of this plant were really branches: see
'Revue Horticole,' 1867, and 'Report. Bot. Congress,' London, 1866, p. 124.
CHAPTER II.
MULTIPLICATION OF FOLIAR ORGANS.
[381] 'Bull. Acad. Belg.,' xvii, part ii, p. 38.
[382] 'Bull. Soc. Bot. Fr.,' vol. iv, 1857. p. 760.
[383] Ibid., vol. viii, 1861, p. 159.
[384] See 'Gard. Chron.,' July, 1866, p. 656, and Clusius, 'Plant. Rar.,' lib. 2, p. 143, Tulipa
serotina πολυκλαδης, minor, &c. Hort. Eysttett. Plant. Vern.,' fol. 12.
[385] 'Jaeger de Pini sylvestris monstrositate,' Stuttgardt, 1828.
[386] Cited in 'Bull. Soc. Bot. Fr.,' xiv, p. 265.
[387] Duval Jouve, 'Hist. Nat. Equiset. Fr.,' tab. 8, also Milde, 'Nov. Act. Acad. Nat. Cur.,' t.
xxvi, part 2. For branched inflorescence of orchids, see 'Reichenbach Proc. Lond. Bot.
Congress,' 1866, p. 121.
[388] 'Bull. Soc. Bot. Fr.,' vi, 266, vii, 457. Irmisch, 'Knollen und Zwiebelgew.,' tab. 7, figs.
10, 11.
[389] 'Flora,' 1831, p. 5, tab. i; see also Hanstein, 'Flora,' 1857, p. 513. Schlechtendal, 'Bot.
Zeit.,' xviii, p. 381.
[390] 'Bull. Soc. Bot. Fr.,' ix, p. 8.
[391] It will be seen, from what has been just said, that in some of the cases where the axile
organs, branches, &c., appear to be multiplied, the increased number is due to subdivision
rather than to renewed formation (see Fission). Of this last description is an instance which
came under the writer's notice after the section relating to that subject was in print, and which
may therefore here be alluded to. The instance is that of the subdivision of the leaf-like organs
of Sciadopitys verticillata. In one instance the pseudo leaf divided, and from the division
proceeded a little axis, bearing at its summit a verticil of pseudo leaves. This division and
formation of new axes and verticils affords ample confirmation of the opinion thrown out by
Professor Alexander Dickson, that the apparent leaves of this plant were really branches: see
'Revue Horticole,' 1867, and 'Report. Bot. Congress,' London, 1866, p. 124.
CHAPTER II.
MULTIPLICATION OF FOLIAR ORGANS.
Page 447
The cases referrible to this head may be ranged under two sections
according as the increase is due to plurality of ordinarily single organs, or to
an increase in the number of verticils or whorls.
When, in place of a single leaf organ two or more are really or in
appearance present the occurrence may be due to one of several causes;
among them may be mentioned an actual formation of parts in unwonted
number, hypertrophy or enation, chorisis or fission, disjunction, adhesion of
one leaf to another or to the stem, as in some of the leaves called
"geminate," wherein the two leaves, though apparently in juxtaposition, yet
originate from different parts of the stem, but by coalescence or lack of
separation produce the impression as if they sprang from the same node. In
the adult state it is not always possible to ascertain with certainty to which
of these causes the increase in the number of leaves is due, though a clue to
the real state of things may be gained from attention to the distribution of
the veins, to the arrangement or phyllotaxy of the leaves, the size and
position of the supernumerary organs, &c.
The term "phyllomania," as ordinarily used, is applied to an unwonted
development of leafy tissue, as in some begonias where the scales or
ramenta are replaced by small leaflets, or as in some cabbage leaves, from
the surface of which project, at right angles to the primary plane, other
secondary leafy plates; but these are, strictly speaking, cases of hypertrophy
(see Hypertrophy).
Those instances in which the actual number of leaves is increased, so that in
place of one there are more leaflets, may be included under the term
"pleiophylly," which may serve to designate both the appearance of two or
more leaves in the place usually occupied by a single one, and also those
normally compound leaves in which the number of leaflets is greater than
usual.
according as the increase is due to plurality of ordinarily single organs, or to
an increase in the number of verticils or whorls.
When, in place of a single leaf organ two or more are really or in
appearance present the occurrence may be due to one of several causes;
among them may be mentioned an actual formation of parts in unwonted
number, hypertrophy or enation, chorisis or fission, disjunction, adhesion of
one leaf to another or to the stem, as in some of the leaves called
"geminate," wherein the two leaves, though apparently in juxtaposition, yet
originate from different parts of the stem, but by coalescence or lack of
separation produce the impression as if they sprang from the same node. In
the adult state it is not always possible to ascertain with certainty to which
of these causes the increase in the number of leaves is due, though a clue to
the real state of things may be gained from attention to the distribution of
the veins, to the arrangement or phyllotaxy of the leaves, the size and
position of the supernumerary organs, &c.
The term "phyllomania," as ordinarily used, is applied to an unwonted
development of leafy tissue, as in some begonias where the scales or
ramenta are replaced by small leaflets, or as in some cabbage leaves, from
the surface of which project, at right angles to the primary plane, other
secondary leafy plates; but these are, strictly speaking, cases of hypertrophy
(see Hypertrophy).
Those instances in which the actual number of leaves is increased, so that in
place of one there are more leaflets, may be included under the term
"pleiophylly," which may serve to designate both the appearance of two or
more leaves in the place usually occupied by a single one, and also those
normally compound leaves in which the number of leaflets is greater than
usual.
Page 448
The increased number of leaves in a whorl may well be designated as
"polyphylly," using the word in the same sense as in ordinary descriptive
botany, while "pleiotaxy" may be applied to those cases in which the
number of whorls is increased.
Fig. 183.—Supernumerary leaflet, Ulmus campestris.
Pleiophylly.—As above stated, this term is proposed to designate those
cases in which there is an absolute increase in the number of leaves starting
from one particular point, as well as those in which the number of leaflets in
a compound leaf is preternaturally increased. The simplest cases are such as
are figured in the adjacent cuts, wherein, in place of a single leaf, two are
produced in the elm. In the one case the new leaflet springs from the apex
of the petiole and partially fills the space consequent on the obliquity of the
base of the leaf. In the other it would seem as if two distinct leaves emerged
from the stem in juxtaposition. This is probably due to a lateral chorisis or
subdivision of the primitive tubercle or growing point, followed by a like
"polyphylly," using the word in the same sense as in ordinary descriptive
botany, while "pleiotaxy" may be applied to those cases in which the
number of whorls is increased.
Fig. 183.—Supernumerary leaflet, Ulmus campestris.
Pleiophylly.—As above stated, this term is proposed to designate those
cases in which there is an absolute increase in the number of leaves starting
from one particular point, as well as those in which the number of leaflets in
a compound leaf is preternaturally increased. The simplest cases are such as
are figured in the adjacent cuts, wherein, in place of a single leaf, two are
produced in the elm. In the one case the new leaflet springs from the apex
of the petiole and partially fills the space consequent on the obliquity of the
base of the leaf. In the other it would seem as if two distinct leaves emerged
from the stem in juxtaposition. This is probably due to a lateral chorisis or
subdivision of the primitive tubercle or growing point, followed by a like
Page 449
subdivision of the vascular bundle supplying it. There are certain varieties
of elm that very generally present this anomaly on their rank, coarse,
growing shoots. In these cases the new growths have the same direction as
the primary one, but in other cases the supplementary production is exactly
reversed in direction. Thus, in the common hazel (Corylus) a second
smaller leaf proceeding from the end of the leaf-stalk at the base of the
primary one may frequently be seen. M. Germain de Saint Pierre records an
instance in a mulberry leaf, from the base of which proceeded a large leafy
expansion divided into two tubular, horn-like projections, and in the centre
a thread-like process representing the midrib and terminated by a small two-
lipped limb.[392] Dr. Ferdinand Müller speaks of a leaf of Pomaderris
elliptica as bearing a secondary leaf on its under surface.[393]
Fig. 184.—Supernumerary leaf, Ulmus montana.
of elm that very generally present this anomaly on their rank, coarse,
growing shoots. In these cases the new growths have the same direction as
the primary one, but in other cases the supplementary production is exactly
reversed in direction. Thus, in the common hazel (Corylus) a second
smaller leaf proceeding from the end of the leaf-stalk at the base of the
primary one may frequently be seen. M. Germain de Saint Pierre records an
instance in a mulberry leaf, from the base of which proceeded a large leafy
expansion divided into two tubular, horn-like projections, and in the centre
a thread-like process representing the midrib and terminated by a small two-
lipped limb.[392] Dr. Ferdinand Müller speaks of a leaf of Pomaderris
elliptica as bearing a secondary leaf on its under surface.[393]
Fig. 184.—Supernumerary leaf, Ulmus montana.
Page 450
Fig. 185.—Supernumerary leaf of hazel.
The leaves of Heterocentron macrodon have likewise been observed
occasionally to produce leaflets from their upper surface.
To this production of leaves from leaves the late Professor Morren applied
the term "autophyllogeny."[394] The Belgian botanist figures a small perfect
leaf springing from the nerves of the upper surface of the primary leaf in a
species of Miconia. As in the hazel, the direction of the adventitious leaf is
inversely that of the primary one, the upper surface of the supernumerary
leaflet being turned towards the corresponding surface of the normal leaf. A
similar occurrence took place in Gesnera zebrina, but the new growth in
this case sprang from the lower face of the leaf. Morren explains the
appearances in question by supposing that the supplementary leaf is one of
a pair belonging to a bud borne on a slender stalk. This stalk and one of the
bud-leaves are supposed to be inseparably united with the primary leaf. But
there is no reason at all for supposing the existence of adhesion in these
cases; no trace of any such union is to be seen. A much more natural
explanation is that, from some cause or another, development at the apex of
the petiole or on the surface of the nerves, instead of taking place in one
The leaves of Heterocentron macrodon have likewise been observed
occasionally to produce leaflets from their upper surface.
To this production of leaves from leaves the late Professor Morren applied
the term "autophyllogeny."[394] The Belgian botanist figures a small perfect
leaf springing from the nerves of the upper surface of the primary leaf in a
species of Miconia. As in the hazel, the direction of the adventitious leaf is
inversely that of the primary one, the upper surface of the supernumerary
leaflet being turned towards the corresponding surface of the normal leaf. A
similar occurrence took place in Gesnera zebrina, but the new growth in
this case sprang from the lower face of the leaf. Morren explains the
appearances in question by supposing that the supplementary leaf is one of
a pair belonging to a bud borne on a slender stalk. This stalk and one of the
bud-leaves are supposed to be inseparably united with the primary leaf. But
there is no reason at all for supposing the existence of adhesion in these
cases; no trace of any such union is to be seen. A much more natural
explanation is that, from some cause or another, development at the apex of
the petiole or on the surface of the nerves, instead of taking place in one
Page 451
plane only, as usual, takes place in more than one, thus showing the close
relationship, if not the intrinsic identity, between the leaf-stalk and its
continuation, the midrib, with the branch and its subdivisions. The form of
the leaf-stalk and the arrangement of the vascular bundles in a circle in the
case of the hazel, before alluded to, bear out this notion. Such cases are
significant in reference to the notion propounded by M. Casimir de
Candolle, that the leaf is the equivalent of a branch in which the upper
portion of the vascular circle is abortive.[395]
Compound leaves, as has been stated, occasionally produce an extra
number of leaflets; one of the most familiar illustrations of this is in the
case of the four-leaved shamrock (Trifolium repens), which was gathered at
night-time during the full moon by sorceresses, who mixed it with vervain
and other ingredients, while young girls in search of a token of perfect
happiness made quest of the plant by day. Linné, who in this matter, at any
rate, had less than his usual feeling for romance, says of the four-leaved
trefoil that it differs no more from the ordinary trefoil than a man with six
fingers differs from one provided with the ordinary number. It should be
stated that five and six adventitious leaflets are found almost as frequently
as four.
Walpers describes a case where the leaf of T. repens bore seven leaflets.
Schlechtendal alludes to a similar increase in number in Cytisus Laburnum,
and many other instances might be cited.
For figures or descriptions of four-leaved shamrocks the reader is referred to Lobel, 'Stirp.
Advers.,' Nov., p. 382. Tabernæmontanus 'Krauterbuch,' S. 222. Schlechtendal, 'Bot. Zeit.,'
ix, p. 583, xiv, p. 71. Maugin, 'Bull. Soc Bot. Fr.,' 1866, t. xiii, p. 279. See also Cramer,
'Bildungsabweich,' p. 92. Walpers, 'Linnæa,' 1840, p. 362 (7-leaved). Schlechtendal, 'Bot.
Zeit.,' 1844, p. 457, Cytisus. Wigand, 'Flora,' 1856, p. 706.
Frondiferous leaves have much the appearance of branches provided with
leaves, and they may be compared with those instances in which an
relationship, if not the intrinsic identity, between the leaf-stalk and its
continuation, the midrib, with the branch and its subdivisions. The form of
the leaf-stalk and the arrangement of the vascular bundles in a circle in the
case of the hazel, before alluded to, bear out this notion. Such cases are
significant in reference to the notion propounded by M. Casimir de
Candolle, that the leaf is the equivalent of a branch in which the upper
portion of the vascular circle is abortive.[395]
Compound leaves, as has been stated, occasionally produce an extra
number of leaflets; one of the most familiar illustrations of this is in the
case of the four-leaved shamrock (Trifolium repens), which was gathered at
night-time during the full moon by sorceresses, who mixed it with vervain
and other ingredients, while young girls in search of a token of perfect
happiness made quest of the plant by day. Linné, who in this matter, at any
rate, had less than his usual feeling for romance, says of the four-leaved
trefoil that it differs no more from the ordinary trefoil than a man with six
fingers differs from one provided with the ordinary number. It should be
stated that five and six adventitious leaflets are found almost as frequently
as four.
Walpers describes a case where the leaf of T. repens bore seven leaflets.
Schlechtendal alludes to a similar increase in number in Cytisus Laburnum,
and many other instances might be cited.
For figures or descriptions of four-leaved shamrocks the reader is referred to Lobel, 'Stirp.
Advers.,' Nov., p. 382. Tabernæmontanus 'Krauterbuch,' S. 222. Schlechtendal, 'Bot. Zeit.,'
ix, p. 583, xiv, p. 71. Maugin, 'Bull. Soc Bot. Fr.,' 1866, t. xiii, p. 279. See also Cramer,
'Bildungsabweich,' p. 92. Walpers, 'Linnæa,' 1840, p. 362 (7-leaved). Schlechtendal, 'Bot.
Zeit.,' 1844, p. 457, Cytisus. Wigand, 'Flora,' 1856, p. 706.
Frondiferous leaves have much the appearance of branches provided with
leaves, and they may be compared with those instances in which an
Page 452
adventitious bud is placed on the surface or edges of the leaves, as in
Gesnera, Cardamine, &c. In truth, the two conditions merge one into the
other, as in some begonias, where the ramenta often become leaf-like and
bear small bulbils in the axil.
When frondiferous leaves die the appendages die also, but when a true bud
has been formed on a leaf it does not of necessity die with the leaf that
bears it, but separates from it and continues to grow independently.
Increased number of stipules, spathes, &c.—Seringe relates the
occasional presence of two or three additional stipules upon the leaf-stalks
of Salix fragilis, and even makes a variety (Salix pendula, var.
multistipulata).
An increase in the number of the spathes has been often noticed in
Arads[396]. Prof. Alex. Braun has studied this subject in some detail[397]. In
Calla palustris the shoot which continues the growth of the plant proceeds
from the axil of the last leaf but one; the very last leaf producing no bud,
but if accidentally a shoot is developed in this latter situation it produces
flowers at once. No leaves are formed, but, on the contrary, two or three
spathes surround the spadix, so that the presence of an increased number of
spathes in this plant is associated with the development of a side shoot from
the axil of the last leaf, the situation whence, under natural circumstances,
no shoot at all issues. The supernumerary spathes are not always on the
same level, but may be separated by a considerable interval. They vary very
much in size, and sometimes assume the form and appearance of leaves.
Similar anomalies occur in other Arads as Arum maculatum, Richardia
æthiopica, and Anthurium Scherzerianum, frequently combined with a leaf-
like appearance of the spathes and sometimes with a subdivision of the
spadix into two or three branches.
Gesnera, Cardamine, &c. In truth, the two conditions merge one into the
other, as in some begonias, where the ramenta often become leaf-like and
bear small bulbils in the axil.
When frondiferous leaves die the appendages die also, but when a true bud
has been formed on a leaf it does not of necessity die with the leaf that
bears it, but separates from it and continues to grow independently.
Increased number of stipules, spathes, &c.—Seringe relates the
occasional presence of two or three additional stipules upon the leaf-stalks
of Salix fragilis, and even makes a variety (Salix pendula, var.
multistipulata).
An increase in the number of the spathes has been often noticed in
Arads[396]. Prof. Alex. Braun has studied this subject in some detail[397]. In
Calla palustris the shoot which continues the growth of the plant proceeds
from the axil of the last leaf but one; the very last leaf producing no bud,
but if accidentally a shoot is developed in this latter situation it produces
flowers at once. No leaves are formed, but, on the contrary, two or three
spathes surround the spadix, so that the presence of an increased number of
spathes in this plant is associated with the development of a side shoot from
the axil of the last leaf, the situation whence, under natural circumstances,
no shoot at all issues. The supernumerary spathes are not always on the
same level, but may be separated by a considerable interval. They vary very
much in size, and sometimes assume the form and appearance of leaves.
Similar anomalies occur in other Arads as Arum maculatum, Richardia
æthiopica, and Anthurium Scherzerianum, frequently combined with a leaf-
like appearance of the spathes and sometimes with a subdivision of the
spadix into two or three branches.
Page 453
Engelmann relates the occurrence of an increased number of glumes in
Bromus velutinus associated with suppression of the flowers.
Polyphylly.—As previously explained, this term is here applied to those
cases in which the members of any particular whorl are increased in
number, the whorls themselves not necessarily being augmented.
The simplest cases of this kind are those in which we meet with an unusual
number of leaves in a whorl.
Increased number of leaves in a whorl.—This may arise from actual
multiplication, or from lateral chorisis, or fission. The true nature of the
case may usually be ascertained by an examination of the distribution of the
veins of the leaves, or of the fibrous cords of the stem, by the relative
position of the supernumerary organs, &c.
Among plants with normally opposite leaves the following occasionally
produce them in whorls of three:—Lonicera brachypoda, L. Xylosteum,
Weigela rosea, Cornus mas, Vinca minor, &c.
Paris quadrifolia may frequently be met with five leaves in its whorl, or
even six.[398]
Increased number of bracts.—This is not of infrequent occurrence; one of
the most curious instances is that recorded by Mr. Edwards[399] in
Cerastium glomeratum, where, in place of the usual pair of bracts at the
base of the head of flowers, there was a whorl of six or eight, forming an
involucre. The flowers in this case were apetalous and imperfect.
Polyphylly of the calyx.—This may occur without any other perceptible
change, while at other times the number of the other parts of the flower is
proportionately increased. In a flower of a plum six sepals in place of five
Bromus velutinus associated with suppression of the flowers.
Polyphylly.—As previously explained, this term is here applied to those
cases in which the members of any particular whorl are increased in
number, the whorls themselves not necessarily being augmented.
The simplest cases of this kind are those in which we meet with an unusual
number of leaves in a whorl.
Increased number of leaves in a whorl.—This may arise from actual
multiplication, or from lateral chorisis, or fission. The true nature of the
case may usually be ascertained by an examination of the distribution of the
veins of the leaves, or of the fibrous cords of the stem, by the relative
position of the supernumerary organs, &c.
Among plants with normally opposite leaves the following occasionally
produce them in whorls of three:—Lonicera brachypoda, L. Xylosteum,
Weigela rosea, Cornus mas, Vinca minor, &c.
Paris quadrifolia may frequently be met with five leaves in its whorl, or
even six.[398]
Increased number of bracts.—This is not of infrequent occurrence; one of
the most curious instances is that recorded by Mr. Edwards[399] in
Cerastium glomeratum, where, in place of the usual pair of bracts at the
base of the head of flowers, there was a whorl of six or eight, forming an
involucre. The flowers in this case were apetalous and imperfect.
Polyphylly of the calyx.—This may occur without any other perceptible
change, while at other times the number of the other parts of the flower is
proportionately increased. In a flower of a plum six sepals in place of five
Page 454
sometimes exist; a precisely similar occurrence in the flowers of the elder
(Sambucus), the Fuchsia, and of Œnanthe crocata, may occasionally be
met with. In the latter case, indeed, there are sometimes as many as ten
segments to the calyx, and this without the other parts of the flower being
correspondingly augmented. Among monocotyledons a similar increase is
not uncommon, as in Tulipa, Allium, Iris, Narcissus, &c.
In some plants there seems to exist normally much variation in the number
of parts; thus in some species of Lacistema in adjacent flowers the calyx
may be found with four, five, or six segments.
Most of these cases of polyphylly affecting the calyx may be explained by
lateral chorisis or fission.
Polyphylly of the corolla.—This may happen in connection with similar
alterations in the calyx and stamens, or sometimes as an isolated
occurrence. In the latter case it may be due to lateral chorisis, to
substitution, or to the development of organs usually suppressed; thus, when
in aconites we meet with four or five horn-like nectaries (petals) instead of
two only, as usual, the supernumerary ones are accounted for by the
inordinate development of parts which ordinarily are in an abortive or
rudimentary state only. This is borne out by what happens in Balsamineæ.
In the common garden balsam the fifth petal is occasionally present, while
in Hydrocera triflora this petal is always present.
In a flower of a Cyclamen recently examined there were ten petals in one
series, the additional five being evidently due to the subdivision of the five
primary ones; the natural circular plan of the flower was here replaced by
an elliptical one. A similar occurrence takes place in the flowers of maples
(Acer), which sometimes show an increased number of parts in their floral
whorls and an elliptical outline. Whether the additional organs in this last
(Sambucus), the Fuchsia, and of Œnanthe crocata, may occasionally be
met with. In the latter case, indeed, there are sometimes as many as ten
segments to the calyx, and this without the other parts of the flower being
correspondingly augmented. Among monocotyledons a similar increase is
not uncommon, as in Tulipa, Allium, Iris, Narcissus, &c.
In some plants there seems to exist normally much variation in the number
of parts; thus in some species of Lacistema in adjacent flowers the calyx
may be found with four, five, or six segments.
Most of these cases of polyphylly affecting the calyx may be explained by
lateral chorisis or fission.
Polyphylly of the corolla.—This may happen in connection with similar
alterations in the calyx and stamens, or sometimes as an isolated
occurrence. In the latter case it may be due to lateral chorisis, to
substitution, or to the development of organs usually suppressed; thus, when
in aconites we meet with four or five horn-like nectaries (petals) instead of
two only, as usual, the supernumerary ones are accounted for by the
inordinate development of parts which ordinarily are in an abortive or
rudimentary state only. This is borne out by what happens in Balsamineæ.
In the common garden balsam the fifth petal is occasionally present, while
in Hydrocera triflora this petal is always present.
In a flower of a Cyclamen recently examined there were ten petals in one
series, the additional five being evidently due to the subdivision of the five
primary ones; the natural circular plan of the flower was here replaced by
an elliptical one. A similar occurrence takes place in the flowers of maples
(Acer), which sometimes show an increased number of parts in their floral
whorls and an elliptical outline. Whether the additional organs in this last
Page 455
case are the result of complete lateral chorisis or of multiplication proper I
do not know.
Orchids are very subject to an increase in the number of their labella. As
illustrations may be cited an instance recorded by Mr. J. T. Moggridge in a
flower of Ophrys insectifera, and in which there were two labella without
any other visible deviation from the ordinary conformation.[400]
I am indebted to Mr. Hemsley for the communication of a similar specimen
in O. apifera, in which there were two divergent lips, each with the same
peculiar markings. One of the sepals in this flower was adherent to one of
the lateral petals. This augmentation of the labella depends sometimes on
the separation, one from the other, of the elements of which the lip is
composed, at other times on the development, in the guise of lips, of
stamens which are usually suppressed (see p. 380).
The following enumeration will suffice to show the genera in which an
increased number of petals or perianth-segments in any given whorl most
frequently occurs.
Anemone!
Ranunculus!
Aconitum!
Raphanus.
Bunias.
Saponaria.
Dianthus!
Pelargonium!
Hibiscus.
Fuchsia.
Sarothamnus!
Lotus!
Ulex!
Prunus!
Trifolium.
do not know.
Orchids are very subject to an increase in the number of their labella. As
illustrations may be cited an instance recorded by Mr. J. T. Moggridge in a
flower of Ophrys insectifera, and in which there were two labella without
any other visible deviation from the ordinary conformation.[400]
I am indebted to Mr. Hemsley for the communication of a similar specimen
in O. apifera, in which there were two divergent lips, each with the same
peculiar markings. One of the sepals in this flower was adherent to one of
the lateral petals. This augmentation of the labella depends sometimes on
the separation, one from the other, of the elements of which the lip is
composed, at other times on the development, in the guise of lips, of
stamens which are usually suppressed (see p. 380).
The following enumeration will suffice to show the genera in which an
increased number of petals or perianth-segments in any given whorl most
frequently occurs.
Anemone!
Ranunculus!
Aconitum!
Raphanus.
Bunias.
Saponaria.
Dianthus!
Pelargonium!
Hibiscus.
Fuchsia.
Sarothamnus!
Lotus!
Ulex!
Prunus!
Trifolium.
Page 456
Œnanthe and Umbellif. pl.!
Sambucus!
Bryonia.
Campanula.
Solanum.
Veronica.
Cyclamen!
Primula!
Anagallis!
Plumbago.
Jasminum.
Syringa!
Tradescantia.
Iris.
Tigridia.
Narcissus.
Tulipa.
Convallaria!
Paris!
Hyacinthus!
Allium!
Ornithogalum.
Orchideæ, sp. pl.!
For other illustrations see multiplication of whorls, petalody; see also Moquin, loc. cit., p.
350. Engelmann, loc. cit., p. 20, § 18. Cramer, loc. cit., p. 25.
Polyphylly of the andrœcium.—An increased number of stamens
frequently accompanies the corresponding alterations in other whorls, and
seems, if anything, to be more frequent among monocotyledonous plants
than among dicotyledonous ones; thus, we occasionally find tetramerous
flowers in Crocus, Hyacinthus, Tulipa, Iris, Tigridia, &c., and more rarely
in Yucca (Y. flexilis[401]).
The increased number of stamens in a single whorl may result from a
development of organs usually suppressed, and constitute a form of regular
peloria as in Linaria, wherein a fifth stamen is occasionally met with.
Sambucus!
Bryonia.
Campanula.
Solanum.
Veronica.
Cyclamen!
Primula!
Anagallis!
Plumbago.
Jasminum.
Syringa!
Tradescantia.
Iris.
Tigridia.
Narcissus.
Tulipa.
Convallaria!
Paris!
Hyacinthus!
Allium!
Ornithogalum.
Orchideæ, sp. pl.!
For other illustrations see multiplication of whorls, petalody; see also Moquin, loc. cit., p.
350. Engelmann, loc. cit., p. 20, § 18. Cramer, loc. cit., p. 25.
Polyphylly of the andrœcium.—An increased number of stamens
frequently accompanies the corresponding alterations in other whorls, and
seems, if anything, to be more frequent among monocotyledonous plants
than among dicotyledonous ones; thus, we occasionally find tetramerous
flowers in Crocus, Hyacinthus, Tulipa, Iris, Tigridia, &c., and more rarely
in Yucca (Y. flexilis[401]).
The increased number of stamens in a single whorl may result from a
development of organs usually suppressed, and constitute a form of regular
peloria as in Linaria, wherein a fifth stamen is occasionally met with.
Page 457
Among normally didynamous plants such numerical restitution, so to speak,
is not unusual; thus, in Veronica four and five stamens occur. Fresenius has
seen five stamens in Lamium, Mentha, Chelone;[402] Bentham in Melittis,
and other instances are cited under the head of peloria. Chorisis may also
serve to account for some of these cases; thus, Eichler[403] figures a flower
of Matthiola annua with five long stamens instead of four; one of the long
pairs of stamens has here undergone a greater degree of repetition than
usual. De Candolle[404] cites and figures a curious form of Capsella Bursa-
pastoris sent him by Jacquin, and which was to some extent reproduced by
seed. In the flowers of this variety there were no petals, but ten stamens;
hence De Candolle inferred that the petals were here replaced by stamens,
but Moquin[405] objects, and with justice, to this view, as the ten stamens
are all on the same line; he considers the additional stamens to be the result
of chorisis. Buchenau[406] mentions the presence of seven stamens in
another Crucifer, Ionopsidium acaule. Here the supernumerary organ was
placed between two of the long stamens. The effect of chorisis in producing
an augmentation of parts is well seen in some plants that have some of their
flowers provided with staminodes or abortive stamens, and others with
clusters or phalanges of perfect stamens. Thus, in the female flowers of
Liquidambar there are five small staminodes without anthers, whereas in
the male flower the stamens are numerous and grouped together in
phalanges, so that the relation of simple to compound stamens is in this case
readily seen, as also in many Malvaceæ, Sterculiaceæ, Byttneriaceæ,
Tiliaceæ, and Myrtaceæ. It is probably the idea of splitting or dilamination
involved in the word chorisis that has led many English botanists to hesitate
about accepting the notion. Had they looked upon the process as identical
with that by which a branched inflorescence replaces an unbranched one, or
a compound leaf takes the place of a simple one, the objections would not
have been raised with such force. The process consists, in most cases, not so
is not unusual; thus, in Veronica four and five stamens occur. Fresenius has
seen five stamens in Lamium, Mentha, Chelone;[402] Bentham in Melittis,
and other instances are cited under the head of peloria. Chorisis may also
serve to account for some of these cases; thus, Eichler[403] figures a flower
of Matthiola annua with five long stamens instead of four; one of the long
pairs of stamens has here undergone a greater degree of repetition than
usual. De Candolle[404] cites and figures a curious form of Capsella Bursa-
pastoris sent him by Jacquin, and which was to some extent reproduced by
seed. In the flowers of this variety there were no petals, but ten stamens;
hence De Candolle inferred that the petals were here replaced by stamens,
but Moquin[405] objects, and with justice, to this view, as the ten stamens
are all on the same line; he considers the additional stamens to be the result
of chorisis. Buchenau[406] mentions the presence of seven stamens in
another Crucifer, Ionopsidium acaule. Here the supernumerary organ was
placed between two of the long stamens. The effect of chorisis in producing
an augmentation of parts is well seen in some plants that have some of their
flowers provided with staminodes or abortive stamens, and others with
clusters or phalanges of perfect stamens. Thus, in the female flowers of
Liquidambar there are five small staminodes without anthers, whereas in
the male flower the stamens are numerous and grouped together in
phalanges, so that the relation of simple to compound stamens is in this case
readily seen, as also in many Malvaceæ, Sterculiaceæ, Byttneriaceæ,
Tiliaceæ, and Myrtaceæ. It is probably the idea of splitting or dilamination
involved in the word chorisis that has led many English botanists to hesitate
about accepting the notion. Had they looked upon the process as identical
with that by which a branched inflorescence replaces an unbranched one, or
a compound leaf takes the place of a simple one, the objections would not
have been raised with such force. The process consists, in most cases, not so
Page 458
much in actual cleavage of a pre-existing organ as in the development of
new-growing points from the old ones.
An illustration given by Moquin from Dunal[407] goes far to support the
notion here adopted. The majority of the stamens of laurels (Laurus) have,
says M. Dunal, on each side of the base of their filaments a small glandular
bifid appendage; these excrescences are liable to be changed into small
stamens. The male flowers have a four-leaved calyx, and sometimes eight
stamens, each with two glands, four in one row, opposite to the sepals, four
in a second series alternating with the first. More generally two of the
stamens are destitute of glands, but have in their place a perfectly developed
stamen, so that in these latter flowers there are twelve stamens.
M. Clos[408] mentions a flower of rue (Ruta) wherein there were two
stamens joined together below and placed in front of a petal, as in
Peganum.
Buchenau[409] mentions a flower of Lotus uliginosus in which there were
eleven stamens, namely, two free and nine monadelphous; and Hildebrand
describes an analogous increase in a flower of Sarothamnus scoparius in
which, in conjunction with a seven-toothed calyx, there were two carinas
and fourteen stamens. It would seem probable in this case that there was a
coalescence of two flowers at an early date and consequent suppression of
some of the parts of the flower. Whether this was the case or not in this
particular illustration, it is nevertheless certain that many of the recorded
instances of increased number in the organs of a flower are really the results
of a fusion of two or more flowers, though frequently in the adult state but
few traces of the coalescence are to be seen.
Polyphylly of the gynœcium.—Moquin[410] remarks that, as the pistils are,
generally speaking, more or less subject to pressure, owing to their central
new-growing points from the old ones.
An illustration given by Moquin from Dunal[407] goes far to support the
notion here adopted. The majority of the stamens of laurels (Laurus) have,
says M. Dunal, on each side of the base of their filaments a small glandular
bifid appendage; these excrescences are liable to be changed into small
stamens. The male flowers have a four-leaved calyx, and sometimes eight
stamens, each with two glands, four in one row, opposite to the sepals, four
in a second series alternating with the first. More generally two of the
stamens are destitute of glands, but have in their place a perfectly developed
stamen, so that in these latter flowers there are twelve stamens.
M. Clos[408] mentions a flower of rue (Ruta) wherein there were two
stamens joined together below and placed in front of a petal, as in
Peganum.
Buchenau[409] mentions a flower of Lotus uliginosus in which there were
eleven stamens, namely, two free and nine monadelphous; and Hildebrand
describes an analogous increase in a flower of Sarothamnus scoparius in
which, in conjunction with a seven-toothed calyx, there were two carinas
and fourteen stamens. It would seem probable in this case that there was a
coalescence of two flowers at an early date and consequent suppression of
some of the parts of the flower. Whether this was the case or not in this
particular illustration, it is nevertheless certain that many of the recorded
instances of increased number in the organs of a flower are really the results
of a fusion of two or more flowers, though frequently in the adult state but
few traces of the coalescence are to be seen.
Polyphylly of the gynœcium.—Moquin[410] remarks that, as the pistils are,
generally speaking, more or less subject to pressure, owing to their central
Page 459
position, and it may be added owing to their later development, than the
other parts of the flower, they are more subject to suppression than to
multiplication; nevertheless, augmentation in the number of carpels does
occasionally take place, especially when the other parts of the flower are
also augmented in number. Sometimes this increase in the number of
carpels is due to pure multiplication, without any other change. At other
times the increase is due to a substitution of stamens or other organs for
carpels (see Substitutions). In other cases the augmentation seems to be due
to the development of parts usually suppressed; for instance, in
Antirrhinum, where there are usually only two carpels present, but where,
under peculiar circumstances, five may be found—thus rendering the
symmetry complete.[411] In Papilionaceæ, wherein usually only one carpel
is developed, we occasionally find two, or even more, as in Wistaria,
Gleditschia, Trifolium, &c. In Prunus and Amygdalus from two to five
carpels are occasionally to be found,[412] in Mimosa five, in Umbelliferæ
three to five; in some composites, e.g. Spilanthes, five carpels have also
been noticed; in Cruciferæ three and four, in grasses three.[413] The double
cocoa-nut affords an illustration of the development of two carpels out of
three, one only generally arriving at perfection. Triple nuts (Corylus) also
owe their peculiarity to the equal development of all three carpels which
exist in the original flower, but of which, under ordinary circumstances, two
become abortive. It is necessary, however, to distinguish these cases from
those in which two embryos are developed in one seed.
The following list may serve to show in what genera this change has been
most frequently noticed, and it may be said in general terms that Cruciferæ,
Umbelliferæ, and Liliaceæ, are the orders most frequently affected. Cases of
peloria are not included in the subjoined list.
Nigella.
Aquilegia.
Pæonia!
other parts of the flower, they are more subject to suppression than to
multiplication; nevertheless, augmentation in the number of carpels does
occasionally take place, especially when the other parts of the flower are
also augmented in number. Sometimes this increase in the number of
carpels is due to pure multiplication, without any other change. At other
times the increase is due to a substitution of stamens or other organs for
carpels (see Substitutions). In other cases the augmentation seems to be due
to the development of parts usually suppressed; for instance, in
Antirrhinum, where there are usually only two carpels present, but where,
under peculiar circumstances, five may be found—thus rendering the
symmetry complete.[411] In Papilionaceæ, wherein usually only one carpel
is developed, we occasionally find two, or even more, as in Wistaria,
Gleditschia, Trifolium, &c. In Prunus and Amygdalus from two to five
carpels are occasionally to be found,[412] in Mimosa five, in Umbelliferæ
three to five; in some composites, e.g. Spilanthes, five carpels have also
been noticed; in Cruciferæ three and four, in grasses three.[413] The double
cocoa-nut affords an illustration of the development of two carpels out of
three, one only generally arriving at perfection. Triple nuts (Corylus) also
owe their peculiarity to the equal development of all three carpels which
exist in the original flower, but of which, under ordinary circumstances, two
become abortive. It is necessary, however, to distinguish these cases from
those in which two embryos are developed in one seed.
The following list may serve to show in what genera this change has been
most frequently noticed, and it may be said in general terms that Cruciferæ,
Umbelliferæ, and Liliaceæ, are the orders most frequently affected. Cases of
peloria are not included in the subjoined list.
Nigella.
Aquilegia.
Pæonia!
Page 460
Delphinium!
Iberis.
Diplotaxis.
Lunaria.
Ricotiana.
Octadenia.
Draba!
Lepidium.
*Cheiranthus!
Dianthus.
Brassica!
Parnassia.
*Acer!
Ptelea.
Citrus!
Philadelphus.
Prunus!
Amygdalus!
Cratægus!
Fuchsia!
Trapa!
Cassia.
Cercis.
Medicago.
*Phaseolus!
Wistaria.
Gleditschia.
Affonsea.
Trifolium!
Archidendron.
Mimosa.
Robinia.
Diphaca.
Cœsalpinia.
Vicia.
Anthyllis.
Cucurbita.
Passiflora!
Sambucus!
Iberis.
Diplotaxis.
Lunaria.
Ricotiana.
Octadenia.
Draba!
Lepidium.
*Cheiranthus!
Dianthus.
Brassica!
Parnassia.
*Acer!
Ptelea.
Citrus!
Philadelphus.
Prunus!
Amygdalus!
Cratægus!
Fuchsia!
Trapa!
Cassia.
Cercis.
Medicago.
*Phaseolus!
Wistaria.
Gleditschia.
Affonsea.
Trifolium!
Archidendron.
Mimosa.
Robinia.
Diphaca.
Cœsalpinia.
Vicia.
Anthyllis.
Cucurbita.
Passiflora!
Sambucus!
Page 461
*Œnanthe!
Daucus!
Angelica!
Heracleum!
Silaus.
Carum.
Thysselinum.
Campanula!
Spilanthes.
Chrysanthemum.
Anagallis.
Primula!
Fraxinus!
Lycium.
Cobæa.
Datura!
Solanum!
Sesamum.
Sideritis.
Coleus.
Veronica!
*Digitalis!
Antirrhinum!
Linaria.
Gloxinia!
Symphytum.
Anchusa.
Polygonum.
Euphorbia.
Cneorum.
Mercurialis!
Chenopodium.
Suæda.
Beta.
Corylus!
Lambertia.
Cocos!
Tigridia.
Tulipa!
Daucus!
Angelica!
Heracleum!
Silaus.
Carum.
Thysselinum.
Campanula!
Spilanthes.
Chrysanthemum.
Anagallis.
Primula!
Fraxinus!
Lycium.
Cobæa.
Datura!
Solanum!
Sesamum.
Sideritis.
Coleus.
Veronica!
*Digitalis!
Antirrhinum!
Linaria.
Gloxinia!
Symphytum.
Anchusa.
Polygonum.
Euphorbia.
Cneorum.
Mercurialis!
Chenopodium.
Suæda.
Beta.
Corylus!
Lambertia.
Cocos!
Tigridia.
Tulipa!
Page 462
Iris!
Narcissus!
Allium!
Ornithogalum.
Gagea!
Tradescantia!
Schœnodon.
Bambuseæ.
A few additional references may here be given to papers where an increased number of
carpels is described:—Engelmann, 'De Antholys,' § 17, p. 19. Bernhardi, 'Flora,' 1838, p.
129. Schkuhr., 'Bot. Handb.,' t. 179. Godron, 'Ann. Sc. Nat.,' ser. 5, vol. ii, p. 280, tab. xviii,
pluricarpellary Crucifers. Weber, 'Verhandl. Nat. Hist. Vereins. Rhein. Pruss.,' &c., 1860,
Cerasus, &c., &c. Baillon, 'Adansonia,' iv, p. 71, Trifolium. Schlechtendal, 'Bot. Zeit.,' xv,
p. 67, Datura, three-celled fruit; 'Bot. Zeit.,' xiii, p. 823, Phaseolus, double pistil—a
common case. Cramer, 'Bildungsabweich,' p. 99, reference to several leguminous plants
with polycarpellary pistils. Munro, Gen., 'Linn. Trans.,' vol. xxvi, p. 26, Bambuseæ. Alph.
de Candolle, 'Neue Denkschrift,' Cheiranthus. Schimper, 'Flora,' 1829, ii, p. 433. Wigand,
'Bot. Untersuch.' Fleischer, 'Missbild. Cultur Pfl.' Cramer, 'Bildungsabweich,' p. 65,
Umbelliferæ.
Polyphylly of the flower in general.—Although, for the sake of
convenience, multiplication has here been treated of as it affects the
members of individual whorls of the flower, yet it must be remembered
that, in general, the augmentation is not confined to one whorl, but affects
several; thus, if the sepals are increased, the petals are likely to be so
likewise, and so forth. One of the most curious illustrations of this is that
recorded by Mr. Berkeley[414] in a plum, wherein there was an increased
number of sepals, a corresponding augmentation in the petals, while the
pistil was composed of two and sometimes three carpels distinct from the
calyx and from each other. In the flowers there did not appear to be any
definite relation in the position of the parts either with reference one to
another or to the axis.
Narcissus!
Allium!
Ornithogalum.
Gagea!
Tradescantia!
Schœnodon.
Bambuseæ.
A few additional references may here be given to papers where an increased number of
carpels is described:—Engelmann, 'De Antholys,' § 17, p. 19. Bernhardi, 'Flora,' 1838, p.
129. Schkuhr., 'Bot. Handb.,' t. 179. Godron, 'Ann. Sc. Nat.,' ser. 5, vol. ii, p. 280, tab. xviii,
pluricarpellary Crucifers. Weber, 'Verhandl. Nat. Hist. Vereins. Rhein. Pruss.,' &c., 1860,
Cerasus, &c., &c. Baillon, 'Adansonia,' iv, p. 71, Trifolium. Schlechtendal, 'Bot. Zeit.,' xv,
p. 67, Datura, three-celled fruit; 'Bot. Zeit.,' xiii, p. 823, Phaseolus, double pistil—a
common case. Cramer, 'Bildungsabweich,' p. 99, reference to several leguminous plants
with polycarpellary pistils. Munro, Gen., 'Linn. Trans.,' vol. xxvi, p. 26, Bambuseæ. Alph.
de Candolle, 'Neue Denkschrift,' Cheiranthus. Schimper, 'Flora,' 1829, ii, p. 433. Wigand,
'Bot. Untersuch.' Fleischer, 'Missbild. Cultur Pfl.' Cramer, 'Bildungsabweich,' p. 65,
Umbelliferæ.
Polyphylly of the flower in general.—Although, for the sake of
convenience, multiplication has here been treated of as it affects the
members of individual whorls of the flower, yet it must be remembered
that, in general, the augmentation is not confined to one whorl, but affects
several; thus, if the sepals are increased, the petals are likely to be so
likewise, and so forth. One of the most curious illustrations of this is that
recorded by Mr. Berkeley[414] in a plum, wherein there was an increased
number of sepals, a corresponding augmentation in the petals, while the
pistil was composed of two and sometimes three carpels distinct from the
calyx and from each other. In the flowers there did not appear to be any
definite relation in the position of the parts either with reference one to
another or to the axis.
Page 463
Fig. 186.—Plum. Increased number of parts in the calycine, corolline, and carpellary whorls
respectively.
In Primulaceæ this general augmentation has been frequently noticed.[415]
Among Orchideæ the instance related by Dr. Seubert is worth alluding to
here. This botanist observed and figured a flower of Orchis palustris with
tetramerous arrangement of parts, that is to say there were four outer
segments to the perianth, four petals, of which two were lip-like, four
stamens, three of which were rudimentary, and an ovary with four parietal
placentæ.[416]
The following list will serve to show in what plants this general
augmentation of parts has been observed most frequently:
Ranunculus.
Clematis!
Delphinium.
respectively.
In Primulaceæ this general augmentation has been frequently noticed.[415]
Among Orchideæ the instance related by Dr. Seubert is worth alluding to
here. This botanist observed and figured a flower of Orchis palustris with
tetramerous arrangement of parts, that is to say there were four outer
segments to the perianth, four petals, of which two were lip-like, four
stamens, three of which were rudimentary, and an ovary with four parietal
placentæ.[416]
The following list will serve to show in what plants this general
augmentation of parts has been observed most frequently:
Ranunculus.
Clematis!
Delphinium.
Page 464
Brassica!
Ruta.
Acer!
Prunus!
Rosa!
Rubus.
Philadelphus!
Chrysosplenium.
Umbelliferæ, sp. pl.!
*Fuchsia!
Œnothera.
Adoxa.
Bryonia.
Cucumis!
Campanula!
Sambucus!
*Primula!
Anagallis!
Lycium.
Solanum.
Symphytum.
Syringa!
Linaria.
Chenopodium.
*Paris!
Convallaria!
Allium.
*Lilium!
*Tulipa!
Ornithogalum.
*Gagea!
Tradescantia!
Orchideæ, sp. pl.!
Increased number of ovules or seeds.—This appears not to be of very
frequent occurrence, at least in those plants where the number of these
organs is normally small; where, as in Primula, the ovules and seeds are
produced in large quantities, it is not practicable to ascertain whether the
Ruta.
Acer!
Prunus!
Rosa!
Rubus.
Philadelphus!
Chrysosplenium.
Umbelliferæ, sp. pl.!
*Fuchsia!
Œnothera.
Adoxa.
Bryonia.
Cucumis!
Campanula!
Sambucus!
*Primula!
Anagallis!
Lycium.
Solanum.
Symphytum.
Syringa!
Linaria.
Chenopodium.
*Paris!
Convallaria!
Allium.
*Lilium!
*Tulipa!
Ornithogalum.
*Gagea!
Tradescantia!
Orchideæ, sp. pl.!
Increased number of ovules or seeds.—This appears not to be of very
frequent occurrence, at least in those plants where the number of these
organs is normally small; where, as in Primula, the ovules and seeds are
produced in large quantities, it is not practicable to ascertain whether the
Page 465
number be augmented or not in any particular case. Very probably, the
attachment or source of origin of the ovules determines, in some measure,
their number. Thus, in the case of marginal placentation the number must be
limited by the narrow space from which they proceed, whereas in parietal
and free central placentation the ovules are generally numerous. In the latter
case, however, it will be remembered that solitary ovules are not rare. An
increased number of ovules is generally remarked in conjunction with some
other change, such as a foliaceous condition of the carpel, in which the
margins are disunited. In such cases the ovules may occupy the margin or
may be placed a short distance within it, as in the case of some open carpels
of Ranunculus Ficaria,[417] and in which two ovules were borne in shallow
depressions on the upper or inner surface of the open carpel and supplied
with vascular cords from the central bundle or midrib. The outer coating of
the ovule here contained barred or spiral fusiform vessels derived from the
source just indicated.
In the very common cases where the pistil of Trifolium repens becomes
foliaceous (see Frondescence), the outer ovules are generally two or more
instead of being solitary. So, also, in the Rose with polliniferous ovules (see
p. 274). Among Umbelliferæ affected with frondescence of the pistil a
similar increase in the number of ovules takes place. It will be borne in
mind that in most, if not all, these cases the structure of the ovule is itself
imperfect.[418]
What are called in popular parlance double almonds or double nuts
(Corylus) are cases where two seeds are developed in place of one.
In the 'Revue Horticole,' 1867, p. 382, mention is made of a bush which
produces these double nuts each year—in fact, it never produces any single-
seeded fruit. The plant was a chance seedling, perhaps itself the offspring of
a double-seeded parent. It would be interesting to observe if the character
attachment or source of origin of the ovules determines, in some measure,
their number. Thus, in the case of marginal placentation the number must be
limited by the narrow space from which they proceed, whereas in parietal
and free central placentation the ovules are generally numerous. In the latter
case, however, it will be remembered that solitary ovules are not rare. An
increased number of ovules is generally remarked in conjunction with some
other change, such as a foliaceous condition of the carpel, in which the
margins are disunited. In such cases the ovules may occupy the margin or
may be placed a short distance within it, as in the case of some open carpels
of Ranunculus Ficaria,[417] and in which two ovules were borne in shallow
depressions on the upper or inner surface of the open carpel and supplied
with vascular cords from the central bundle or midrib. The outer coating of
the ovule here contained barred or spiral fusiform vessels derived from the
source just indicated.
In the very common cases where the pistil of Trifolium repens becomes
foliaceous (see Frondescence), the outer ovules are generally two or more
instead of being solitary. So, also, in the Rose with polliniferous ovules (see
p. 274). Among Umbelliferæ affected with frondescence of the pistil a
similar increase in the number of ovules takes place. It will be borne in
mind that in most, if not all, these cases the structure of the ovule is itself
imperfect.[418]
What are called in popular parlance double almonds or double nuts
(Corylus) are cases where two seeds are developed in place of one.
In the 'Revue Horticole,' 1867, p. 382, mention is made of a bush which
produces these double nuts each year—in fact, it never produces any single-
seeded fruit. The plant was a chance seedling, perhaps itself the offspring of
a double-seeded parent. It would be interesting to observe if the character
Page 466
be retained by the original plant, and whether it can be perpetuated by seed
or by grafting.
It is necessary to distinguish in the case of the nut between additional seeds
or ovules, as just described, and the double, triple, or fourfold nuts that are
occasionally met with, and which are the result either of actual
multiplication of the carpels or of the continued development of some of the
carpels which, under ordinary circumstances cease to grow (see ante, p.
364). In the case of a ripe nut with two seeds it might be impossible to tell
whether the adventitious seed were the product of multiplication, or
whether it belonged, in the first instance, to the same carpel as that
producing the fellow-seed, or to a different and now obliterated ovary. In all
probability, however, the second seed would be accounted for by the
development of two seeds in one carpellary cavity.
There is still another condition occasionally met with in the almond, and
which must be discriminated from the more common multiplication of the
seed, and which is the multiplication of the embryos within the seed, and
which furnishes the subject of the succeeding paragraph.
Increased number of embryos.—A ripe seed usually contains but a single
embryo, although in the ovular state preparation is commonly made for
more; and, indeed, in certain natural orders plurality of embryos in the same
seed does occur, as in Cycadeæ and Coniferæ. In the seeds of the orange
(Citrus), in those of some Euphorbiaceæ, &c., there are frequently two or
more additional embryos. A similar occurrence has been recorded in the
mango, for a specimen of which I am indebted to the Rev. Mr. Parish, of
Moulmein.[419]
Plurality of embryos has also been observed in—
Raphanus sativus.
or by grafting.
It is necessary to distinguish in the case of the nut between additional seeds
or ovules, as just described, and the double, triple, or fourfold nuts that are
occasionally met with, and which are the result either of actual
multiplication of the carpels or of the continued development of some of the
carpels which, under ordinary circumstances cease to grow (see ante, p.
364). In the case of a ripe nut with two seeds it might be impossible to tell
whether the adventitious seed were the product of multiplication, or
whether it belonged, in the first instance, to the same carpel as that
producing the fellow-seed, or to a different and now obliterated ovary. In all
probability, however, the second seed would be accounted for by the
development of two seeds in one carpellary cavity.
There is still another condition occasionally met with in the almond, and
which must be discriminated from the more common multiplication of the
seed, and which is the multiplication of the embryos within the seed, and
which furnishes the subject of the succeeding paragraph.
Increased number of embryos.—A ripe seed usually contains but a single
embryo, although in the ovular state preparation is commonly made for
more; and, indeed, in certain natural orders plurality of embryos in the same
seed does occur, as in Cycadeæ and Coniferæ. In the seeds of the orange
(Citrus), in those of some Euphorbiaceæ, &c., there are frequently two or
more additional embryos. A similar occurrence has been recorded in the
mango, for a specimen of which I am indebted to the Rev. Mr. Parish, of
Moulmein.[419]
Plurality of embryos has also been observed in—
Raphanus sativus.
Page 467
*Citrus Aurantium!
Diosma, sp.
Hypericum perforatum.
Triphasia aurantiaca.
*Æsculus Hippocastanum!
Euonymus latifolius.
*Mangifera indica!
Eugenia Jambos.
Amygdalus vulgaris!
Vicia, sp.
Cassia, sp.
*Viscum album!
Daucus Carota.
Ardisia serrulata!
Cynanchum nigrum.
fuscatum.
Euphorbia rosea.
Cœlebogyne ilicifolia.
Allium fragrans.
Funckia, sp.
Carex maritima.
Zea Mays.
See Schauer's translation of Moquin-Tandon, 'El. Terat. Veget.,' p. 245, adnot., and 'Al.
Braun Polyembryonie.'
Increased number of the cotyledons.—Although the presence of one or of
two cotyledons in the embryo is generally accepted as a valuable means of
separating flowering plants into two primary groups, yet, like all other
means of discrimination, it occasionally fails, and, indeed, almost always
requires to be taken in conjunction with some other character. There are
cases among flowering plants where the embryo is homogeneous in its
structure, there are others in which the number of the cotyledons is more
than two. Thus, in some seeds of Cola acuminata the cotyledons vary in
number from two to five. I have not been able to ascertain precisely
whether this multiplication of the cotyledons is characteristic of all the
Diosma, sp.
Hypericum perforatum.
Triphasia aurantiaca.
*Æsculus Hippocastanum!
Euonymus latifolius.
*Mangifera indica!
Eugenia Jambos.
Amygdalus vulgaris!
Vicia, sp.
Cassia, sp.
*Viscum album!
Daucus Carota.
Ardisia serrulata!
Cynanchum nigrum.
fuscatum.
Euphorbia rosea.
Cœlebogyne ilicifolia.
Allium fragrans.
Funckia, sp.
Carex maritima.
Zea Mays.
See Schauer's translation of Moquin-Tandon, 'El. Terat. Veget.,' p. 245, adnot., and 'Al.
Braun Polyembryonie.'
Increased number of the cotyledons.—Although the presence of one or of
two cotyledons in the embryo is generally accepted as a valuable means of
separating flowering plants into two primary groups, yet, like all other
means of discrimination, it occasionally fails, and, indeed, almost always
requires to be taken in conjunction with some other character. There are
cases among flowering plants where the embryo is homogeneous in its
structure, there are others in which the number of the cotyledons is more
than two. Thus, in some seeds of Cola acuminata the cotyledons vary in
number from two to five. I have not been able to ascertain precisely
whether this multiplication of the cotyledons is characteristic of all the
Page 468
seeds of particular trees, or whether some only are thus affected. Some
fruits that I examined bore out the latter view, as in the same pod were
seeds with two, three, and four cotyledons respectively.
I have also seen three cotyledons present in embryo-plants of Correa,
Cratægus Oxyacantha, Dianthus sinensis, Daucus Carota, Cerasus Lauro-
cerasus. De Candolle alludes to a case of the kind in the bean, and figures a
species of Solanum with three cotyledons.[420] Jaeger alludes to a similar
instance in Apium Petroselinum;[421] Ehrenberg to one in the marigold
(Calendula);[422] Reinsch to an analogous appearance in the beech (Fagus),
associated with a union of the margins of two out of the three cotyledons,
and of those of two out of the three leaves next adjacent.[423] This fusion
seems frequently to accompany increase in the number of cotyledons. It
was so in the Correa, and in the Cratægus previously mentioned. Some of
these cases may be accounted for by chorisis or by a cleavage of the
original cotyledons, as happens, according to Duchartre,[424] in some
Coniferæ, which he considers to be improperly termed polycotyledonous.
Whether this holds good in the Loranths, where (Nuytsia, Psittacanthus) an
appearance of polycotyledony exists, is not stated. In the case of the rue
(Ruta) figured by M. A. de Jussieu[425] this splitting of one cotyledon into
two is sufficiently evident, as is also the case in the sycamore (Acer pseudo-
platanus), seedlings of which may often be met with divided cotyledons.
In other instances a fusion of two embryo plants may give rise to a similar
appearance, as in the Euphorbia and Sinapis found by M. Alph. de Candolle
(see ante, p. 56).
Pleiotaxy or multiplication of whorls.—In the preceding section notice
has been taken of the increased number of parts in a single whorl, but an
augmentation of the number of distinct whorls is still more frequently met
with. Many of the so-called double flowers owe their peculiarity to this
fruits that I examined bore out the latter view, as in the same pod were
seeds with two, three, and four cotyledons respectively.
I have also seen three cotyledons present in embryo-plants of Correa,
Cratægus Oxyacantha, Dianthus sinensis, Daucus Carota, Cerasus Lauro-
cerasus. De Candolle alludes to a case of the kind in the bean, and figures a
species of Solanum with three cotyledons.[420] Jaeger alludes to a similar
instance in Apium Petroselinum;[421] Ehrenberg to one in the marigold
(Calendula);[422] Reinsch to an analogous appearance in the beech (Fagus),
associated with a union of the margins of two out of the three cotyledons,
and of those of two out of the three leaves next adjacent.[423] This fusion
seems frequently to accompany increase in the number of cotyledons. It
was so in the Correa, and in the Cratægus previously mentioned. Some of
these cases may be accounted for by chorisis or by a cleavage of the
original cotyledons, as happens, according to Duchartre,[424] in some
Coniferæ, which he considers to be improperly termed polycotyledonous.
Whether this holds good in the Loranths, where (Nuytsia, Psittacanthus) an
appearance of polycotyledony exists, is not stated. In the case of the rue
(Ruta) figured by M. A. de Jussieu[425] this splitting of one cotyledon into
two is sufficiently evident, as is also the case in the sycamore (Acer pseudo-
platanus), seedlings of which may often be met with divided cotyledons.
In other instances a fusion of two embryo plants may give rise to a similar
appearance, as in the Euphorbia and Sinapis found by M. Alph. de Candolle
(see ante, p. 56).
Pleiotaxy or multiplication of whorls.—In the preceding section notice
has been taken of the increased number of parts in a single whorl, but an
augmentation of the number of distinct whorls is still more frequently met
with. Many of the so-called double flowers owe their peculiarity to this
Page 469
condition. The distinction between the two modes in which the parts of the
flower are increased in number has been pointed out by Engelmann,
Moquin, and others, and the two seem to require distinctive epithets; hence
the application of the terms polyphylly and pleiotaxy, as here proposed.
Pleiotaxy in the bracts.—An increase in the number of bracts has been met
with very constantly in a species of Mæsa, and in a peculiar variety of
carnation, called the wheat-ear carnation.[426] In some of these cases the
increase in the number of bracts is attended by a corresponding suppression
in the other parts of the flower. Such a condition has been frequently met
with in Gentiana Amarella, where the bracts are increased in number,
coloured purple, and destitute of any true floral organs. A similar condition
exists in some varieties of Plantago major (var. paniculata), as has been
previously stated, p. 109.
Fig. 187.—Wheat-ear carnation. The appearance is due to the multiplication of the bracts and the
suppression of the other parts of the flower.
flower are increased in number has been pointed out by Engelmann,
Moquin, and others, and the two seem to require distinctive epithets; hence
the application of the terms polyphylly and pleiotaxy, as here proposed.
Pleiotaxy in the bracts.—An increase in the number of bracts has been met
with very constantly in a species of Mæsa, and in a peculiar variety of
carnation, called the wheat-ear carnation.[426] In some of these cases the
increase in the number of bracts is attended by a corresponding suppression
in the other parts of the flower. Such a condition has been frequently met
with in Gentiana Amarella, where the bracts are increased in number,
coloured purple, and destitute of any true floral organs. A similar condition
exists in some varieties of Plantago major (var. paniculata), as has been
previously stated, p. 109.
Fig. 187.—Wheat-ear carnation. The appearance is due to the multiplication of the bracts and the
suppression of the other parts of the flower.
Page 470
It has been noticed also in the common pea, Pisum sativum, and M.
Lortet[427] records a case of the kind in Erica multiflora, the flowers of
which, under ordinary circumstances, are arranged in clusters, but in this
case the pedicels were more closely crowded than usual, and were covered
for their whole length with small rose-coloured bracts arranged in irregular
whorls, the upper ones sometimes enclosing imperfect flowers. In the
'Gardeners' Chronicle,' 1865, p. 769, is figured a corresponding instance of
Delphinium Consolida, in which the bracts were greatly increased in
number, petaloid, and, at the same time, the central organs of the flower
were wholly wanting.
Fig. 188.—Delphinium Consolida. Multiplication of bracts at the expense of the other parts of the
flower.
Lortet[427] records a case of the kind in Erica multiflora, the flowers of
which, under ordinary circumstances, are arranged in clusters, but in this
case the pedicels were more closely crowded than usual, and were covered
for their whole length with small rose-coloured bracts arranged in irregular
whorls, the upper ones sometimes enclosing imperfect flowers. In the
'Gardeners' Chronicle,' 1865, p. 769, is figured a corresponding instance of
Delphinium Consolida, in which the bracts were greatly increased in
number, petaloid, and, at the same time, the central organs of the flower
were wholly wanting.
Fig. 188.—Delphinium Consolida. Multiplication of bracts at the expense of the other parts of the
flower.
Page 471
Fig. 189.—Multiplication of bracts, &c., Pelargonium.
In flowers of Pelargonium may occasionally be seen a repetition of the
whorls of bracts, in conjunction with suppression and diminished size of
some of the other portions of the flower (fig. 189).
The common foxglove (Digitalis purpurea) has likewise occasionally been
observed subject to a similar malformation.
Cornus mas and C. suecica sometimes show a triple involucre.[428]
Irmish[429] records an analogous case in Anemone Hepatica, wherein the
involucre was doubled. Similar augmentation occurs in cultivated
Anemone. In addition to the plants already mentioned, Engelmann[430]
mentions as having produced bracts in unwonted numbers, Lythrum
Salicaria, Plantago major, Veronica spicata, Echium vulgare, Melilotus
arvensis, and Rubus fruticosus.
In flowers of Pelargonium may occasionally be seen a repetition of the
whorls of bracts, in conjunction with suppression and diminished size of
some of the other portions of the flower (fig. 189).
The common foxglove (Digitalis purpurea) has likewise occasionally been
observed subject to a similar malformation.
Cornus mas and C. suecica sometimes show a triple involucre.[428]
Irmish[429] records an analogous case in Anemone Hepatica, wherein the
involucre was doubled. Similar augmentation occurs in cultivated
Anemone. In addition to the plants already mentioned, Engelmann[430]
mentions as having produced bracts in unwonted numbers, Lythrum
Salicaria, Plantago major, Veronica spicata, Echium vulgare, Melilotus
arvensis, and Rubus fruticosus.
Page 472
It must here be remarked that this great number of the bracts occurs
naturally in such plants as Godoya, in which the bracts, or, as some consider
them, the segments of the calyx, are very numerous, and arranged in several
overlapping segments.
In some of the cultivated double varieties of Nigella the finely divided
involucral bracts are repeated over and over again, but on a diminished
scale, to the exclusion of all the other parts of the flower.
Pleiotaxy or repetition of the calyx.—The true calyx is very seldom
affected in this manner, unless such organs as the epicalyx of mallows,
Potentilla, &c., be considered as really parts of the calyx.
In Linaria vulgaris Rœper observed a calyx consisting of a double series,
each of five sepals, in conjunction with other changes.[431] It is also
common in double columbines, delphiniums, nigellas, &c.
In the 'Revue Horticole,' 1867, p. 71, fig. 9, is described and figured by M.
B. Verlot a curious variety of vine grown for years in the Botanic Garden at
Grenoble, under the name of the double-flowered vine. The place of the
flower is occupied by a large number of successive whorls of sepals
disposed in regular order, and without any trace of the other portions of the
flower. It is, in fact, more like a leaf-bud than a flower. The outermost
whorls of this flower open at the time when the ordinary flowers of vines
do; the second series are gradually produced, and expand about the time
when the ovaries of the normal flowers begin to swell; a third series then
gradually forms, and so on, until frost puts a stop to the growth. This
malformation, it appears, is produced annually in certain varieties of vine,
and may be perpetuated by cuttings.
The flower of the St. Valèry apple, already alluded to under the head of
sepalody, might equally well be placed here. It is not very material whether
naturally in such plants as Godoya, in which the bracts, or, as some consider
them, the segments of the calyx, are very numerous, and arranged in several
overlapping segments.
In some of the cultivated double varieties of Nigella the finely divided
involucral bracts are repeated over and over again, but on a diminished
scale, to the exclusion of all the other parts of the flower.
Pleiotaxy or repetition of the calyx.—The true calyx is very seldom
affected in this manner, unless such organs as the epicalyx of mallows,
Potentilla, &c., be considered as really parts of the calyx.
In Linaria vulgaris Rœper observed a calyx consisting of a double series,
each of five sepals, in conjunction with other changes.[431] It is also
common in double columbines, delphiniums, nigellas, &c.
In the 'Revue Horticole,' 1867, p. 71, fig. 9, is described and figured by M.
B. Verlot a curious variety of vine grown for years in the Botanic Garden at
Grenoble, under the name of the double-flowered vine. The place of the
flower is occupied by a large number of successive whorls of sepals
disposed in regular order, and without any trace of the other portions of the
flower. It is, in fact, more like a leaf-bud than a flower. The outermost
whorls of this flower open at the time when the ordinary flowers of vines
do; the second series are gradually produced, and expand about the time
when the ovaries of the normal flowers begin to swell; a third series then
gradually forms, and so on, until frost puts a stop to the growth. This
malformation, it appears, is produced annually in certain varieties of vine,
and may be perpetuated by cuttings.
The flower of the St. Valèry apple, already alluded to under the head of
sepalody, might equally well be placed here. It is not very material whether
Page 473
the second whorl of organs be regarded as a repetition of the calyx or as a
row of petals in the guise of sepals.
Engelmann[432] cites the following plants as occasionally presenting a
repetition of the calyx, in most cases with a suppression of the other floral
whorls:—Stachys lanata, Myosotis palustris, Veronica media, Aquilegia
vulgaris, Nigella damascena, Campanula rapunculoides.
Pleiotaxy in the perianth.—Increase in the number of whorls in the
perianth is common in lilies, narcissus, hyacinths, &c. It may be also met
with occasionally among orchids. The lily of the valley (Convallaria
maialis) seems also to be particularly subject to an increase in the number
of parts of which its perianth consists, the augmentation being due partly to
repetition or pleiotaxy, partly to the substitution of petaloid segments for
stamens and pistils.[433]
In this place may also be mentioned the curious deviation from the ordinary
structure occasionally met with in Lilium candidum, and known in English
gardens as the double white lily. In this case there are no true flowers, but a
large number of petal-like segments disposed in an irregular spiral manner
at the extremity of the stem, some of the uppermost being occasionally
verticillate.[434]
row of petals in the guise of sepals.
Engelmann[432] cites the following plants as occasionally presenting a
repetition of the calyx, in most cases with a suppression of the other floral
whorls:—Stachys lanata, Myosotis palustris, Veronica media, Aquilegia
vulgaris, Nigella damascena, Campanula rapunculoides.
Pleiotaxy in the perianth.—Increase in the number of whorls in the
perianth is common in lilies, narcissus, hyacinths, &c. It may be also met
with occasionally among orchids. The lily of the valley (Convallaria
maialis) seems also to be particularly subject to an increase in the number
of parts of which its perianth consists, the augmentation being due partly to
repetition or pleiotaxy, partly to the substitution of petaloid segments for
stamens and pistils.[433]
In this place may also be mentioned the curious deviation from the ordinary
structure occasionally met with in Lilium candidum, and known in English
gardens as the double white lily. In this case there are no true flowers, but a
large number of petal-like segments disposed in an irregular spiral manner
at the extremity of the stem, some of the uppermost being occasionally
verticillate.[434]
Page 474
Fig. 190.—Double white lily. Multiplication of perianth-segments and other changes.
Pleiotaxy of the corolla.—With reference to double flowers, it was
remarked by Linné that polypetalous flowers were, as he said, multiplied,
while monopetalous flowers were duplicated, or triplicated, as the case may
be,[435] a statement that is true in the main, though it requires modification.
In the case of polypetalous, or rather dialypetalous flowers, the petals may
be very largely increased by multiplication, as in roses, anemones, pinks,
&c. In the last-named genus the number is often so much increased that the
calyx splits from the tension exercised on it by the increasing mass within.
This multiplication may happen without any metamorphy or substitution of
petals for stamens, though, in the majority of cases, it is associated with
such a change. It is curious to observe in some of these flowers that the total
number of parts is not greatly increased; thus, in some of the double-
flowered Leguminosæ, such as Ulex europæus and Lotus corniculatus, the
petals are repeated once or twice, the stamens are petalodic, but reduced in
Pleiotaxy of the corolla.—With reference to double flowers, it was
remarked by Linné that polypetalous flowers were, as he said, multiplied,
while monopetalous flowers were duplicated, or triplicated, as the case may
be,[435] a statement that is true in the main, though it requires modification.
In the case of polypetalous, or rather dialypetalous flowers, the petals may
be very largely increased by multiplication, as in roses, anemones, pinks,
&c. In the last-named genus the number is often so much increased that the
calyx splits from the tension exercised on it by the increasing mass within.
This multiplication may happen without any metamorphy or substitution of
petals for stamens, though, in the majority of cases, it is associated with
such a change. It is curious to observe in some of these flowers that the total
number of parts is not greatly increased; thus, in some of the double-
flowered Leguminosæ, such as Ulex europæus and Lotus corniculatus, the
petals are repeated once or twice, the stamens are petalodic, but reduced in
Page 475
number, while the carpels are usually entirely wanting. Thus, owing to the
diminished number of parts in the inner whorls of the flower, these very
double-looking blooms do not contain any greatly increased number of
parts.[436]
Flowers that, under ordinary circumstances, are gamopetalous, become, in
some instances, multiplied by the formation of additional segments, just as
in the case of polypetalous corollas; but in these cases the corollas become
polypetalous, their petals do not cohere one with another. Among double
flowers of this character may be mentioned Campanula rotundifolia,
Gardenia sp., Nerium Oleander, Serissa sp., Arbutus Unedo, &c. The
change is associated with petalody of the stamens and pistils.
A more frequent change among the monopetalous orders is the duplication
or triplication of the corolla, in consequence of which there appear to be a
series of corollas enclosed one within the other, the lobes of which
generally alternate with one another, but which sometimes are superposed.
This happens occasionally in the primrose (Primula acaulis), and
constitutes the variety called by the gardeners "hose in hose."
The same condition occurs frequently in some species of Datura and
Campanula.
diminished number of parts in the inner whorls of the flower, these very
double-looking blooms do not contain any greatly increased number of
parts.[436]
Flowers that, under ordinary circumstances, are gamopetalous, become, in
some instances, multiplied by the formation of additional segments, just as
in the case of polypetalous corollas; but in these cases the corollas become
polypetalous, their petals do not cohere one with another. Among double
flowers of this character may be mentioned Campanula rotundifolia,
Gardenia sp., Nerium Oleander, Serissa sp., Arbutus Unedo, &c. The
change is associated with petalody of the stamens and pistils.
A more frequent change among the monopetalous orders is the duplication
or triplication of the corolla, in consequence of which there appear to be a
series of corollas enclosed one within the other, the lobes of which
generally alternate with one another, but which sometimes are superposed.
This happens occasionally in the primrose (Primula acaulis), and
constitutes the variety called by the gardeners "hose in hose."
The same condition occurs frequently in some species of Datura and
Campanula.
Page 476
Fig. 191.—Campanula rotundifolia. "Double flowers" resulting from dialysis and multiplication of
the petals.
In Antirrhinum majus double flowers of this character sometimes occur; the
outermost corolla is normal, the succeeding ones usually have their petals
separate one from the other; the stamens are sometimes present, sometimes
absent, and at other times petalodic. Similar occurrences may be met with
in labiates and jasmines, and in Erica hyemalis.
Mr. W. B. Hemsley has kindly furnished me with flowers of a similar kind
occurring in wild specimens of Epacris impressa,[437] and there are
analogous phenomena in the common honeysuckle (Lonicera
Periclymenum), in which three corollas and no stamens often occur.
This duplication may either be accounted for on the theory of chorisis
above alluded to, or by supposing that the extra corolline whorl is due to a
series of confluent petalodic stamens; that the latter is the true explanation,
the petals.
In Antirrhinum majus double flowers of this character sometimes occur; the
outermost corolla is normal, the succeeding ones usually have their petals
separate one from the other; the stamens are sometimes present, sometimes
absent, and at other times petalodic. Similar occurrences may be met with
in labiates and jasmines, and in Erica hyemalis.
Mr. W. B. Hemsley has kindly furnished me with flowers of a similar kind
occurring in wild specimens of Epacris impressa,[437] and there are
analogous phenomena in the common honeysuckle (Lonicera
Periclymenum), in which three corollas and no stamens often occur.
This duplication may either be accounted for on the theory of chorisis
above alluded to, or by supposing that the extra corolline whorl is due to a
series of confluent petalodic stamens; that the latter is the true explanation,
Page 477
in certain cases at least, is shown by some flowers of Datura fastuosa, in
which the second corolla was partially staminal in its appearance, and bore
nearly perfect anthers, in addition to the five ordinary stamens, which were
unaltered either in form or position. Some partially virescent honeysuckle
flowers have a similar structure.
There are other cases of apparent multiplication or duplication, due,
probably, rather to the formation of outgrowths from the petals than to
actual augmentation of their number. These excrescences occur sometimes
on the inner surface of the petals, or of the corolla; at other times on the
outer surface, as in some gloxinias, &c. This matter will be more fully
treated of under the head of hypertrophy and enation.
Pleiotaxy of the andrœcium.—An increase in the number of whorls in the
stamens is very common, especially in cases where the number of circles of
stamens is naturally large. The augmentation of the number of stamens is
still more frequent where these organs are arranged, not in verticils, but in
one continuous spiral line.
In Cruciferæ there is always an indication of two whorls of stamens, and
this indication is rendered even more apparent in some varieties
accidentally met with. So in Saponaria, in Dianthus, and other
Caryophylleæ, three and four verticils of stamens have been met with. In
Lonicera Periclymenum a second whorl of stamens more or less petalodic
sometimes occurs.
Moquin mentions a variety of Rubus fruticosus in which nearly 900 petaloid
organs existed in the place of the twenty-five or thirty stamens natural to the
plant, the other organs of the flower being in their ordinary condition, with
the exception of the pistil, which did not attain its full size. Baillon records
the occasional existence of two rows of stamens in Ditaxis lancifolia.
which the second corolla was partially staminal in its appearance, and bore
nearly perfect anthers, in addition to the five ordinary stamens, which were
unaltered either in form or position. Some partially virescent honeysuckle
flowers have a similar structure.
There are other cases of apparent multiplication or duplication, due,
probably, rather to the formation of outgrowths from the petals than to
actual augmentation of their number. These excrescences occur sometimes
on the inner surface of the petals, or of the corolla; at other times on the
outer surface, as in some gloxinias, &c. This matter will be more fully
treated of under the head of hypertrophy and enation.
Pleiotaxy of the andrœcium.—An increase in the number of whorls in the
stamens is very common, especially in cases where the number of circles of
stamens is naturally large. The augmentation of the number of stamens is
still more frequent where these organs are arranged, not in verticils, but in
one continuous spiral line.
In Cruciferæ there is always an indication of two whorls of stamens, and
this indication is rendered even more apparent in some varieties
accidentally met with. So in Saponaria, in Dianthus, and other
Caryophylleæ, three and four verticils of stamens have been met with. In
Lonicera Periclymenum a second whorl of stamens more or less petalodic
sometimes occurs.
Moquin mentions a variety of Rubus fruticosus in which nearly 900 petaloid
organs existed in the place of the twenty-five or thirty stamens natural to the
plant, the other organs of the flower being in their ordinary condition, with
the exception of the pistil, which did not attain its full size. Baillon records
the occasional existence of two rows of stamens in Ditaxis lancifolia.
Page 478
Increased number of stamens in orchids, &c.—Various deviations from
the ordinary type of orchid structure have been already alluded to under the
head of displacement, fusion, peloria, substitution, &c., but the alterations
presented by the andrœcium in this family are so important in reference to
what is considered its natural conformation, that it seems desirable, in this
place, to enter upon the teratological appearances presented by the
andrœcium in this order, in somewhat greater detail than usual. The
ordinary structure of the flower with its three sepals, two petals, labellum,
column; and inferior ovary, is well known. Such a conformation would be
wholly anomalous and inexplicable were it not that the real number and
arrangement of parts have been revealed by various workers labouring to
the same end in different fields. Thus, Robert Brown, Link, Bauer, Darwin,
and others, paid special attention to the minute anatomy and mode of
distribution of the vessels; Irmisch, Crueger, Payer, and others, to the
evolution of the flower; Lindley, St. Hilaire, and Reichenbach, to the
comparison of the completed structures in the various genera and species;
while the teratological observers have been numerous, as will be seen from
the selected references cited at the end of this paragraph and in other places.
The result of this manifold study has been a pretty general agreement that
the structure of the order (omitting minor details) is as follows:—A six-
parted perianth in two rows, the outer three (sepals) generally regular and
equal in shape; of the inner three (petals or tepals) two are regular, and one,
the labellum very irregular, consisting not only of a petal, but of two
abortive stamens incorporated with it. The column is considered to be made
up of one perfect and three abortive stamens, in inseparable connection with
three styles. By some, however, it is supposed that all the stamens are
confluent with the column and none with the lip.
the ordinary type of orchid structure have been already alluded to under the
head of displacement, fusion, peloria, substitution, &c., but the alterations
presented by the andrœcium in this family are so important in reference to
what is considered its natural conformation, that it seems desirable, in this
place, to enter upon the teratological appearances presented by the
andrœcium in this order, in somewhat greater detail than usual. The
ordinary structure of the flower with its three sepals, two petals, labellum,
column; and inferior ovary, is well known. Such a conformation would be
wholly anomalous and inexplicable were it not that the real number and
arrangement of parts have been revealed by various workers labouring to
the same end in different fields. Thus, Robert Brown, Link, Bauer, Darwin,
and others, paid special attention to the minute anatomy and mode of
distribution of the vessels; Irmisch, Crueger, Payer, and others, to the
evolution of the flower; Lindley, St. Hilaire, and Reichenbach, to the
comparison of the completed structures in the various genera and species;
while the teratological observers have been numerous, as will be seen from
the selected references cited at the end of this paragraph and in other places.
The result of this manifold study has been a pretty general agreement that
the structure of the order (omitting minor details) is as follows:—A six-
parted perianth in two rows, the outer three (sepals) generally regular and
equal in shape; of the inner three (petals or tepals) two are regular, and one,
the labellum very irregular, consisting not only of a petal, but of two
abortive stamens incorporated with it. The column is considered to be made
up of one perfect and three abortive stamens, in inseparable connection with
three styles. By some, however, it is supposed that all the stamens are
confluent with the column and none with the lip.
Page 479
Fig. 192.—Diagram showing the arrangement of parts in an orchid flower. According to Crüger, the
stamens A 2, A 3, should be distinct from the lip. The uppermost figure 2 should have been 1. (See
text.)
In either case it is admitted that there are six stamens in two rows. The first
row consists of one posterior stamen, which is generally perfect, and two
abortive stamens incorporated with the labellum. The second row also
consists of three stamens, all of which are usually abortive and inseparable
from the column. Traces of them may occasionally be met with in the form
of tubercles or wing-like processes from the column. In Cypripedium, while
the ordinary stamen of the outer row is deficient, two of the inner series are
present. The diagram, fig. 192, will serve to show the arrangement of the
parts as above described. + represents the situation of the stem or axis; on
the opposite side is the bract; between these are placed the sepals, one
posterior or next the axis (incorrectly numbered 2 in the plan), two lateral 1,
1; next in order follow the petals, 2, 2, 2, two lateral and somewhat
posterior, one larger (the lip), anterior; the outer series of stamens are
stamens A 2, A 3, should be distinct from the lip. The uppermost figure 2 should have been 1. (See
text.)
In either case it is admitted that there are six stamens in two rows. The first
row consists of one posterior stamen, which is generally perfect, and two
abortive stamens incorporated with the labellum. The second row also
consists of three stamens, all of which are usually abortive and inseparable
from the column. Traces of them may occasionally be met with in the form
of tubercles or wing-like processes from the column. In Cypripedium, while
the ordinary stamen of the outer row is deficient, two of the inner series are
present. The diagram, fig. 192, will serve to show the arrangement of the
parts as above described. + represents the situation of the stem or axis; on
the opposite side is the bract; between these are placed the sepals, one
posterior or next the axis (incorrectly numbered 2 in the plan), two lateral 1,
1; next in order follow the petals, 2, 2, 2, two lateral and somewhat
posterior, one larger (the lip), anterior; the outer series of stamens are
Page 480
represented by A 1, A 2, A 3, the two latter being fused with the labellum; a
1, a 2, a 3 represent the position of the inner verticil of stamens, while s, s, s
denote the three carpels. It is foreign to the purpose of this book to detail
the varied evidence in support of this explanation of the homologies of
orchid flowers.[438] All that can be done in these pages is to set forth the
evidence furnished by teratology as to this matter—evidence for the most
part accumulated and recorded without any special reference to any theory
of orchid structure.
The following details all refer to flowers in which the number of stamens in
orchidaceous plants was increased beyond what is necessary. They are
arranged with reference to the number of adventitious organs, beginning
with those in which the number was smallest, and proceeding thence to
those in which it was greatest. In some cases it has not been possible to
ascertain whether the adventitious organs were really restorations of the
numerical symmetry, substitutions of one part for another, stamen for petal,
&c., or wholly adventitious productions. Unless otherwise stated, the
interpretation put upon the facts thus recorded is that of the present writer,
and not necessarily that of the original observer.
Mr. J. T. Moggridge has described and figured a flower of Ophrys insectifera in which
there was a vestige of a second stamen present, probably one of the inner series fig. 192
(a2).[439] The same observer also records the presence of a second anther between the
lobes of the normal one. This can hardly be referred to either of the typical stamens, but
would seem to be a perverted development of the rostellum.[440]
Rœper is stated by Cramer[441] to have seen a specimen of Orchis morio with two
stamens.
In a flower of Habenaria chlorantha, described by the late Professor Henslow,[442] the
outer three stamens are suppressed, while two of the inner group are present, as happens
normally in Cypripedium.
A flower of Cattleya violacea afforded a similar illustration; but in this case only one of the
inner stamens was developed, and this in the form of a small petal, partly adherent to the
1, a 2, a 3 represent the position of the inner verticil of stamens, while s, s, s
denote the three carpels. It is foreign to the purpose of this book to detail
the varied evidence in support of this explanation of the homologies of
orchid flowers.[438] All that can be done in these pages is to set forth the
evidence furnished by teratology as to this matter—evidence for the most
part accumulated and recorded without any special reference to any theory
of orchid structure.
The following details all refer to flowers in which the number of stamens in
orchidaceous plants was increased beyond what is necessary. They are
arranged with reference to the number of adventitious organs, beginning
with those in which the number was smallest, and proceeding thence to
those in which it was greatest. In some cases it has not been possible to
ascertain whether the adventitious organs were really restorations of the
numerical symmetry, substitutions of one part for another, stamen for petal,
&c., or wholly adventitious productions. Unless otherwise stated, the
interpretation put upon the facts thus recorded is that of the present writer,
and not necessarily that of the original observer.
Mr. J. T. Moggridge has described and figured a flower of Ophrys insectifera in which
there was a vestige of a second stamen present, probably one of the inner series fig. 192
(a2).[439] The same observer also records the presence of a second anther between the
lobes of the normal one. This can hardly be referred to either of the typical stamens, but
would seem to be a perverted development of the rostellum.[440]
Rœper is stated by Cramer[441] to have seen a specimen of Orchis morio with two
stamens.
In a flower of Habenaria chlorantha, described by the late Professor Henslow,[442] the
outer three stamens are suppressed, while two of the inner group are present, as happens
normally in Cypripedium.
A flower of Cattleya violacea afforded a similar illustration; but in this case only one of the
inner stamens was developed, and this in the form of a small petal, partly adherent to the
Page 481
column.
In Dendrobium normale, Falconer, not only is the perianth regular, but the column is
triandrous,[443] the three stamens (according to the diagram of its structure given by
Lindley) pertaining to the outer row.
In a specimen of Dendrobium hœmoglossum kindly forwarded from Ceylon by Mr.
Thwaites there were three stamens present, of which one posterior belonged to the outer
series A 1, and two lateral to the inner a 1, a 2, fig. 192.
M. His observed, several years in succession, some flowers of a species of Ophrys with
three sepals, no lateral petals, one lip, and three perfect stamens. In this case probably the
two supernumerary stamens were petals which had assumed an anther-like character.
Wydler describes a flower of Ophrys aranifera in which one outer and two inner stamens
were present.[444] I have myself met with three such flowers in the same species. The
stamens present were A 1, a 1, a 2.
Dr. J. E. Gray exhibited at the Botanical Society of London, in August, 1843, a specimen of
Ophrys apifera with a triandrous column, the supernumerary anthers belonging, apparently,
to the inner whorl.
In his 'Catalogue of the Plants of South Kent,' p. 56, tab. iv, f. 16, the Rev. G. E. Smith
describes and figures a flower of O. aranifera with a triandrous column, seemingly of the
same kind as that spoken of by Dr. Gray.
Mr. Moggridge met with a triandrous flower in the same species, and refers the appearance
to "a fusion of two flowers, accompanied by suppression and modification."[445] As,
however, no details are given in support of this opinion, it may be conjectured that the two
additional stamens were members of the inner whorl a 1, a 2, and thus the conformation
would be the same as in the flowers just mentioned. The figures given by Mr. Moggridge
bear out this latter view, while they lend no support to the hypothesis advanced by him.
Nevertheless, no decided opinion can be pronounced by those who have not had the
opportunity of examining the flowers in question.
Alphonse de Candolle[446] figures a flower of Maxillaria in exactly the same condition, so
far as the stamens are concerned, as in the Ophrys flowers just mentioned. It is curious to
observe that in many of these cases the two lateral petals are suppressed.
Von Martius mentions the occurrence of three anthers (naturaliter conformatæ) in Orchis
morio.[447] Richard, as cited by Moquin-Tandon, Lindley, and others, describes and
figures a peloria of Orchis latifolia with regular triandrous flowers.[448]
In Dendrobium normale, Falconer, not only is the perianth regular, but the column is
triandrous,[443] the three stamens (according to the diagram of its structure given by
Lindley) pertaining to the outer row.
In a specimen of Dendrobium hœmoglossum kindly forwarded from Ceylon by Mr.
Thwaites there were three stamens present, of which one posterior belonged to the outer
series A 1, and two lateral to the inner a 1, a 2, fig. 192.
M. His observed, several years in succession, some flowers of a species of Ophrys with
three sepals, no lateral petals, one lip, and three perfect stamens. In this case probably the
two supernumerary stamens were petals which had assumed an anther-like character.
Wydler describes a flower of Ophrys aranifera in which one outer and two inner stamens
were present.[444] I have myself met with three such flowers in the same species. The
stamens present were A 1, a 1, a 2.
Dr. J. E. Gray exhibited at the Botanical Society of London, in August, 1843, a specimen of
Ophrys apifera with a triandrous column, the supernumerary anthers belonging, apparently,
to the inner whorl.
In his 'Catalogue of the Plants of South Kent,' p. 56, tab. iv, f. 16, the Rev. G. E. Smith
describes and figures a flower of O. aranifera with a triandrous column, seemingly of the
same kind as that spoken of by Dr. Gray.
Mr. Moggridge met with a triandrous flower in the same species, and refers the appearance
to "a fusion of two flowers, accompanied by suppression and modification."[445] As,
however, no details are given in support of this opinion, it may be conjectured that the two
additional stamens were members of the inner whorl a 1, a 2, and thus the conformation
would be the same as in the flowers just mentioned. The figures given by Mr. Moggridge
bear out this latter view, while they lend no support to the hypothesis advanced by him.
Nevertheless, no decided opinion can be pronounced by those who have not had the
opportunity of examining the flowers in question.
Alphonse de Candolle[446] figures a flower of Maxillaria in exactly the same condition, so
far as the stamens are concerned, as in the Ophrys flowers just mentioned. It is curious to
observe that in many of these cases the two lateral petals are suppressed.
Von Martius mentions the occurrence of three anthers (naturaliter conformatæ) in Orchis
morio.[447] Richard, as cited by Moquin-Tandon, Lindley, and others, describes and
figures a peloria of Orchis latifolia with regular triandrous flowers.[448]
Page 482
The writer has examined, in the Royal Gardens at Kew, a flower of Cattleya crispa in
which were three stamens, the central one normal; the two lateral ones, belonging probably
to the inner whorl, were in appearance like the lateral petals, and one of them was adherent
to the central perfect column. Duchartre[449] mentions a flower of Cattleya Forbesii in
which there were two labella in addition to the ordinary one, the column being in its
normal condition. From the analogy of other cases it would appear as if the additional
labella in this instance were the representatives of two stamens of the outer whorl. Beer
likewise has put on record the existence of a triandrous Cattleya.[450]
A specimen of Catasetum eburneum forwarded by Mr. Wilson Saunders was normal so far
as the sepals and two lateral petals were concerned, but the anterior petal or labellum was
flat and in form quite like the two lateral ones; the column was normal and in the situation
of the two anterior stamens of the outer series A 2, A 3, were two labella of the usual form
(fig. 156, p. 291). Perhaps the Oncidium represented at p. 68, fig. 29, may also be
explained on the supposition that the two lateral lobes of the labellum in this flower were
the representatives of stamens.
In Fig. 193 is shown the arrangement of parts in a flower of Ophrys aranifera. Here there
were three sepals, two lateral petals, one of which was adherent to the side of the column;
the central labellum was seemingly deficient, but there were two pseudo-labella placed
laterally in the position of the two antero-lateral stamens of the outer series (A 2, A 3).
Within these was another perfect stamen occupying the position of the anterior stamen of
the inner series (a 3). In another flower of the same species, gathered at the same time (fig.
194), there were three sepals not at all different from those of the normal flower. The three
petals next in succession were also, in form and position, in their ordinary state. In colour,
however, the two upper lateral petals differed from what is customary, in having the same
purplish-brown tint which characterises the lip. Within these petals, at the upper part of the
flower, there was the ordinary column, and at the opposite side, alternating with the petals
before mentioned, two additional lip-like petals, one provided with a half-anther containing
a single perfectly formed pollen-mass (A 2, A 3). It is, perhaps, worthy of notice that the
arrangement of the coloured spots on the true labellum, and that on the adventitious lips,
replacing the two lower of the outer stamens, were not of a similar character. The
supernumerary lips had the π-shaped marking which is so common in this species, while
the true lip was, as to its spots, much more like O. apifera. Alternating with this last whorl
were three columns, all apparently perfectly formed and differing only from the ordinary
one in their smaller size and corresponding to a 1, a 2, a 3. The ovary in this flower was
two-celled, with four parietal placentas, thus giving an appearance as though there had
been a fusion of two or more flowers associated with suppression and other changes. The
position of the supernumerary organs and the absence of any positive sign of fusion in the
bracts or other part of the flower, seemed, however, to negative the idea of fusion.[451]
which were three stamens, the central one normal; the two lateral ones, belonging probably
to the inner whorl, were in appearance like the lateral petals, and one of them was adherent
to the central perfect column. Duchartre[449] mentions a flower of Cattleya Forbesii in
which there were two labella in addition to the ordinary one, the column being in its
normal condition. From the analogy of other cases it would appear as if the additional
labella in this instance were the representatives of two stamens of the outer whorl. Beer
likewise has put on record the existence of a triandrous Cattleya.[450]
A specimen of Catasetum eburneum forwarded by Mr. Wilson Saunders was normal so far
as the sepals and two lateral petals were concerned, but the anterior petal or labellum was
flat and in form quite like the two lateral ones; the column was normal and in the situation
of the two anterior stamens of the outer series A 2, A 3, were two labella of the usual form
(fig. 156, p. 291). Perhaps the Oncidium represented at p. 68, fig. 29, may also be
explained on the supposition that the two lateral lobes of the labellum in this flower were
the representatives of stamens.
In Fig. 193 is shown the arrangement of parts in a flower of Ophrys aranifera. Here there
were three sepals, two lateral petals, one of which was adherent to the side of the column;
the central labellum was seemingly deficient, but there were two pseudo-labella placed
laterally in the position of the two antero-lateral stamens of the outer series (A 2, A 3).
Within these was another perfect stamen occupying the position of the anterior stamen of
the inner series (a 3). In another flower of the same species, gathered at the same time (fig.
194), there were three sepals not at all different from those of the normal flower. The three
petals next in succession were also, in form and position, in their ordinary state. In colour,
however, the two upper lateral petals differed from what is customary, in having the same
purplish-brown tint which characterises the lip. Within these petals, at the upper part of the
flower, there was the ordinary column, and at the opposite side, alternating with the petals
before mentioned, two additional lip-like petals, one provided with a half-anther containing
a single perfectly formed pollen-mass (A 2, A 3). It is, perhaps, worthy of notice that the
arrangement of the coloured spots on the true labellum, and that on the adventitious lips,
replacing the two lower of the outer stamens, were not of a similar character. The
supernumerary lips had the π-shaped marking which is so common in this species, while
the true lip was, as to its spots, much more like O. apifera. Alternating with this last whorl
were three columns, all apparently perfectly formed and differing only from the ordinary
one in their smaller size and corresponding to a 1, a 2, a 3. The ovary in this flower was
two-celled, with four parietal placentas, thus giving an appearance as though there had
been a fusion of two or more flowers associated with suppression and other changes. The
position of the supernumerary organs and the absence of any positive sign of fusion in the
bracts or other part of the flower, seemed, however, to negative the idea of fusion.[451]
Page 483
Fig. 193.—Diagram showing the arrangement of parts in a malformed flower of Ophrys
aranifera (see p. 384).
aranifera (see p. 384).
Page 484
Fig. 194.—Malformed flower of Ophrys aranifera with two supernumerary lips and three
additional stamens.
A similar illustration, for a knowledge of which the writer is indebted to the kindness of
Professor Asa Gray and Mr. Darwin, occurred in some specimens of Pogonia
ophioglossoides collected by Dr. J. H. Paine in a bog near Utica, New York. It will be seen
from the following description that these flowers presented an almost precisely similar
condition to those of the Ophrys aranifera just mentioned. "The peculiarities of these
flowers," writes Professor Gray, "are that they have three labella, and that the column is
resolved into small petaloid organs. The blossom is normal as to the proper perianth,
except that the labellum is unusually papillose, bearded almost to the base. The points of
interest are, first, that the two accessory labella are just in the position of the two
suppressed stamens of the outer series, viz. of a2 and a3, as represented in the diagram, fig.
192; and there is a small petaloid body on the other side of the flower, answering to the
other stamen, a1. Secondly, in one of the blossoms, and less distinctly in another, two
lateral stamens of the inner series (a1 and a2) are represented each by a slender naked
filament. There are remaining petaloid bodies enough to answer for the third stamen of the
inner series and for the stigmas, but their order is not well to be made out in the dried
specimens." It may here be mentioned that Isochilus is normally triandrous.
A tetrandrous flower of Cypripedium has also been recorded.
In Isochilus, according to Cruger, there are often five stamens, and there are several,
besides those already mentioned, in which six more or less perfect stamens have been seen
—of these the following may be taken as illustrations. A hexandrous flower of Orchis
militaris has been recorded by Kirschleger,[452] and in the accompanying diagram (fig.
195), from Cramer,[453] of a monstrous flower of Orchis mascula, there is one perfect
stamen of the outer row and two lip-like stamens of the same series, while the inner verticil
comprises one perfect and two abortive stamens.
additional stamens.
A similar illustration, for a knowledge of which the writer is indebted to the kindness of
Professor Asa Gray and Mr. Darwin, occurred in some specimens of Pogonia
ophioglossoides collected by Dr. J. H. Paine in a bog near Utica, New York. It will be seen
from the following description that these flowers presented an almost precisely similar
condition to those of the Ophrys aranifera just mentioned. "The peculiarities of these
flowers," writes Professor Gray, "are that they have three labella, and that the column is
resolved into small petaloid organs. The blossom is normal as to the proper perianth,
except that the labellum is unusually papillose, bearded almost to the base. The points of
interest are, first, that the two accessory labella are just in the position of the two
suppressed stamens of the outer series, viz. of a2 and a3, as represented in the diagram, fig.
192; and there is a small petaloid body on the other side of the flower, answering to the
other stamen, a1. Secondly, in one of the blossoms, and less distinctly in another, two
lateral stamens of the inner series (a1 and a2) are represented each by a slender naked
filament. There are remaining petaloid bodies enough to answer for the third stamen of the
inner series and for the stigmas, but their order is not well to be made out in the dried
specimens." It may here be mentioned that Isochilus is normally triandrous.
A tetrandrous flower of Cypripedium has also been recorded.
In Isochilus, according to Cruger, there are often five stamens, and there are several,
besides those already mentioned, in which six more or less perfect stamens have been seen
—of these the following may be taken as illustrations. A hexandrous flower of Orchis
militaris has been recorded by Kirschleger,[452] and in the accompanying diagram (fig.
195), from Cramer,[453] of a monstrous flower of Orchis mascula, there is one perfect
stamen of the outer row and two lip-like stamens of the same series, while the inner verticil
comprises one perfect and two abortive stamens.
Page 485
Fig. 195.—Diagram of flower of Orchis mascula with two additional lips, two perfect and
two imperfect stamens (after Cramer).
Morren[454] describes some flowers of Orchis morio in which there were three sepals,
three petals, and within the latter two other ternary series of petals; this would seem to be a
case of petalody of all six stamens. Morren, however, seems to have considered the
additional segments as repetitions of the corolline whorl, though he describes a central
mass as the column bearing a "souvenir of the anther." Nevertheless, there is no decisive
evidence either in his figure or his description in support of his opinion as to the nature of
the central mass, which might be a distorted condition of the styles, or, as is more probable,
a rudimentary and irregular flower. Morren also describes another flower of the same plant
in which there were three sepals, two lateral petals partially lip-like in aspect, a third
labellum normal, two additional labella representing the two anterior stamens of the outer
whorl, while more or less developed rudiments of the remaining four stamens also exist.
While, in most cases, the supernumerary stamens can, by reason of their relative position,
their complete or partial antheriferous nature, be safely referred to one or other of the six
stamens, making up a typical orchid flower, there are other specimens in which the
additional stamens are altogether adventitious, and do not admit of reference to the
homologue. Thus it was in a specimen of Odontoglossum Alexandræ examined by the
writer, and in which, within a normally constructed perianth, there were six columns, all
polliniferous, but arranged in so confused and complicated a manner that it was impossible
two imperfect stamens (after Cramer).
Morren[454] describes some flowers of Orchis morio in which there were three sepals,
three petals, and within the latter two other ternary series of petals; this would seem to be a
case of petalody of all six stamens. Morren, however, seems to have considered the
additional segments as repetitions of the corolline whorl, though he describes a central
mass as the column bearing a "souvenir of the anther." Nevertheless, there is no decisive
evidence either in his figure or his description in support of his opinion as to the nature of
the central mass, which might be a distorted condition of the styles, or, as is more probable,
a rudimentary and irregular flower. Morren also describes another flower of the same plant
in which there were three sepals, two lateral petals partially lip-like in aspect, a third
labellum normal, two additional labella representing the two anterior stamens of the outer
whorl, while more or less developed rudiments of the remaining four stamens also exist.
While, in most cases, the supernumerary stamens can, by reason of their relative position,
their complete or partial antheriferous nature, be safely referred to one or other of the six
stamens, making up a typical orchid flower, there are other specimens in which the
additional stamens are altogether adventitious, and do not admit of reference to the
homologue. Thus it was in a specimen of Odontoglossum Alexandræ examined by the
writer, and in which, within a normally constructed perianth, there were six columns, all
polliniferous, but arranged in so confused and complicated a manner that it was impossible
Page 486
to make out any definite relation in their position. There was nothing to indicate a fusion of
flowers, but rather an extension of the centre of the flower, and consequent displacement of
the stamens, &c. Again, the existence of adventitious stamens does not necessarily imply
the development of organs usually suppressed, inasmuch as they may result from the
assumption by the lateral petals of staminal characteristics.
Nevertheless, as far as teratology is concerned, specimens may be found in which some or
all of the usually suppressed stamens of Orchidaceæ may be found. These stamens may be
all perfect (polliniferous), or, as is more frequently the case, more or less petal-like.
Moreover, when the stamens are petalodic, the form assumed is usually that of the
labellum.
The presence of stamens in undue numbers in orchids is very generally, but not always,
attended by some coincident malformation, of which the most frequent is cohesion of two
or more sepals, and consequent displacement or adhesion of one petal to the side of the
column. Petalody of the styles and median prolification are also sometimes found in
association with an augmented number of stamens.
flowers, but rather an extension of the centre of the flower, and consequent displacement of
the stamens, &c. Again, the existence of adventitious stamens does not necessarily imply
the development of organs usually suppressed, inasmuch as they may result from the
assumption by the lateral petals of staminal characteristics.
Nevertheless, as far as teratology is concerned, specimens may be found in which some or
all of the usually suppressed stamens of Orchidaceæ may be found. These stamens may be
all perfect (polliniferous), or, as is more frequently the case, more or less petal-like.
Moreover, when the stamens are petalodic, the form assumed is usually that of the
labellum.
The presence of stamens in undue numbers in orchids is very generally, but not always,
attended by some coincident malformation, of which the most frequent is cohesion of two
or more sepals, and consequent displacement or adhesion of one petal to the side of the
column. Petalody of the styles and median prolification are also sometimes found in
association with an augmented number of stamens.
Page 487
Fig. 196.—Increased number of carpels, tulip.
Page 488
Fig. 197.—Fruit of St. Valery apple cut lengthwise.
Pleiotaxy of the gynœcium.—An increase in the number of whorls of
which the pistil consists is not of very frequent occurrence. Generally after
the formation of the whorl of carpels, the energy of the growing point
ceases, or if by chance it be continued, the result is more generally the
production of a new flower-bud (median prolification) than the repetition of
the carpellary series. It is necessary also to distinguish between the veritable
augmentation of the pistil and the semblance of it, brought about by the
substitution of carpels for some other organs, as pistillody of the stamens,
and even of the segments of the perianth, is not very unfrequent, as has
already been stated under the head of substitution. Again, the increased
number of carpels which is sometimes met with in such flowers, as
Magnolia or Delphinium, where the ovaries are arranged in spiral series, is
not strictly referable to the present category.
The orange is one of the plants most frequently subject to an augmentation
in the number of carpellary whorls; sometimes this is due to the stamens
assuming the guise of carpels, but at other times the increase occurs without
any alteration in the stamens or other organs. If the adventitious carpels be
exposed, they are covered with yellow rind, while those portions that are
covered by the primary carpels are destitute of rind. Some varieties of the
double tulip are very subject to a similar change, but, in this case, the petals
and the stamens very frequently become more or less carpellary in their
nature. Fig. 196 represents an increased number of whorls of carpels in the
variety called "rex rubrorum," the segments of the perianth having been
removed.
In the St. Valery apple, already referred to, there is a second whorl of
carpels above the first, a fact which has been made use of to explain the
similar structure of the pomegranate.
Pleiotaxy of the gynœcium.—An increase in the number of whorls of
which the pistil consists is not of very frequent occurrence. Generally after
the formation of the whorl of carpels, the energy of the growing point
ceases, or if by chance it be continued, the result is more generally the
production of a new flower-bud (median prolification) than the repetition of
the carpellary series. It is necessary also to distinguish between the veritable
augmentation of the pistil and the semblance of it, brought about by the
substitution of carpels for some other organs, as pistillody of the stamens,
and even of the segments of the perianth, is not very unfrequent, as has
already been stated under the head of substitution. Again, the increased
number of carpels which is sometimes met with in such flowers, as
Magnolia or Delphinium, where the ovaries are arranged in spiral series, is
not strictly referable to the present category.
The orange is one of the plants most frequently subject to an augmentation
in the number of carpellary whorls; sometimes this is due to the stamens
assuming the guise of carpels, but at other times the increase occurs without
any alteration in the stamens or other organs. If the adventitious carpels be
exposed, they are covered with yellow rind, while those portions that are
covered by the primary carpels are destitute of rind. Some varieties of the
double tulip are very subject to a similar change, but, in this case, the petals
and the stamens very frequently become more or less carpellary in their
nature. Fig. 196 represents an increased number of whorls of carpels in the
variety called "rex rubrorum," the segments of the perianth having been
removed.
In the St. Valery apple, already referred to, there is a second whorl of
carpels above the first, a fact which has been made use of to explain the
similar structure of the pomegranate.
Page 489
The tomato (Lycopersicum esculentum) is another plant in which an
adventitious series is frequently produced, and generally in combination
with the primary series.
In the Chinese primrose (Primula sinensis) a supernumerary whorl is
frequently met with, generally associated with other changes in the
construction and arrangement of the parts of the flower.
M. de Candolle[455] mentions a flower of Gentiana purpurea with four
carpels in one series, and five others in the circle immediately above them.
Wigand[456] alludes to an instance wherein there was a second pair of
carpels above the first in Vinca herbacea. Dr. Sankey has forwarded flowers
of a Pelargonium having a double series of carpels, eight in the outer row,
five in the inner, and this condition is stated to exist in the flowers of the
same plant for two years consecutively. In Aquilegia I have met with a
similar increase in the whorls of carpels.[457] Meissner records a similar
augmentation in Polygonum orientale.[458]
Wigand[459] describes and figures a flower of Vinca minor, in which there
were two carpels intervening between the ordinary pair, and a similar
illustration has been observed by the writer in Allamanda cathartica.
Eichler[460] has put on record a similar case in a capparid.
Marchand[461] mentions a polycarpellary berberid (Epimedium
Musschianum). The supernumerary carpels in this flower were placed on a
short axis, which originated in the axils of the stamens, and as these latter
organs were present in their usual number and position, the adventitious
carpels could not be considered as resulting from a transformation, or
substitution of carpels for stamens.
adventitious series is frequently produced, and generally in combination
with the primary series.
In the Chinese primrose (Primula sinensis) a supernumerary whorl is
frequently met with, generally associated with other changes in the
construction and arrangement of the parts of the flower.
M. de Candolle[455] mentions a flower of Gentiana purpurea with four
carpels in one series, and five others in the circle immediately above them.
Wigand[456] alludes to an instance wherein there was a second pair of
carpels above the first in Vinca herbacea. Dr. Sankey has forwarded flowers
of a Pelargonium having a double series of carpels, eight in the outer row,
five in the inner, and this condition is stated to exist in the flowers of the
same plant for two years consecutively. In Aquilegia I have met with a
similar increase in the whorls of carpels.[457] Meissner records a similar
augmentation in Polygonum orientale.[458]
Wigand[459] describes and figures a flower of Vinca minor, in which there
were two carpels intervening between the ordinary pair, and a similar
illustration has been observed by the writer in Allamanda cathartica.
Eichler[460] has put on record a similar case in a capparid.
Marchand[461] mentions a polycarpellary berberid (Epimedium
Musschianum). The supernumerary carpels in this flower were placed on a
short axis, which originated in the axils of the stamens, and as these latter
organs were present in their usual number and position, the adventitious
carpels could not be considered as resulting from a transformation, or
substitution of carpels for stamens.
Page 490
Lastly, the instance cited by Dr. Allman[462] in Saxifraga Geum may be
alluded to. Here there was a row of adventitious carpels between the
stamens and pistils, the backs of the carpels being turned towards the axis
of the flowers. Dr. Allman explains the presence of the supernumerary parts
by the supposed production of a whorl of secondary axes between the
stamens and the centre of the flower. These axes are further supposed to
bear imperfect flowers, of which the additional carpels are the only traces,
but this explanation seems forced.
In addition to the references already cited the following may be given:
Duchartre, 'Ann. Sc. Nat.,' 4 ser., vii, p. 23 (Tulip).
Ferrari, 'Hesperides,' pp. 271, 395, 405. Duchartre, 'Ann. Sc. Nat.,' 4 ser., 1844, vol. i, p.
294. Maout, 'Leçons Elément.,' vol. ii, pp. 488–9. Clos, 'Ann. Sc. Nat.,' 1865, p. 317
(Citrus Aurantium).
Clos, 'Bull. Soc. Bot. Fr.,' vol. xiii; 'Rev. Bibl.,' p. 75. Pasquale, 'Reddicont Accad. Sc. Fis.
e Math. Napoli.' Octr. 1866 (Solanum Lycopersicum).
On the general subject of multiplication, in addition to previous citations,
the reader may be referred to A. P. de Candolle, 'Théorie Elément. Bot.,' ed.
3, p. 89.
Increased number of flowers in an inflorescence.—This happens
generally as a result of over luxuriant growth, and scarcely demands notice
here, being rather referable to variation than to malformation. The increased
number of florets in the spikelets of some grasses has already been alluded
to (p. 351). Thus spikelets of wheat occasionally produce more than the
three florets which are proper to them.[463] It will be remembered that in
this as in many other grasses there are rudimentary florets, and it is no
matter for surprise that these florets should occasionally be fully developed.
FOOTNOTES:
alluded to. Here there was a row of adventitious carpels between the
stamens and pistils, the backs of the carpels being turned towards the axis
of the flowers. Dr. Allman explains the presence of the supernumerary parts
by the supposed production of a whorl of secondary axes between the
stamens and the centre of the flower. These axes are further supposed to
bear imperfect flowers, of which the additional carpels are the only traces,
but this explanation seems forced.
In addition to the references already cited the following may be given:
Duchartre, 'Ann. Sc. Nat.,' 4 ser., vii, p. 23 (Tulip).
Ferrari, 'Hesperides,' pp. 271, 395, 405. Duchartre, 'Ann. Sc. Nat.,' 4 ser., 1844, vol. i, p.
294. Maout, 'Leçons Elément.,' vol. ii, pp. 488–9. Clos, 'Ann. Sc. Nat.,' 1865, p. 317
(Citrus Aurantium).
Clos, 'Bull. Soc. Bot. Fr.,' vol. xiii; 'Rev. Bibl.,' p. 75. Pasquale, 'Reddicont Accad. Sc. Fis.
e Math. Napoli.' Octr. 1866 (Solanum Lycopersicum).
On the general subject of multiplication, in addition to previous citations,
the reader may be referred to A. P. de Candolle, 'Théorie Elément. Bot.,' ed.
3, p. 89.
Increased number of flowers in an inflorescence.—This happens
generally as a result of over luxuriant growth, and scarcely demands notice
here, being rather referable to variation than to malformation. The increased
number of florets in the spikelets of some grasses has already been alluded
to (p. 351). Thus spikelets of wheat occasionally produce more than the
three florets which are proper to them.[463] It will be remembered that in
this as in many other grasses there are rudimentary florets, and it is no
matter for surprise that these florets should occasionally be fully developed.
FOOTNOTES:
Page 491
[392] 'Bull. Soc. Bot. Fr.,' vol. vii, 1860, p. 587.
[393] 'Fragment. Phyt. Austral.,' part xx, p. 270.
[394] 'Bull. Acad. Belg.,' xvi, pt. i, p. 60, "Fuchsia," p. 125, c. ic.
[395] "Théorie de la feuille," 'Arch. des Sciences Bibl. Univers.,' 1868.
[396] See Engelmann, 'De Antholysi,' p. 16, section 12.
[397] Verhandl. des Botanisch. Vereins Brandenburg,' 1859, 1 heft.
[398] See Henslow. 'Mag. Nat, Hist.' 1832, vol. v, p. 429.
[399] 'Phytologist,' September, 1857.
[400] Seemann's 'Journal of Botany,' iv, p. 168, t. 47, f. 3.
[401] 'Illust. Hortic.,' 1866, misc., p. 97.
[402] See Fresenius, 'Mus. Senkenb.,' bd. 2, p. 43. Schlechtendal, 'Bot. Zeit.,' iv, pp. 403,
492, Veronica tetrandra.
[403] 'Flora,' 1865, tab. 6, fig. 8.
[404] 'Org. Veget.,' t. i, p. 497, pl. 42, f. 3.
[405] 'El. Ter. Veg.,' p. 354.
[406] Cited in "Rev. Bibl." of 'Bull. Soc. Bot. Fr.,' 1866, p. 171.
[407] Loc. cit., 351.
[408] 'Mém. Acad. Toulous.,' vi, 1862, ex 'Bull. Soc. Bot. Fr.,' "Rev. Bibl.," vol. ix, 1862. p.
127.
[409] 'Flora.' 1857. p. 289.
[410] L. c., p. 354.
[411] Giraud, 'Ed. Phil. Mag.,' Dec., 1839.
[412] See Cerasus Caproniana, D. C. 'Plant. Rar. Hort. Genev.,' tab. 18.
[413] Nees, 'Linnæa,' v, p. 679, tab. 11 (Schœnodorus).
[414] 'Gard. Chron.,' 1852, p. 452.
[393] 'Fragment. Phyt. Austral.,' part xx, p. 270.
[394] 'Bull. Acad. Belg.,' xvi, pt. i, p. 60, "Fuchsia," p. 125, c. ic.
[395] "Théorie de la feuille," 'Arch. des Sciences Bibl. Univers.,' 1868.
[396] See Engelmann, 'De Antholysi,' p. 16, section 12.
[397] Verhandl. des Botanisch. Vereins Brandenburg,' 1859, 1 heft.
[398] See Henslow. 'Mag. Nat, Hist.' 1832, vol. v, p. 429.
[399] 'Phytologist,' September, 1857.
[400] Seemann's 'Journal of Botany,' iv, p. 168, t. 47, f. 3.
[401] 'Illust. Hortic.,' 1866, misc., p. 97.
[402] See Fresenius, 'Mus. Senkenb.,' bd. 2, p. 43. Schlechtendal, 'Bot. Zeit.,' iv, pp. 403,
492, Veronica tetrandra.
[403] 'Flora,' 1865, tab. 6, fig. 8.
[404] 'Org. Veget.,' t. i, p. 497, pl. 42, f. 3.
[405] 'El. Ter. Veg.,' p. 354.
[406] Cited in "Rev. Bibl." of 'Bull. Soc. Bot. Fr.,' 1866, p. 171.
[407] Loc. cit., 351.
[408] 'Mém. Acad. Toulous.,' vi, 1862, ex 'Bull. Soc. Bot. Fr.,' "Rev. Bibl.," vol. ix, 1862. p.
127.
[409] 'Flora.' 1857. p. 289.
[410] L. c., p. 354.
[411] Giraud, 'Ed. Phil. Mag.,' Dec., 1839.
[412] See Cerasus Caproniana, D. C. 'Plant. Rar. Hort. Genev.,' tab. 18.
[413] Nees, 'Linnæa,' v, p. 679, tab. 11 (Schœnodorus).
[414] 'Gard. Chron.,' 1852, p. 452.
Page 492
[415] See Cramer, 'Bildungsabweich.' pp. 16, 24.
[416] 'Linnæa,' 1842, p. 389, c. ic.
[417] Seemann's 'Journal of Botany,' 1867, vol. v, p. 158.
[418] Cramer, 'Bildungsabweich,' p. 66, Astrantia major, Eryngium, to which may be added
Daucus, Heracleum, &c.
[419] See also Reinwardt, 'Nov. Act. Acad. Nat. Cur.,' 12, 1, 37; and Masters, 'Journ. Linn.
Soc.,' vi, p. 24.
[420] 'Organog. Veget.,' tab. 53.
[421] 'Missbild.,' p. 206.
[422] Ehrenberg, 'Flora,' 1846, p. 704.
[423] 'Flora,' 1860, tab. 7.
[424] 'Ann. Sc. Nat.,' 3 ser., t. x, p. 207.
[425] 'Mem. Mus.,' xii. t. 17.
[426] 'Nov. Act. Acad. Nat. Cur.,' xv, tab. xxviii, f. 3; 'Bot. Mag.,' t. 1622. "Caryophyllus
spicam frumenti referens." A similar malformation in Dianthus barbatus is not uncommon. It
has lately been introduced into gardens under the name of Dianthus "mousseux," but is not
likely to find favour with gardeners.
[427] 'Bull. Soc. Bot. France,' t. vi, 1859. p. 268.
[428] Weber, 'Verhandl. Nat. Hist. Vereins. Rhein. Pruss.,' 1860.
[429] 'Bot. Zeit.,' 1848, p. 217.
[430] 'De Anthol.,' p. 17, § 12.
[431] 'Linnæa,' vol. ii, 1827, p. 85.
[432] 'De Antholysi,' p. 17, tab. iii, f. 15, 16; Weinmann, 'Phytanth. iconogr.,' nro. 292.
[433] See Hildebrand, 'Bot. Zeit.,' 1862, p. 209, tab. viii; Cramer, 'Bildungsabweich.,' p. 7,
tab. xiii; Engelmann, 'De Antholysi,' p. 18, &c. For similar changes in Gagea arvensis see
Wirtgen, 'Flora,' 1838, t. xxi. p. 350, and 'Flora.' 1846, p. 353. Some of these are cases of
synanthy.
[416] 'Linnæa,' 1842, p. 389, c. ic.
[417] Seemann's 'Journal of Botany,' 1867, vol. v, p. 158.
[418] Cramer, 'Bildungsabweich,' p. 66, Astrantia major, Eryngium, to which may be added
Daucus, Heracleum, &c.
[419] See also Reinwardt, 'Nov. Act. Acad. Nat. Cur.,' 12, 1, 37; and Masters, 'Journ. Linn.
Soc.,' vi, p. 24.
[420] 'Organog. Veget.,' tab. 53.
[421] 'Missbild.,' p. 206.
[422] Ehrenberg, 'Flora,' 1846, p. 704.
[423] 'Flora,' 1860, tab. 7.
[424] 'Ann. Sc. Nat.,' 3 ser., t. x, p. 207.
[425] 'Mem. Mus.,' xii. t. 17.
[426] 'Nov. Act. Acad. Nat. Cur.,' xv, tab. xxviii, f. 3; 'Bot. Mag.,' t. 1622. "Caryophyllus
spicam frumenti referens." A similar malformation in Dianthus barbatus is not uncommon. It
has lately been introduced into gardens under the name of Dianthus "mousseux," but is not
likely to find favour with gardeners.
[427] 'Bull. Soc. Bot. France,' t. vi, 1859. p. 268.
[428] Weber, 'Verhandl. Nat. Hist. Vereins. Rhein. Pruss.,' 1860.
[429] 'Bot. Zeit.,' 1848, p. 217.
[430] 'De Anthol.,' p. 17, § 12.
[431] 'Linnæa,' vol. ii, 1827, p. 85.
[432] 'De Antholysi,' p. 17, tab. iii, f. 15, 16; Weinmann, 'Phytanth. iconogr.,' nro. 292.
[433] See Hildebrand, 'Bot. Zeit.,' 1862, p. 209, tab. viii; Cramer, 'Bildungsabweich.,' p. 7,
tab. xiii; Engelmann, 'De Antholysi,' p. 18, &c. For similar changes in Gagea arvensis see
Wirtgen, 'Flora,' 1838, t. xxi. p. 350, and 'Flora.' 1846, p. 353. Some of these are cases of
synanthy.
Page 493
[434] Schlechtendal, 'Bot. Zeit.,' xx, 1862, p. 301.
[435] 'Phil. Bot.,' § 126.
[436] C. Morren, 'Bull. Acad. Belg.,' xix, part ii, p. 17.
[437] 'Seemann's Journal of Botany,' iii, p. 354.
[438] On this point the reader will find an excellent summary in Lindley's 'Vegetable
Kingdom,' cd. iii, p. 183a, and in Darwin, 'Fertilisation of Orchids,' p. 292. See also
Crüger,'Journ. Linn. Soc.,' t. viii, p. 134.
[439] 'Seemann's Journal of Botany,' vol. iv, p. 168, tab. 47.
[440] Ibid., t. iv. 1866, p. 168, t. xlvii, f. 1.
[441] 'Bildungsabweich,' p. 8; see also 'Bot. Zeit.,' 1852, p. 425.
[442] 'Journ. Linn. Soc.,' t. ii, p. 104. tab. 1, fig. B.
[443] Lindl., "Orchid. Ind.," 'Jour. Linn. Soc.,' iii, p. 9.
[444] 'Arch. Bot.,' ii, p. 300, tab. xvi, f. 11.
[445] 'Seemann's Journal of Botany,' v, p. 318, tab. lxxii, figs. A 4, 4 a.
[446] "Monstr. Veg.," in 'Neue Denkschrift,' p. 17, tab. vii.
[447] 'Flora,' t. viii, 1825, p. 736.
[448] 'Mem. Soc. d'Hist. Nat.,' ii, 1, p. 212, tab. iii.
[449] 'Bull. Soc. Bot. Fr.,' t. vii, 1860, p. 26.
[450] 'Beitr. Morphol. und Biol. Orchid.,' quoted by Cramer; 'Bildungsabweich,' p. 9.
[451] Masters, 'Journ. Linn. Soc.,' viii, p. 207. See also Rodigas, 'Bull. Soc. Bot. Belg.,' iv, p.
266, for similar changes in Cypripedium Hookeræ.
[452] Kirschleger, 'Flora,' 1844, p. 131.
[453] 'Bildungsabweich,' p. 11, tab. xiv, f. 3.
[454] 'Bull. Acad. Roy. Belg.,' t. xix, part 2, p. 171.
[455] 'Organogr. Végét.,' t. i, p. 509, tab. 40, figs. 6, 7.
[435] 'Phil. Bot.,' § 126.
[436] C. Morren, 'Bull. Acad. Belg.,' xix, part ii, p. 17.
[437] 'Seemann's Journal of Botany,' iii, p. 354.
[438] On this point the reader will find an excellent summary in Lindley's 'Vegetable
Kingdom,' cd. iii, p. 183a, and in Darwin, 'Fertilisation of Orchids,' p. 292. See also
Crüger,'Journ. Linn. Soc.,' t. viii, p. 134.
[439] 'Seemann's Journal of Botany,' vol. iv, p. 168, tab. 47.
[440] Ibid., t. iv. 1866, p. 168, t. xlvii, f. 1.
[441] 'Bildungsabweich,' p. 8; see also 'Bot. Zeit.,' 1852, p. 425.
[442] 'Journ. Linn. Soc.,' t. ii, p. 104. tab. 1, fig. B.
[443] Lindl., "Orchid. Ind.," 'Jour. Linn. Soc.,' iii, p. 9.
[444] 'Arch. Bot.,' ii, p. 300, tab. xvi, f. 11.
[445] 'Seemann's Journal of Botany,' v, p. 318, tab. lxxii, figs. A 4, 4 a.
[446] "Monstr. Veg.," in 'Neue Denkschrift,' p. 17, tab. vii.
[447] 'Flora,' t. viii, 1825, p. 736.
[448] 'Mem. Soc. d'Hist. Nat.,' ii, 1, p. 212, tab. iii.
[449] 'Bull. Soc. Bot. Fr.,' t. vii, 1860, p. 26.
[450] 'Beitr. Morphol. und Biol. Orchid.,' quoted by Cramer; 'Bildungsabweich,' p. 9.
[451] Masters, 'Journ. Linn. Soc.,' viii, p. 207. See also Rodigas, 'Bull. Soc. Bot. Belg.,' iv, p.
266, for similar changes in Cypripedium Hookeræ.
[452] Kirschleger, 'Flora,' 1844, p. 131.
[453] 'Bildungsabweich,' p. 11, tab. xiv, f. 3.
[454] 'Bull. Acad. Roy. Belg.,' t. xix, part 2, p. 171.
[455] 'Organogr. Végét.,' t. i, p. 509, tab. 40, figs. 6, 7.
Page 494
[456] 'Flora,' 1856, p. 715.
[457] 'Linn. Trans.,' t. xxiii, p. 364, tab. 34, fig. 5.
[458] 'Monog. Polygon,' pl. 3, K. f. 12.
[459] 'Flora,' 1856, tab. viii.
[460] Ibid., 1865, tab. ix, f. 6.
[461] 'Adansonia,' vol. iv, 1864, p. 127.
[462] 'Ann. Nat. Hist.,' 1845, vol. xvi, p. 126.
[463] See Schlechtendal, 'Bot. Zeit.,' t. xviii, p. 381 (Triticum); also 'Flora,' t. xiv, 1831, p. 5
(Avena).
PART II.
DIMINISHED NUMBER OF ORGANS.
A diminution in the number of parts is generally due to suppression, using
that word as the equivalent of non-development. It corresponds thus in
meaning with the Fehlschlagen of the Germans, the avortement complète of
Moquin and other French writers. It differs from atrophy, or partial
abortion, inasmuch as the latter terms apply to instances wherein there has
been a partial development, and in which evolution has gone on to a certain
extent, but has, from some cause or other, been checked. These cases will
be found under the head of diminished size of organs. As the word abortion
is used by different authors in different ways, it is the more necessary to be
as precise as possible in the application of the term. In the present work
abortion is used to apply to cases wherein parts have been formed, but
wherein growth has been arrested at a certain stage, and which, therefore,
have either remained in statu quo, while the surrounding parts have
[457] 'Linn. Trans.,' t. xxiii, p. 364, tab. 34, fig. 5.
[458] 'Monog. Polygon,' pl. 3, K. f. 12.
[459] 'Flora,' 1856, tab. viii.
[460] Ibid., 1865, tab. ix, f. 6.
[461] 'Adansonia,' vol. iv, 1864, p. 127.
[462] 'Ann. Nat. Hist.,' 1845, vol. xvi, p. 126.
[463] See Schlechtendal, 'Bot. Zeit.,' t. xviii, p. 381 (Triticum); also 'Flora,' t. xiv, 1831, p. 5
(Avena).
PART II.
DIMINISHED NUMBER OF ORGANS.
A diminution in the number of parts is generally due to suppression, using
that word as the equivalent of non-development. It corresponds thus in
meaning with the Fehlschlagen of the Germans, the avortement complète of
Moquin and other French writers. It differs from atrophy, or partial
abortion, inasmuch as the latter terms apply to instances wherein there has
been a partial development, and in which evolution has gone on to a certain
extent, but has, from some cause or other, been checked. These cases will
be found under the head of diminished size of organs. As the word abortion
is used by different authors in different ways, it is the more necessary to be
as precise as possible in the application of the term. In the present work
abortion is used to apply to cases wherein parts have been formed, but
wherein growth has been arrested at a certain stage, and which, therefore,
have either remained in statu quo, while the surrounding parts have
Page 495
increased, or have, from pressure or other causes, actually diminished in
size.
In practice, however, it is not always possible to discriminate between those
instances in which there has been a true suppression, an absolute non-
development of any particular organ, and those in which it has been formed,
and has grown for a time, but has afterwards ceased to do so, and has been
gradually obliterated by the pressure exercised by the constantly increasing
bulk of adjacent parts, or possibly has become incorporated with them. In
the adult flower the appearances are the same, though the causes may have
been different.
CHAPTER I.
SUPPRESSION OF AXILE ORGANS.
Absolute suppression of the main axis is tantamount to the non-existence of
the plant, so that the terms "acaulescent," "acaulosia," etc, must be
considered relatively only, and must be taken to signify an atrophied or
diminished size of the stem, arising from the non-development of the
internodes.
The absence of lateral branches or divisions of the axis is of frequent
occurrence, and is dependent on such causes as the following:—deficient
supply of nutriment, position against a wall or other obstacle, close
crowding of individual plants, too great or too little light, too rich or too
poor a soil, &c.
Probably the absence of the swollen portion below the flower in the case of
many proliferous roses, double-flowered apples, as already referred to, may
be dependent on the non-development of the extremity of the peduncle or
size.
In practice, however, it is not always possible to discriminate between those
instances in which there has been a true suppression, an absolute non-
development of any particular organ, and those in which it has been formed,
and has grown for a time, but has afterwards ceased to do so, and has been
gradually obliterated by the pressure exercised by the constantly increasing
bulk of adjacent parts, or possibly has become incorporated with them. In
the adult flower the appearances are the same, though the causes may have
been different.
CHAPTER I.
SUPPRESSION OF AXILE ORGANS.
Absolute suppression of the main axis is tantamount to the non-existence of
the plant, so that the terms "acaulescent," "acaulosia," etc, must be
considered relatively only, and must be taken to signify an atrophied or
diminished size of the stem, arising from the non-development of the
internodes.
The absence of lateral branches or divisions of the axis is of frequent
occurrence, and is dependent on such causes as the following:—deficient
supply of nutriment, position against a wall or other obstacle, close
crowding of individual plants, too great or too little light, too rich or too
poor a soil, &c.
Probably the absence of the swollen portion below the flower in the case of
many proliferous roses, double-flowered apples, as already referred to, may
be dependent on the non-development of the extremity of the peduncle or
Page 496
flower-stalk. Thus, in a double-flowered apple recently examined, there was
a sort of involucel of five perfect leaves, then five sepals surrounding an
equal number of petals, numerous stamens, and five styles, but not a trace
of an expanded axis, nor of any portion of the carpels, except the styles. The
views taken as to the nature of this and similar malformations must depend
on the opinion held as to the nature of inferior pistils, and on the share, if
any, that the expanded axis takes in their production. As elsewhere said, the
evidence furnished by teratology is conflicting, but there seems little or
nothing to invalidate the notion that the end of the flower-stalk and the base
of the calyx may, to a varying extent, in different cases, jointly be
concerned in the formation of the so-called calyx-tube and of the inferior
ovary. Obviously it is not proper to apply to all cases where there is an
inferior ovary the same explanation as to how it is brought about.
As these pages are passing through the press, M. Casimir de Candolle has
published a different explanation as to the nature of the hip of the rose,
having been led to his opinion by the conclusion that he has arrived at, that
the leaf is to be considered in the light of a flattened branch, whose upper or
posterior surface is more or less completely atrophied.
According to M. de Candolle, the calyx-tube, in the case of the rose, is
neither a whorl of leaves, nor a concave axis in the ordinary sense in which
those terms are used, but is rather to be considered as a ring-like projection
from an axis arrested in its ulterior development. The secondary projections
from the original one correspond to an equal number of vascular bundles,
and develope into the sepals, petals, stamens, and ovaries. If these organs
remained in a rudimentary condition, the tube of the calyx would be
reduced to the condition of a sheathing leaf. The rose flower, then,
according to M. de Candolle, may be considered as a sheathing leaf, whose
fibro-vascular system is complete, and from which all possible primary
projections are developed.[464]
a sort of involucel of five perfect leaves, then five sepals surrounding an
equal number of petals, numerous stamens, and five styles, but not a trace
of an expanded axis, nor of any portion of the carpels, except the styles. The
views taken as to the nature of this and similar malformations must depend
on the opinion held as to the nature of inferior pistils, and on the share, if
any, that the expanded axis takes in their production. As elsewhere said, the
evidence furnished by teratology is conflicting, but there seems little or
nothing to invalidate the notion that the end of the flower-stalk and the base
of the calyx may, to a varying extent, in different cases, jointly be
concerned in the formation of the so-called calyx-tube and of the inferior
ovary. Obviously it is not proper to apply to all cases where there is an
inferior ovary the same explanation as to how it is brought about.
As these pages are passing through the press, M. Casimir de Candolle has
published a different explanation as to the nature of the hip of the rose,
having been led to his opinion by the conclusion that he has arrived at, that
the leaf is to be considered in the light of a flattened branch, whose upper or
posterior surface is more or less completely atrophied.
According to M. de Candolle, the calyx-tube, in the case of the rose, is
neither a whorl of leaves, nor a concave axis in the ordinary sense in which
those terms are used, but is rather to be considered as a ring-like projection
from an axis arrested in its ulterior development. The secondary projections
from the original one correspond to an equal number of vascular bundles,
and develope into the sepals, petals, stamens, and ovaries. If these organs
remained in a rudimentary condition, the tube of the calyx would be
reduced to the condition of a sheathing leaf. The rose flower, then,
according to M. de Candolle, may be considered as a sheathing leaf, whose
fibro-vascular system is complete, and from which all possible primary
projections are developed.[464]
Page 497
If, as M. de Candolle considers, the leaf and the branch differ merely in the
fact that the vascular system is complete in the latter, and partly atrophied in
the former, it would surely be better to consider the "calyx-tube" of the rose
as a concave axis rather than as a leaf, seeing that he admits the fibro-
vascular system to be complete in the case of the rose.
With reference to this point the reader is referred to Mr. Bentham's account
of the morphology and homologies of the Myrtaceæ in the 'Journal of the
Linnean Society,' vol. x, p. 105. See also ante, pp. 71, 77.
Some doubts also exist as to the nature of the beak or columella of such
fruits as those of Geraniaceæ, Malvaceæ, Umbelliferæ, Euphorbiaceæ, &c.
The nature of the organ in question may probably be different in the several
orders named; at any rate the subject cannot be discussed in this place, and
it is mentioned here because, now and then, it happens that the organ in
question is completely wanting, and hence affords an illustration of
suppression.
FOOTNOTES:
[464] 'Théorie de la feuille.' p. 24.
CHAPTER II.
SUPPRESSION OF FOLIAR ORGANS.
This subject may be considered, according as the separate leaves of the
stem or of the flower are affected, and according as either the number of
members of distinct whorls, or that of the whorls themselves, is diminished.
fact that the vascular system is complete in the latter, and partly atrophied in
the former, it would surely be better to consider the "calyx-tube" of the rose
as a concave axis rather than as a leaf, seeing that he admits the fibro-
vascular system to be complete in the case of the rose.
With reference to this point the reader is referred to Mr. Bentham's account
of the morphology and homologies of the Myrtaceæ in the 'Journal of the
Linnean Society,' vol. x, p. 105. See also ante, pp. 71, 77.
Some doubts also exist as to the nature of the beak or columella of such
fruits as those of Geraniaceæ, Malvaceæ, Umbelliferæ, Euphorbiaceæ, &c.
The nature of the organ in question may probably be different in the several
orders named; at any rate the subject cannot be discussed in this place, and
it is mentioned here because, now and then, it happens that the organ in
question is completely wanting, and hence affords an illustration of
suppression.
FOOTNOTES:
[464] 'Théorie de la feuille.' p. 24.
CHAPTER II.
SUPPRESSION OF FOLIAR ORGANS.
This subject may be considered, according as the separate leaves of the
stem or of the flower are affected, and according as either the number of
members of distinct whorls, or that of the whorls themselves, is diminished.
Page 498
The terms aphylly, meiophylly, and meiotaxy may be employed, according
as the individual leaves are altogether wanting, or with reference to the
diminished number of parts in a whorl, or a decrease in the verticils.
Aphylly.—Entire suppression of the leaves is a rare phenomenon. Under
ordinary circumstances it occurs in most Cactaceæ, in some of the
succulent Euphorbias, and other similar plants, where the epidermal layers
of the stem fulfil the functions of leaves. But even in these plants leaf-like
organs are present in some stage or another of the plant's life.
Partial suppression of the leaf occurs sometimes in compound leaves, some
or other of the leaflets of which are occasionally suppressed. Sometimes, as
Moquin remarks, it is the terminal leaflet which is wanting, when the
appearance is that of Cliffortia, at other times the lateral leaflets are
deficient, as in Citrus or Phyllarthron. Ononis monophylla and Fragaria
monophylla may be cited as instances of the suppression of the lateral
leaflets. If the blade of the leaf disappears entirely, we have then an
analogous condition to that of the phyllodineous acacias.
With reference to the strawberry just mentioned, Duchesne, 'Hist. Nat.
Frais.,' p. 133, says that this was a seedling raised from the fraisier des bois,
and the characters of which were reproduced by seed, and have now
become fixed. The monophyllous condition has been considered to be the
result of fusion of two or more leaflets, but however true this may be in
some cases, it is not the case with this strawberry. M. Paillot states that he
has found the variety in a wild state.[465]
In like manner varieties of the following plants occur with simple leaves,
Rosa berberifolia (Lowea), Rubus Idæus, Robinia pseudacacia, Fraxinus
excelsior, Sambucus nigra, Juglans nigra, &c.
as the individual leaves are altogether wanting, or with reference to the
diminished number of parts in a whorl, or a decrease in the verticils.
Aphylly.—Entire suppression of the leaves is a rare phenomenon. Under
ordinary circumstances it occurs in most Cactaceæ, in some of the
succulent Euphorbias, and other similar plants, where the epidermal layers
of the stem fulfil the functions of leaves. But even in these plants leaf-like
organs are present in some stage or another of the plant's life.
Partial suppression of the leaf occurs sometimes in compound leaves, some
or other of the leaflets of which are occasionally suppressed. Sometimes, as
Moquin remarks, it is the terminal leaflet which is wanting, when the
appearance is that of Cliffortia, at other times the lateral leaflets are
deficient, as in Citrus or Phyllarthron. Ononis monophylla and Fragaria
monophylla may be cited as instances of the suppression of the lateral
leaflets. If the blade of the leaf disappears entirely, we have then an
analogous condition to that of the phyllodineous acacias.
With reference to the strawberry just mentioned, Duchesne, 'Hist. Nat.
Frais.,' p. 133, says that this was a seedling raised from the fraisier des bois,
and the characters of which were reproduced by seed, and have now
become fixed. The monophyllous condition has been considered to be the
result of fusion of two or more leaflets, but however true this may be in
some cases, it is not the case with this strawberry. M. Paillot states that he
has found the variety in a wild state.[465]
In like manner varieties of the following plants occur with simple leaves,
Rosa berberifolia (Lowea), Rubus Idæus, Robinia pseudacacia, Fraxinus
excelsior, Sambucus nigra, Juglans nigra, &c.
Page 499
In one instance seen by the writer every portion of the leaf of a rose was
deficient, except the stipules and a small portion of the petiole. (See
abortion.)
Meiophylly.—A diminished number of leaves in a whorl occasionally takes
place; thus, in some of the Stellatæ, and frequently in Paris quadrifolia, the
number of leaves in the verticil is reduced. Care must be exercised in such
instances that an apparent diminution arising from a fusion of two or more
leaves be not confounded with suppression.
Meiophylly of the calyx or perianth.—A lessened number of sepals is not
a very common occurrence among dicotyledonous plants. Seringe figures a
proliferous flower of Arabis alpina with two sepals only, and a similar
occurrence has been noticed in Diplotaxis tenuifolia.
In Cattleya violacea the writer has met with a flower in which the
uppermost sepal was entirely wanting, while two of the lateral petals were
fused together. Moquin records that in some of the flowers of
Chenopodiaceæ, in which the inflorescence is dense, a suppression of two
or three sepals sometimes occurs. The species mentioned are Ambrina
ambrosioides, Chenopodium glaucum, and Blitum polymorphum.
Meiophylly of the corolla.—Suppression of one or more petals is of more
frequent occurrence than the corresponding deficiency in the case of the
sepals. Among Caryophyllaceæ imperfection as regards the numerical
symmetry of the flower is not uncommon, as in species of Cerastium,
Sagina, Dianthus, &c. In Ranunculaceæ the petals are likewise not
unfrequently partially or wholly suppressed. A familiar illustration of this is
afforded by Ranunculus auricomus, in which it is the exception to find the
corolla perfect.[466] Some varieties of Corchorus acutangulus in west
tropical Africa are likewise subject to the same peculiarity. Amongst
deficient, except the stipules and a small portion of the petiole. (See
abortion.)
Meiophylly.—A diminished number of leaves in a whorl occasionally takes
place; thus, in some of the Stellatæ, and frequently in Paris quadrifolia, the
number of leaves in the verticil is reduced. Care must be exercised in such
instances that an apparent diminution arising from a fusion of two or more
leaves be not confounded with suppression.
Meiophylly of the calyx or perianth.—A lessened number of sepals is not
a very common occurrence among dicotyledonous plants. Seringe figures a
proliferous flower of Arabis alpina with two sepals only, and a similar
occurrence has been noticed in Diplotaxis tenuifolia.
In Cattleya violacea the writer has met with a flower in which the
uppermost sepal was entirely wanting, while two of the lateral petals were
fused together. Moquin records that in some of the flowers of
Chenopodiaceæ, in which the inflorescence is dense, a suppression of two
or three sepals sometimes occurs. The species mentioned are Ambrina
ambrosioides, Chenopodium glaucum, and Blitum polymorphum.
Meiophylly of the corolla.—Suppression of one or more petals is of more
frequent occurrence than the corresponding deficiency in the case of the
sepals. Among Caryophyllaceæ imperfection as regards the numerical
symmetry of the flower is not uncommon, as in species of Cerastium,
Sagina, Dianthus, &c. In Ranunculaceæ the petals are likewise not
unfrequently partially or wholly suppressed. A familiar illustration of this is
afforded by Ranunculus auricomus, in which it is the exception to find the
corolla perfect.[466] Some varieties of Corchorus acutangulus in west
tropical Africa are likewise subject to the same peculiarity. Amongst
Page 500
Papilionaceæ absence of the carina or of the alæ is not uncommon, as in
Trifolium repens, Faba vulgaris, &c.
Moquin relates a case of the kind in the haricot bean, in which the carina
was entirely absent, and another in the pea, where both carina and alæ were
missing, thus reducing the flower to the condition that is normal in
Amorpha and Afzelia. Suppression of the upper lip in such flowers as
Calceolaria has been termed by Morren "apilary."
In Orchidaceæ entire absence of the labellum, frequently without any other
perceptible change, is of common occurrence. The writer has seen
numerous specimens of the kind in Ophrys apifera and O. aranifera; also in
Dendrobium nobile, Ærides odoratum, Cypripedium villosum, Listera
ovata, &c. Morren[467] mentions analogous deficiencies in Zygopetalum
maxillare, Calanthe sp., and Cattleya Forbesii. In most of these there was
also a fusion of the two lower sepals, which were so twisted out of place as
to occupy the situation usually held by the labellum. At the same time the
column was partially atrophied. To this deficiency of the lip the author just
quoted proposed to apply the term acheilary, α-χειλαριον. Mr. Moggridge
has communicated to the author an account of certain flowers of Ophrys
aranifera, in which the petals were deficient, sometimes completely, at
other times one or two only were present.
Meiophylly of the andrœcium.—Suppression of one or more stamens,
independently of like defects in other whorls, is not uncommon, even as a
normal occurrence, e.g. in Carlemannia, where the flower, though regular,
has only two stamens, and other similar deficiencies are common in
Dilleniads.
Seringe relates the occurrence of suppression of some of the stamens in
Diplotaxis tenuifolia,[468] St. Hilaire in Cardamine hirsuta, others in C.
Trifolium repens, Faba vulgaris, &c.
Moquin relates a case of the kind in the haricot bean, in which the carina
was entirely absent, and another in the pea, where both carina and alæ were
missing, thus reducing the flower to the condition that is normal in
Amorpha and Afzelia. Suppression of the upper lip in such flowers as
Calceolaria has been termed by Morren "apilary."
In Orchidaceæ entire absence of the labellum, frequently without any other
perceptible change, is of common occurrence. The writer has seen
numerous specimens of the kind in Ophrys apifera and O. aranifera; also in
Dendrobium nobile, Ærides odoratum, Cypripedium villosum, Listera
ovata, &c. Morren[467] mentions analogous deficiencies in Zygopetalum
maxillare, Calanthe sp., and Cattleya Forbesii. In most of these there was
also a fusion of the two lower sepals, which were so twisted out of place as
to occupy the situation usually held by the labellum. At the same time the
column was partially atrophied. To this deficiency of the lip the author just
quoted proposed to apply the term acheilary, α-χειλαριον. Mr. Moggridge
has communicated to the author an account of certain flowers of Ophrys
aranifera, in which the petals were deficient, sometimes completely, at
other times one or two only were present.
Meiophylly of the andrœcium.—Suppression of one or more stamens,
independently of like defects in other whorls, is not uncommon, even as a
normal occurrence, e.g. in Carlemannia, where the flower, though regular,
has only two stamens, and other similar deficiencies are common in
Dilleniads.
Seringe relates the occurrence of suppression of some of the stamens in
Diplotaxis tenuifolia,[468] St. Hilaire in Cardamine hirsuta, others in C.
Page 501
sylvatica.
In Caryophyllaceæ suppression of one or more stamens has been observed
in Mollugo cerviana, Arenaria tetraquetra, Cerastium, &c.[469] Among
violets the writer has observed numerous flowers in which two or three
stamens were suppressed. Chatin[470] alludes to a similar reduction in
Tropæolum, while in flowers that are usually didynamous absence of two or
more of the stamens is not unfrequent, e.g. in Antirrhinum, Digitalis, while
in a flower of Catalpa a solitary perfect stamen, and a complete absence of
the sterile ones usually present, have been observed. This might have been
anticipated from the frequent deficiencies in the staminal whorl in these
plants under what are considered to be normal conditions. Reduction of the
staminal whorl is also not unfrequent in Trifolium repens and T. hybridum,
and has been seen in Delphinium, &c.[471]
Meiophylly of the gynœcium.—Numerical inequality in the case of the
pistil, as compared with the other whorls of the flower, is of such common
occurrence, under ordinary circumstances, that in some text-books it is
looked on as the normal condition, and a flower which is isomerous in the
outer whorls is by some writers not considered numerically irregular if the
number of the carpels does not coincide with that of the other organs.
But in this place it is only necessary to allude to deviations from the number
of carpels that are ordinarily found in the particular species under
observation. As illustrations the following may be cited:—Arenaria
tetraqueta, which has normally three styles, and a six-valved capsule, has
been seen with two styles, and a four or five-valved capsule. Moquin relates
an instance in Polygala vulgaris where there was but a single carpel, a
condition analogous to that which occurs normally in the allied genus
Mozinna. Reseda luteola occasionally occurs with two carpels only, while
In Caryophyllaceæ suppression of one or more stamens has been observed
in Mollugo cerviana, Arenaria tetraquetra, Cerastium, &c.[469] Among
violets the writer has observed numerous flowers in which two or three
stamens were suppressed. Chatin[470] alludes to a similar reduction in
Tropæolum, while in flowers that are usually didynamous absence of two or
more of the stamens is not unfrequent, e.g. in Antirrhinum, Digitalis, while
in a flower of Catalpa a solitary perfect stamen, and a complete absence of
the sterile ones usually present, have been observed. This might have been
anticipated from the frequent deficiencies in the staminal whorl in these
plants under what are considered to be normal conditions. Reduction of the
staminal whorl is also not unfrequent in Trifolium repens and T. hybridum,
and has been seen in Delphinium, &c.[471]
Meiophylly of the gynœcium.—Numerical inequality in the case of the
pistil, as compared with the other whorls of the flower, is of such common
occurrence, under ordinary circumstances, that in some text-books it is
looked on as the normal condition, and a flower which is isomerous in the
outer whorls is by some writers not considered numerically irregular if the
number of the carpels does not coincide with that of the other organs.
But in this place it is only necessary to allude to deviations from the number
of carpels that are ordinarily found in the particular species under
observation. As illustrations the following may be cited:—Arenaria
tetraqueta, which has normally three styles, and a six-valved capsule, has
been seen with two styles, and a four or five-valved capsule. Moquin relates
an instance in Polygala vulgaris where there was but a single carpel, a
condition analogous to that which occurs normally in the allied genus
Mozinna. Reseda luteola occasionally occurs with two carpels only, while
Page 502
Aconites, Delphiniums, Nigellas, and Pæonies frequently experience a like
diminution in their pistil.
In a flower of Papaver Rhæas the writer has recently met with an ovary
with four stigmas and four parietal placentæ only, and to Mr. Worthington
Smith he is indebted for sketches of crocus blooms with two, and in one
instance only a solitary carpel.
Moquin cites the fruit of a wild bramble (Rubus) in which all the little
drupes which go to make up the ordinary fruit were absent, except one,
which thus resembled a small cherry. In Cratægus the pistil is similarly
reduced to a single carpel, as in C. monogyna.
The writer has on more than one occasion met with walnuts (Juglans) with
a single valve and a single suture.[472] If the ovary of Juglans normally
consisted of two valvate carpels, the instances just alluded to might possibly
be explained by the suppression of one carpel, but the ovary in Juglans is at
first one-celled according to M. Casimir de Candolle.
Among monocotyledons Convallaria majalis may be mentioned as very
liable to suffer diminution in the number of its carpels, either separately or
in association with other changes.[473]
Meiophylly of the flower as a whole.—In the preceding sections a
reduction in the parts of each individual whorl has been considered without
reference to similar diminution in neighbouring verticils. It more commonly
happens, nevertheless, that a defect in one series is attended by a
corresponding imperfection in adjoining ones. Thus trimerous fuchsias and
tetramerous jasmines may frequently be met with, and Turpin describes a
tetramerous flower of Cobæa scandens. Perhaps monocotyledonous plants
are more subject to this numerical reduction of the parts of several verticils
than are other flowering plants. Thus, in both Lilium lancifolium and L.
diminution in their pistil.
In a flower of Papaver Rhæas the writer has recently met with an ovary
with four stigmas and four parietal placentæ only, and to Mr. Worthington
Smith he is indebted for sketches of crocus blooms with two, and in one
instance only a solitary carpel.
Moquin cites the fruit of a wild bramble (Rubus) in which all the little
drupes which go to make up the ordinary fruit were absent, except one,
which thus resembled a small cherry. In Cratægus the pistil is similarly
reduced to a single carpel, as in C. monogyna.
The writer has on more than one occasion met with walnuts (Juglans) with
a single valve and a single suture.[472] If the ovary of Juglans normally
consisted of two valvate carpels, the instances just alluded to might possibly
be explained by the suppression of one carpel, but the ovary in Juglans is at
first one-celled according to M. Casimir de Candolle.
Among monocotyledons Convallaria majalis may be mentioned as very
liable to suffer diminution in the number of its carpels, either separately or
in association with other changes.[473]
Meiophylly of the flower as a whole.—In the preceding sections a
reduction in the parts of each individual whorl has been considered without
reference to similar diminution in neighbouring verticils. It more commonly
happens, nevertheless, that a defect in one series is attended by a
corresponding imperfection in adjoining ones. Thus trimerous fuchsias and
tetramerous jasmines may frequently be met with, and Turpin describes a
tetramerous flower of Cobæa scandens. Perhaps monocotyledonous plants
are more subject to this numerical reduction of the parts of several verticils
than are other flowering plants. Thus, in both Lilium lancifolium and L.
Page 503
auratum the writer has frequently met with pentamerous flowers. In
Convallaria maialis a like deviation not unfrequently occurs.[474] M.
Delavaud has recorded a similar occurrence in a tulip.[475]
Dimerous crocuses may also sometimes be met with. In one flower of this
nature the segments of the perianth were arranged in decussating pairs, and
the four stamens were united by their filaments so as to form two pairs.
M. Fournier mentions something of the same kind in the flower of an Iris.
[476]
Orchids seem peculiarly liable to the decrease in the number of their floral
organs. Prillieux[477] mentions a flower of Cattleya amethystina wherein
each whorl of the perianth consisted of two opposite segments.
The same observer has put on record instances of a similar kind in
Epidendrum Stamfordianum. In one flower of the last-named species the
perianth consisted of one sepal only, and one lip-like petal placed opposite
to it.[478] Morren[479] describes a flower of Cypripedium insigne, in which
there were two sepals and two petals. Of a similar character was the flower
found by Mr. J. A. Paine, and described in the following terms by Professor
Asa Gray in the 'American Journal of Science,' July, 1866:—"The plant"
(Cypripedium candidum) "bears two flowers: the axillary one is normal; the
terminal one exhibits the following peculiarities. The lower part of the bract
forms a sheath which encloses the ovary. The labellum is wanting; and there
are two sterile stamens, the supernumerary one being opposite the other, i.e.
on the side of the style where the labellum belongs. Accordingly the first
impression would be that the labellum is here transformed into a sterile
stamen. The latter, however, agrees with the normal sterile stamen in its
insertion as well as in shape, being equally adnate to the base of the style.
Moreover, the anteposed sepal is exactly like the other, has a good midrib
Convallaria maialis a like deviation not unfrequently occurs.[474] M.
Delavaud has recorded a similar occurrence in a tulip.[475]
Dimerous crocuses may also sometimes be met with. In one flower of this
nature the segments of the perianth were arranged in decussating pairs, and
the four stamens were united by their filaments so as to form two pairs.
M. Fournier mentions something of the same kind in the flower of an Iris.
[476]
Orchids seem peculiarly liable to the decrease in the number of their floral
organs. Prillieux[477] mentions a flower of Cattleya amethystina wherein
each whorl of the perianth consisted of two opposite segments.
The same observer has put on record instances of a similar kind in
Epidendrum Stamfordianum. In one flower of the last-named species the
perianth consisted of one sepal only, and one lip-like petal placed opposite
to it.[478] Morren[479] describes a flower of Cypripedium insigne, in which
there were two sepals and two petals. Of a similar character was the flower
found by Mr. J. A. Paine, and described in the following terms by Professor
Asa Gray in the 'American Journal of Science,' July, 1866:—"The plant"
(Cypripedium candidum) "bears two flowers: the axillary one is normal; the
terminal one exhibits the following peculiarities. The lower part of the bract
forms a sheath which encloses the ovary. The labellum is wanting; and there
are two sterile stamens, the supernumerary one being opposite the other, i.e.
on the side of the style where the labellum belongs. Accordingly the first
impression would be that the labellum is here transformed into a sterile
stamen. The latter, however, agrees with the normal sterile stamen in its
insertion as well as in shape, being equally adnate to the base of the style.
Moreover, the anteposed sepal is exactly like the other, has a good midrib
Page 504
and an entire point. As the two sterile stamens are anteposed to the two
sepals, so are the two fertile stamens to the two petals, and the latter are
adnate to the style a little higher than the former. The style is longer than
usual, is straight and erect; the broad, disciform stigma therefore faces
upwards; it is oval and symmetrical, and a light groove across its middle
shows it to be dimerous. The placentæ, accordingly, are only two. The
groove on the stigma and the placentæ are in line with the fertile stamens.
Here, therefore, is a symmetrical and complete, regular, but dimerous
orchideous flower, the first verticil of stamens not antheriferous, the second
antheriferous, the carpels alternate with these; and here we have clear (and
perhaps the first direct) demonstration that the orchideous type of flower
has two stamineal verticils, as Brown always insisted."
Fig. 198.—Regular dimerous flower of Calanthe vestita.
sepals, so are the two fertile stamens to the two petals, and the latter are
adnate to the style a little higher than the former. The style is longer than
usual, is straight and erect; the broad, disciform stigma therefore faces
upwards; it is oval and symmetrical, and a light groove across its middle
shows it to be dimerous. The placentæ, accordingly, are only two. The
groove on the stigma and the placentæ are in line with the fertile stamens.
Here, therefore, is a symmetrical and complete, regular, but dimerous
orchideous flower, the first verticil of stamens not antheriferous, the second
antheriferous, the carpels alternate with these; and here we have clear (and
perhaps the first direct) demonstration that the orchideous type of flower
has two stamineal verticils, as Brown always insisted."
Fig. 198.—Regular dimerous flower of Calanthe vestita.
Page 505
Fig. 199.—Regular dimerous flower of Odontoglossum Alexandræ.
Dr. Moore, of Glasnevin, kindly forwarded to the writer a flower of
Calanthe vestita (fig. 198), in which there were two sepals only, anterior
and posterior, and two petals at right angles to the two sepals. The lip was
entirely wanting, but the column and ovary were in their usual condition. In
Odontoglossum Alexandræ a similar reduction of parts has been observed
by the author (fig. 199).
It is curious to observe in these flowers how precisely one sepal occupies
the position of the labellum, and how the lateral petals are displaced from
the position they usually occupy, so as to form a regular flower, the
segments of which decussate, thus giving rise to a species of regular
peloria.
The genus Mælenia was established on a malformed flower of Orchis of
similar character to those above mentioned.
Dr. Moore, of Glasnevin, kindly forwarded to the writer a flower of
Calanthe vestita (fig. 198), in which there were two sepals only, anterior
and posterior, and two petals at right angles to the two sepals. The lip was
entirely wanting, but the column and ovary were in their usual condition. In
Odontoglossum Alexandræ a similar reduction of parts has been observed
by the author (fig. 199).
It is curious to observe in these flowers how precisely one sepal occupies
the position of the labellum, and how the lateral petals are displaced from
the position they usually occupy, so as to form a regular flower, the
segments of which decussate, thus giving rise to a species of regular
peloria.
The genus Mælenia was established on a malformed flower of Orchis of
similar character to those above mentioned.
Page 506
Meiotaxy of the calyx.—As already mentioned, this term is here employed
to denote those illustrations in which entire whorls are suppressed.
Complete deficiency of the calyx in a dichlamydeous flower seems seldom
or ever to occur; the nearest approach to it would be in those cases where
the calyx is, as it is termed, "obsolete," but here it is chiefly the limb of the
calyx which is atrophied, the lower portion being more or less adherent to
the ovary. In what are termed monochlamydeous flowers both calyx and
corolla are wanting, as in Salicineæ and many other orders.
Meiotaxy of the corolla.—Deficiency of the entire corolla occurs in
conjunction with similar reductions in other organs, or as an isolated
phenomenon in the many apetalous varieties of plants recorded in books.
Deficiency of the corolla was observed in Campanula perfoliata and
Ruellia clandestina by Linné, who calls such blooms flores mutilati.[480]
Drs. Hooker and Thomson relate a similar occurrence in Campanula
canescens and C. colorata. Some plants seem as a normal occurrence to
produce flowers of different construction, and are hence termed dimorphic,
as in many Malpighiaceæ, Violaceæ, Oxalidaceæ, in some of the flowers of
which the petals are altogether wanting, while in others the corolla is
developed as usual. This deficiency of the corolla is frequently, but not
invariably, associated with an increased fertility. Thus, in some violets the
flowers produced in summer, and in which the petals are either entirely
suppressed or are more or less atrophied, are always fertile, while the
blossoms developed in spring, and in which the petals are always present,
are much less fertile. In Oxalis Acetosella there are two forms of flower, the
one with, the other without, petals, but both seem equally fertile. Linné
remarks that many plants which, in warm latitudes, produce a corolla, do
not do so when grown in colder climates. Thus, certain species of
Helianthemum are apetalous in Lapland. In the Pyrenees, according to
Bentham, the flowers of Ajuga iva are constantly deprived of their corolla.
[481]
to denote those illustrations in which entire whorls are suppressed.
Complete deficiency of the calyx in a dichlamydeous flower seems seldom
or ever to occur; the nearest approach to it would be in those cases where
the calyx is, as it is termed, "obsolete," but here it is chiefly the limb of the
calyx which is atrophied, the lower portion being more or less adherent to
the ovary. In what are termed monochlamydeous flowers both calyx and
corolla are wanting, as in Salicineæ and many other orders.
Meiotaxy of the corolla.—Deficiency of the entire corolla occurs in
conjunction with similar reductions in other organs, or as an isolated
phenomenon in the many apetalous varieties of plants recorded in books.
Deficiency of the corolla was observed in Campanula perfoliata and
Ruellia clandestina by Linné, who calls such blooms flores mutilati.[480]
Drs. Hooker and Thomson relate a similar occurrence in Campanula
canescens and C. colorata. Some plants seem as a normal occurrence to
produce flowers of different construction, and are hence termed dimorphic,
as in many Malpighiaceæ, Violaceæ, Oxalidaceæ, in some of the flowers of
which the petals are altogether wanting, while in others the corolla is
developed as usual. This deficiency of the corolla is frequently, but not
invariably, associated with an increased fertility. Thus, in some violets the
flowers produced in summer, and in which the petals are either entirely
suppressed or are more or less atrophied, are always fertile, while the
blossoms developed in spring, and in which the petals are always present,
are much less fertile. In Oxalis Acetosella there are two forms of flower, the
one with, the other without, petals, but both seem equally fertile. Linné
remarks that many plants which, in warm latitudes, produce a corolla, do
not do so when grown in colder climates. Thus, certain species of
Helianthemum are apetalous in Lapland. In the Pyrenees, according to
Bentham, the flowers of Ajuga iva are constantly deprived of their corolla.
[481]
Page 507
Apetalous flowers have been noted most frequently in the following plants:
Aconitum, sp. pl.!
Cardamine impatiens.
Cheiranthus Cheiri!
Viola odorata!
Cerastium vulgatum!
Alsine media.
Stellaria.
Lychnis dioica!
Dianthus barbatus, and other Caryophylleæ.
Helianthemum, sp.!
Oxalis Acetosella.
Balsamineæ.
Malpighiaceæ.
Rosa centifolia.
arvensis!
Cratægus!
Medicago lupulina.
Melilotus officinalis.
Ononis minutissima.
Saxifraga longifolia.
Verbascum Thapsus.
Ajuga iva.
Teucrium Botrys.
Lamium purpureum!
amplexicaule.
Polemonium cæruleum.
Campanula, sp. pl.!
Ruellia clandestina.
Lonicera Periclymenum!
Tradescantia, sp.!
Hymenocallis.
The following references apply some to apetalous and others to dimorphic flowers, but it
must be remembered that the latter plants are not necessarily wanting in petals or stamens,
&c., though the functional activity of the parts may be impaired:
A. de Jussien, 'Monogr. Malpigh.,' pp. 82, 334. Torrey, 'Fl. New York,' i, p. 428. Hooker
and Thomson, 'Journ. Linn. Soc.,' ii, p. 7, Guillemin, 'Archiv. de Botan.,' i, p. 412.
Aconitum, sp. pl.!
Cardamine impatiens.
Cheiranthus Cheiri!
Viola odorata!
Cerastium vulgatum!
Alsine media.
Stellaria.
Lychnis dioica!
Dianthus barbatus, and other Caryophylleæ.
Helianthemum, sp.!
Oxalis Acetosella.
Balsamineæ.
Malpighiaceæ.
Rosa centifolia.
arvensis!
Cratægus!
Medicago lupulina.
Melilotus officinalis.
Ononis minutissima.
Saxifraga longifolia.
Verbascum Thapsus.
Ajuga iva.
Teucrium Botrys.
Lamium purpureum!
amplexicaule.
Polemonium cæruleum.
Campanula, sp. pl.!
Ruellia clandestina.
Lonicera Periclymenum!
Tradescantia, sp.!
Hymenocallis.
The following references apply some to apetalous and others to dimorphic flowers, but it
must be remembered that the latter plants are not necessarily wanting in petals or stamens,
&c., though the functional activity of the parts may be impaired:
A. de Jussien, 'Monogr. Malpigh.,' pp. 82, 334. Torrey, 'Fl. New York,' i, p. 428. Hooker
and Thomson, 'Journ. Linn. Soc.,' ii, p. 7, Guillemin, 'Archiv. de Botan.,' i, p. 412.
Page 508
Michalet, 'Bull. Soc. Bot. Fr.,' vii. p. 465. Müller, 'Bot. Zeit.,' 1857, p. 729. 'Natural History
Review,' July, 1862, p. 235.
Meiotaxy of the andrœcium.—Complete suppression of the stamens
occurs normally in the female flowers of unisexual plants, and, as an
accidental occurrence, is not very uncommon. Erica Tetralix is one of the
plants in which this is said to happen. The variety anandra is said to have
been known in France since 1635. Cornuti speaks of it in his 'Enchiridion.'
In 1860 M. du Parquet discovered it in peaty woods near Nangis (Seine et
Marne).
Many Umbelliferæ, such as Trinia vulgaris, present a like deficiency, while
it is of common occurrence among Rosaceæ and Pomaceæ. In the latter
group the St. Valery apple, so often referred to, is an illustration. To obtain
fruits from this variety it is necessary to apply pollen from another flower, a
proceeding made the occasion of festivity and rejoicing by the villagers in
some parts of France. In some of the Artemisias, especially in Artemisia
Tournefortiana, all the florets have been noticed to be female, owing to the
suppression of the stamens, and this suppression is associated with a change
in the form of florets.[482] Mr. Moggridge has communicated to the author
flowers of Thymus Serpyllum from a plant in which all the stamens were
deficient, the flower being otherwise normal.
M. Dupont has given a list of nineteen species of Chenopodiaceæ in which
female flowers are occasionally produced, owing to the entire suppression
of the staminal whorl.[483]
Flowers the subjects either of regular or irregular peloria, q. v., are often
destitute of some or all their stamens, e.g. Calceolaria, Linaria, Viola, &c.,
while in cases of synanthy suppression of some of the parts of the flower,
and specially of the stamens, is of very common occurrence.
Review,' July, 1862, p. 235.
Meiotaxy of the andrœcium.—Complete suppression of the stamens
occurs normally in the female flowers of unisexual plants, and, as an
accidental occurrence, is not very uncommon. Erica Tetralix is one of the
plants in which this is said to happen. The variety anandra is said to have
been known in France since 1635. Cornuti speaks of it in his 'Enchiridion.'
In 1860 M. du Parquet discovered it in peaty woods near Nangis (Seine et
Marne).
Many Umbelliferæ, such as Trinia vulgaris, present a like deficiency, while
it is of common occurrence among Rosaceæ and Pomaceæ. In the latter
group the St. Valery apple, so often referred to, is an illustration. To obtain
fruits from this variety it is necessary to apply pollen from another flower, a
proceeding made the occasion of festivity and rejoicing by the villagers in
some parts of France. In some of the Artemisias, especially in Artemisia
Tournefortiana, all the florets have been noticed to be female, owing to the
suppression of the stamens, and this suppression is associated with a change
in the form of florets.[482] Mr. Moggridge has communicated to the author
flowers of Thymus Serpyllum from a plant in which all the stamens were
deficient, the flower being otherwise normal.
M. Dupont has given a list of nineteen species of Chenopodiaceæ in which
female flowers are occasionally produced, owing to the entire suppression
of the staminal whorl.[483]
Flowers the subjects either of regular or irregular peloria, q. v., are often
destitute of some or all their stamens, e.g. Calceolaria, Linaria, Viola, &c.,
while in cases of synanthy suppression of some of the parts of the flower,
and specially of the stamens, is of very common occurrence.
Page 509
Suppression of the andrœcium as a teratological occurrence has been most
frequently noticed in the following plants, omitting members of those
families whose floral construction is normally incomplete in the majority of
instances, and exclusive also of cases of substitution. See also under
Heterogamy.
Ranunculus Ficaria!
auricomus!
bulbosus!
Cruciferæ, sp. pl.
Violaceæ, sp. pl.
Honckenya peploides.
Stellaria.
Caryophyllaceæ, sp. pl.
Malpighiaceæ, sp. pl.
Tropæolum majus!
Fragaria vesca!
Rubus, sp.
Pyrus Malus.
Agrimonia vulgaris.
Rosaceæ, sp. pl.
Trifolium hybridum.
repens.
Umbelliferæ, sp. pl.
Onagraceæ, sp. pl.
Hippuris vulgaris.
Callitriche vernalis.
autumnalis.
Lonicera Periclymenum.
Erica Tetralix.
Thymus Serpyllum.
Calceolaria.
Compositæ, sp. pl.
Chenopodiaceæ, sp. pl.
Stratiotes aloides.
Meiotaxy of the gynœcium.—Complete suppression of the pistil is of more
frequent occurrence than that of the stamens, hence more flowers become
frequently noticed in the following plants, omitting members of those
families whose floral construction is normally incomplete in the majority of
instances, and exclusive also of cases of substitution. See also under
Heterogamy.
Ranunculus Ficaria!
auricomus!
bulbosus!
Cruciferæ, sp. pl.
Violaceæ, sp. pl.
Honckenya peploides.
Stellaria.
Caryophyllaceæ, sp. pl.
Malpighiaceæ, sp. pl.
Tropæolum majus!
Fragaria vesca!
Rubus, sp.
Pyrus Malus.
Agrimonia vulgaris.
Rosaceæ, sp. pl.
Trifolium hybridum.
repens.
Umbelliferæ, sp. pl.
Onagraceæ, sp. pl.
Hippuris vulgaris.
Callitriche vernalis.
autumnalis.
Lonicera Periclymenum.
Erica Tetralix.
Thymus Serpyllum.
Calceolaria.
Compositæ, sp. pl.
Chenopodiaceæ, sp. pl.
Stratiotes aloides.
Meiotaxy of the gynœcium.—Complete suppression of the pistil is of more
frequent occurrence than that of the stamens, hence more flowers become
Page 510
accidentally unisexual by suppression of the pistil than by deficiency of the
stamens.
In many Umbelliferæ, e.g. Torilis Anthriscus, Cicuta virosa, the central
flowers are often male, owing to the suppression of the pistil. In many
double flowers, owing to the excessive multiplication of petaloid stamens,
the pistil is suppressed, in which cases it often happens that the flower is
depressed in the centre, as in some garden varieties of Ranunculus.
Schlechtendal, in describing a flower of Colchicum autumnale, in which the
perianth was virescent, says that, although the stamens were present, the
pistil was absent.
In proliferous flowers the pistil is often completely defective, its place
being occupied by the adventitious bud or axis.
As in other cases of like nature, suppression of the pistil is very frequently
consequent on fusion of flowers or other changes. Thus Morren relates an
instance of synanthy in the flowers of Torenia scabra, accompanied by
resorption or disappearance of some parts and spiral torsion of others. The
pistil was entirely absent in this instance.[484]
M. Gaetano Licopoli places on record an instance where the petals and
carpels of Melianthus major were suppressed.[485]
On the whole, the pistil seems less subject to changes of this character than
the andrœcium.
Suppression of the pistil has been most frequently recorded in flowers
(normally bisexual) of—
Ranunculus!
Aconitium!
Delphinium!
stamens.
In many Umbelliferæ, e.g. Torilis Anthriscus, Cicuta virosa, the central
flowers are often male, owing to the suppression of the pistil. In many
double flowers, owing to the excessive multiplication of petaloid stamens,
the pistil is suppressed, in which cases it often happens that the flower is
depressed in the centre, as in some garden varieties of Ranunculus.
Schlechtendal, in describing a flower of Colchicum autumnale, in which the
perianth was virescent, says that, although the stamens were present, the
pistil was absent.
In proliferous flowers the pistil is often completely defective, its place
being occupied by the adventitious bud or axis.
As in other cases of like nature, suppression of the pistil is very frequently
consequent on fusion of flowers or other changes. Thus Morren relates an
instance of synanthy in the flowers of Torenia scabra, accompanied by
resorption or disappearance of some parts and spiral torsion of others. The
pistil was entirely absent in this instance.[484]
M. Gaetano Licopoli places on record an instance where the petals and
carpels of Melianthus major were suppressed.[485]
On the whole, the pistil seems less subject to changes of this character than
the andrœcium.
Suppression of the pistil has been most frequently recorded in flowers
(normally bisexual) of—
Ranunculus!
Aconitium!
Delphinium!
Page 511
Pæonia.
Caryophylleæ!
Umbelliferæ.
Trifolium repens.
hybridum.
Compositæ, sp. pl.
Datura.
Torenia asiatica.
Colchicum autumnale.
Suppression of ovules,—abortion of seeds.—The two cases are taken
together, as the effects are similar, though it must be remembered that in the
one case the ovules at any rate have been formed, but their development has
been arrested, while in the other they have never existed. The precise cause
that has determined the absence of seed cannot in all cases be ascertained in
the adult condition, hence it is convenient to treat the two phenomena under
one head.
Many plants in other than their native climates either produce no fruit at all,
or the fruits that are produced are destitute of seed, e.g. Musa, Artocarpus,
&c. Some of the cultivated varieties of the grape and of the berberry
produce no seeds.
Suppression or abortion of the seed is frequently associated with the
excessive development either in size or number of other portions of the
plant, or with an altered condition, as when carpels become foliaceous and
their margins detached. Hybridisation and cross fertilisation are also well-
known agents in diminishing the number and size of seeds.
Meiotaxy of the parts of the flower in general.—In the preceding sections
suppression has been considered as it affected individual members of a
whorl or separate whorls. It rarely happens, however, that the suppression is
limited in this way. More generally several of the parts of the flower are
simultaneously affected in the same manner.
Caryophylleæ!
Umbelliferæ.
Trifolium repens.
hybridum.
Compositæ, sp. pl.
Datura.
Torenia asiatica.
Colchicum autumnale.
Suppression of ovules,—abortion of seeds.—The two cases are taken
together, as the effects are similar, though it must be remembered that in the
one case the ovules at any rate have been formed, but their development has
been arrested, while in the other they have never existed. The precise cause
that has determined the absence of seed cannot in all cases be ascertained in
the adult condition, hence it is convenient to treat the two phenomena under
one head.
Many plants in other than their native climates either produce no fruit at all,
or the fruits that are produced are destitute of seed, e.g. Musa, Artocarpus,
&c. Some of the cultivated varieties of the grape and of the berberry
produce no seeds.
Suppression or abortion of the seed is frequently associated with the
excessive development either in size or number of other portions of the
plant, or with an altered condition, as when carpels become foliaceous and
their margins detached. Hybridisation and cross fertilisation are also well-
known agents in diminishing the number and size of seeds.
Meiotaxy of the parts of the flower in general.—In the preceding sections
suppression has been considered as it affected individual members of a
whorl or separate whorls. It rarely happens, however, that the suppression is
limited in this way. More generally several of the parts of the flower are
simultaneously affected in the same manner.
Page 512
A few illustrations are all that is necessary to give as to this point.
One of the most familiar instances is that of the cauliflower or broccoli,
where the common flower-stalk is inordinately thickened and fleshy, while
the corolla and inner parts of the flower are usually entirely suppressed; the
four sepals can, however, generally be detected.
Maximowicz describes a Stellaria (Kraschenikovia) in which the upper
flowers are male only, while the lower ones, which ultimately become
buried in the soil, have neither petals, stamens, nor styles, but the walls of
the capsule are fleshy, and enclose numerous seeds.[486]
Kirschleger[487] mentions a variety of Lonicera Caprifolium, which was not
only destitute of petals but of stamens also.
In some species of Muscari and Bellevalia the uppermost flowers of the
raceme show more or less complete suppression of almost all the part of
which the flower normally consists. In those cases where an imperfect
perianth exists, but in which the stamens and pistils are entirely suppressed,
Morren applies the term Cenanthy, κενος, empty.
Complete suppression of the flower.—It is not necessary in this place to
allude to that deficient production of flowers characteristic of what is
termed by gardeners a "sky bloomer." In such plants often the requisite
conditions are not complied with, and the skill of the gardener is shown in
his attempt to discover and allow the plant to avail itself of the necessary
requirements. We need here only allude to those instances in which
provision is made for the production of flowers, and yet they are not
produced. A good illustration of this is afforded by the feather-hyacinth,
Hyacinthus comosus, in which the flowers are almost entirely suppressed,
while the pedicels are inordinately increased in number, and their colour
heightened. Something similar occurs in several allied species, and in
One of the most familiar instances is that of the cauliflower or broccoli,
where the common flower-stalk is inordinately thickened and fleshy, while
the corolla and inner parts of the flower are usually entirely suppressed; the
four sepals can, however, generally be detected.
Maximowicz describes a Stellaria (Kraschenikovia) in which the upper
flowers are male only, while the lower ones, which ultimately become
buried in the soil, have neither petals, stamens, nor styles, but the walls of
the capsule are fleshy, and enclose numerous seeds.[486]
Kirschleger[487] mentions a variety of Lonicera Caprifolium, which was not
only destitute of petals but of stamens also.
In some species of Muscari and Bellevalia the uppermost flowers of the
raceme show more or less complete suppression of almost all the part of
which the flower normally consists. In those cases where an imperfect
perianth exists, but in which the stamens and pistils are entirely suppressed,
Morren applies the term Cenanthy, κενος, empty.
Complete suppression of the flower.—It is not necessary in this place to
allude to that deficient production of flowers characteristic of what is
termed by gardeners a "sky bloomer." In such plants often the requisite
conditions are not complied with, and the skill of the gardener is shown in
his attempt to discover and allow the plant to avail itself of the necessary
requirements. We need here only allude to those instances in which
provision is made for the production of flowers, and yet they are not
produced. A good illustration of this is afforded by the feather-hyacinth,
Hyacinthus comosus, in which the flowers are almost entirely suppressed,
while the pedicels are inordinately increased in number, and their colour
heightened. Something similar occurs in several allied species, and in
Page 513
Bowiea volubilis. The wig plant (Rhus Cotinus) affords another illustration
of the same thing. Some tendrils also owe their appearance to the absence
of flowers, being modified peduncles; proofs of this may frequently be met
with in the case of the vine.
In Lamium album I have seen one of the verticillasters on one side of the
stem completely wanting, the adjacent leaf being, however, as fully formed
as usual.
General remarks on suppression.—On comparing together the various
whorls of the flower in reference to suppression, and, it may be added, to
atrophy, we find that these phenomena occur most rarely in the calyx, more
frequently in the corolla, and very often in the sexual organs and seeds;
hence it would seem as if the uppermost and most central organs, those
most subject to pressure and latest in date of development—formed, that is,
when the formative energies of the plant are most liable to be exhausted—
are the most prone to be suppressed or arrested in their development. When
the plants in which these occurrences happen most frequently are compared
together, it may be seen that partial or entire suppression of the floral
envelopes, calyx, and corolla, is far more commonly met with in the
polypetalous and hypogynous groups than in the gamopetalous or
epigynous series.
The orders in which suppression (speaking generally) occurs most often as
a teratological occurrence are the following:—Ranunculaceæ, Cruciferæ,
Caryophyllaceæ, Violaceæ, Leguminosæ, Onagraceæ, Jasminaceæ,
Orchidaceæ. It will be observed that these are all orders wherein
suppression of the whole or part of the outer floral whorls takes place in
certain genera as a constant occurrence.
Again, it may be remarked that many of these orders show a tendency
towards a regular diminution of the assumed normal number of their parts;
of the same thing. Some tendrils also owe their appearance to the absence
of flowers, being modified peduncles; proofs of this may frequently be met
with in the case of the vine.
In Lamium album I have seen one of the verticillasters on one side of the
stem completely wanting, the adjacent leaf being, however, as fully formed
as usual.
General remarks on suppression.—On comparing together the various
whorls of the flower in reference to suppression, and, it may be added, to
atrophy, we find that these phenomena occur most rarely in the calyx, more
frequently in the corolla, and very often in the sexual organs and seeds;
hence it would seem as if the uppermost and most central organs, those
most subject to pressure and latest in date of development—formed, that is,
when the formative energies of the plant are most liable to be exhausted—
are the most prone to be suppressed or arrested in their development. When
the plants in which these occurrences happen most frequently are compared
together, it may be seen that partial or entire suppression of the floral
envelopes, calyx, and corolla, is far more commonly met with in the
polypetalous and hypogynous groups than in the gamopetalous or
epigynous series.
The orders in which suppression (speaking generally) occurs most often as
a teratological occurrence are the following:—Ranunculaceæ, Cruciferæ,
Caryophyllaceæ, Violaceæ, Leguminosæ, Onagraceæ, Jasminaceæ,
Orchidaceæ. It will be observed that these are all orders wherein
suppression of the whole or part of the outer floral whorls takes place in
certain genera as a constant occurrence.
Again, it may be remarked that many of these orders show a tendency
towards a regular diminution of the assumed normal number of their parts;
Page 514
thus, among Onagraceæ, Circeia and Lopezia may be referred to, the
former normally dimerous, the latter having only one perfect petal. So in
fuchsias, a very common deviation consists in a trimerous and rarely a
dimerous symmetry of the flower.
Although, if the absolute number of genera or orders be counted, there
appears to be little difference in the frequency of the occurrence of
suppression in irregular flowers as contrasted with regular flowers, yet if
the individual instances could be counted in the two groups respectively it
would be found that suppression is more common among irregular than in
regular flowers. Thus, the number of individual instances of flowers in
which the perianth is defective is comparatively large among Violaceæ,
Leguminosæ, and Orchidaceæ. This statement hardly admits of precise
statistical proof; still, it is believed that any observer who pays attention to
the subject must come to the same conclusion. This is but another
illustration of the fact that conditions which are abnormal in one plant
constitute the natural arrangement in others.
As to the suppressions that occur in the case of the sexual organs, and the
relations they bear to dimorphism, diclinism, &c., but little stress has been
laid on them in this place, because their chief interest is in a physiological
point of view, and is treated of in the writings of Mohl, Sprengel, Darwin,
Hildebrand, and others. All that need be said here is, that teratology affords
very numerous illustrations of those intermediate conditions which are also
found, under natural circumstances, between the absolutely unisexual
flowers, male or female, and the structurally hermaphrodite ones.
Rudimentary stamens or pistils are of very common occurrence in
monstrous flowers. See Chapter on Heterogamy, &c.
FOOTNOTES:
[465] 'Rev. Hortic.,' 1866, p. 467.
former normally dimerous, the latter having only one perfect petal. So in
fuchsias, a very common deviation consists in a trimerous and rarely a
dimerous symmetry of the flower.
Although, if the absolute number of genera or orders be counted, there
appears to be little difference in the frequency of the occurrence of
suppression in irregular flowers as contrasted with regular flowers, yet if
the individual instances could be counted in the two groups respectively it
would be found that suppression is more common among irregular than in
regular flowers. Thus, the number of individual instances of flowers in
which the perianth is defective is comparatively large among Violaceæ,
Leguminosæ, and Orchidaceæ. This statement hardly admits of precise
statistical proof; still, it is believed that any observer who pays attention to
the subject must come to the same conclusion. This is but another
illustration of the fact that conditions which are abnormal in one plant
constitute the natural arrangement in others.
As to the suppressions that occur in the case of the sexual organs, and the
relations they bear to dimorphism, diclinism, &c., but little stress has been
laid on them in this place, because their chief interest is in a physiological
point of view, and is treated of in the writings of Mohl, Sprengel, Darwin,
Hildebrand, and others. All that need be said here is, that teratology affords
very numerous illustrations of those intermediate conditions which are also
found, under natural circumstances, between the absolutely unisexual
flowers, male or female, and the structurally hermaphrodite ones.
Rudimentary stamens or pistils are of very common occurrence in
monstrous flowers. See Chapter on Heterogamy, &c.
FOOTNOTES:
[465] 'Rev. Hortic.,' 1866, p. 467.
Page 515
[466] De Rochebrune, 'Bull. Soc. Bot. Fr.,' ix, p. 281. The author points out seven grades
between complete absence of petals and their presence in the normal number in this plant. See
also Gaudin, in 'Koch. Fl. Helv.;' Koch. 'Synops. Fl. Germ.;' Cramer, 'Bildungsabweich,' p. 85.
[467] 'Bull. Acad. Belg.,' t. xix, part 1, p. 255.
[468] 'Bull. Bot.,' i, p. 7, tab. i, f. 7.
[469] See Gay, 'Ann. Sc. Nat.,' iii, p. 27.
[470] 'Ann. Sc. Nat.,' 4 ser., v, p. 305.
[471] Cramer, 'Bildungsabweich,' p. 90.
[472] See also Clos, 'Bull. Soc. Bot. Fr.,' xiii, p. 96, adnot.
[473] See Cramer, 'Bildungsabweich,' p. 7. Hildebrand, 'Bot. Zeit.,' xx, 1862, p. 209.
[474] See Hildebrand, 'Bot. Zeit.,' xx, 1862, p. 209.
[475] 'Bull. Soc. Bot. Fr.,' viii, p. 287.
[476] 'Bull. Soc. Bot. Fr.,' vol. viii, 1861, p. 152.
[477] Ibid., ix, p. 275.
[478] Ibid., 1861, vol. viii, p. 149.
[479] 'Lobelia,' p. 55.
[480] 'Phil. Bot.,' p. 119.
[481] 'Cat. Plant. Pyr,' p. 58.
[482] Moquin-Tandon, loc. cit., p. 328.
[483] For other instances see Chatin in 'Ann. Sc. Nat.,' 4 ser., vol. v, p. 305.
[484] See also Morren. 'Bull. Acad. Belg.,' xv, Fuchsia, p. 67.
[485] Cited in 'Bull. Soc. Bot., France,' t. xiv ("Rev. Bibl."), p. 253.
[486] 'Primit. Flor. Amurens.' p. 57.
[487] 'Flora.' 1848. p. 484.
between complete absence of petals and their presence in the normal number in this plant. See
also Gaudin, in 'Koch. Fl. Helv.;' Koch. 'Synops. Fl. Germ.;' Cramer, 'Bildungsabweich,' p. 85.
[467] 'Bull. Acad. Belg.,' t. xix, part 1, p. 255.
[468] 'Bull. Bot.,' i, p. 7, tab. i, f. 7.
[469] See Gay, 'Ann. Sc. Nat.,' iii, p. 27.
[470] 'Ann. Sc. Nat.,' 4 ser., v, p. 305.
[471] Cramer, 'Bildungsabweich,' p. 90.
[472] See also Clos, 'Bull. Soc. Bot. Fr.,' xiii, p. 96, adnot.
[473] See Cramer, 'Bildungsabweich,' p. 7. Hildebrand, 'Bot. Zeit.,' xx, 1862, p. 209.
[474] See Hildebrand, 'Bot. Zeit.,' xx, 1862, p. 209.
[475] 'Bull. Soc. Bot. Fr.,' viii, p. 287.
[476] 'Bull. Soc. Bot. Fr.,' vol. viii, 1861, p. 152.
[477] Ibid., ix, p. 275.
[478] Ibid., 1861, vol. viii, p. 149.
[479] 'Lobelia,' p. 55.
[480] 'Phil. Bot.,' p. 119.
[481] 'Cat. Plant. Pyr,' p. 58.
[482] Moquin-Tandon, loc. cit., p. 328.
[483] For other instances see Chatin in 'Ann. Sc. Nat.,' 4 ser., vol. v, p. 305.
[484] See also Morren. 'Bull. Acad. Belg.,' xv, Fuchsia, p. 67.
[485] Cited in 'Bull. Soc. Bot., France,' t. xiv ("Rev. Bibl."), p. 253.
[486] 'Primit. Flor. Amurens.' p. 57.
[487] 'Flora.' 1848. p. 484.
Page 516
Page 517
BOOK IV.
DEVIATIONS FROM THE ORDINARY SIZE
AND CONSISTENCE OF ORGANS.
In the animal kingdom the entire adult organism, as well as each of its
separate parts, has certain dimensions, beyond which, under ordinary
circumstances, it does not pass, either in the one direction or the other. It
may not be easy or possible to state what the limits are, but, practically, this
inability to frame a precise limitation is productive of no inconvenience. It
is universally admitted that a certain animal attains such and such
dimensions, and that one organ has a certain proportionate size as
contrasted with another. The same rules hold good in the case of plants,
though in them it is vastly more difficult to ascertain what may be called the
normal dimensions or proportions. Nevertheless observation and experience
soon show what may be termed the average size of each plant, and any
disproportion between the several organs is speedily detected.
When there is a general reduction in size throughout all the organs of a
plant, or throughout all the nutritive organs, stem, leaves, &c., and the
several portions participate in this diminished size, we have what are
generally termed "dwarf varieties," dwarf in comparison, that is, with the
ordinary condition of the plants; on the other hand, if the entire plant, or, at
least, if the whole of one set of organs be increased in size beyond the
DEVIATIONS FROM THE ORDINARY SIZE
AND CONSISTENCE OF ORGANS.
In the animal kingdom the entire adult organism, as well as each of its
separate parts, has certain dimensions, beyond which, under ordinary
circumstances, it does not pass, either in the one direction or the other. It
may not be easy or possible to state what the limits are, but, practically, this
inability to frame a precise limitation is productive of no inconvenience. It
is universally admitted that a certain animal attains such and such
dimensions, and that one organ has a certain proportionate size as
contrasted with another. The same rules hold good in the case of plants,
though in them it is vastly more difficult to ascertain what may be called the
normal dimensions or proportions. Nevertheless observation and experience
soon show what may be termed the average size of each plant, and any
disproportion between the several organs is speedily detected.
When there is a general reduction in size throughout all the organs of a
plant, or throughout all the nutritive organs, stem, leaves, &c., and the
several portions participate in this diminished size, we have what are
generally termed "dwarf varieties," dwarf in comparison, that is, with the
ordinary condition of the plants; on the other hand, if the entire plant, or, at
least, if the whole of one set of organs be increased in size beyond the
Page 518
recognised average, we have large varieties, often qualified by such terms
as macrophylla, longifolia, macrantha, &c. &c. In all these cases either the
entire plant or whole series of organs are alike increased or diminished
beyond average limits; and such variations are often very constant, and are
transmitted by hereditary transmission. It may be supposed that such
deviations may have originated, in the first instance, either from excessive
use, or from disuse, or from the agency of certain conditions promoting or
checking growth, as the case may be; but whether or no, it is certain that
these variations often persist under different conditions, and that they often
retain their distinctive characters side by side with plants presenting the
normal average dimensions. In other cases the variations in size are of a less
general character, and affect certain organs of a whorl in a relative manner,
as, for instance, in the case of didynamous or tetradynamous stamens,
where two or four stamens are longer than their fellows, the long or short
stamens and styles of di- and tri-morphic flowers, &c. These differences are
sometimes connected with the development of parts in succession, and not
simultaneously.
Teratological deviations of size differ from those of which mention has just
been made chiefly in this, that they are more limited in their manifestations.
It is not, as a rule, the whole plant, or the whole series of nutritive or of
reproductive organs, that are affected, but it is certain parts only; the
alteration in size is more a relative change than an absolute one.
For convenience sake the teratological alterations of size may be divided
into those which are the result of increased growth and those which arise
from diminished action. It will be seen, therefore, that in these instances it
is the bulk of the organs that is increased, not their number; moreover, their
development or metamorphosis is not necessarily altered. In connection
with increased size an alteration of consistence is so frequent that the two
phenomena are here taken together. It will be borne in mind that the
as macrophylla, longifolia, macrantha, &c. &c. In all these cases either the
entire plant or whole series of organs are alike increased or diminished
beyond average limits; and such variations are often very constant, and are
transmitted by hereditary transmission. It may be supposed that such
deviations may have originated, in the first instance, either from excessive
use, or from disuse, or from the agency of certain conditions promoting or
checking growth, as the case may be; but whether or no, it is certain that
these variations often persist under different conditions, and that they often
retain their distinctive characters side by side with plants presenting the
normal average dimensions. In other cases the variations in size are of a less
general character, and affect certain organs of a whorl in a relative manner,
as, for instance, in the case of didynamous or tetradynamous stamens,
where two or four stamens are longer than their fellows, the long or short
stamens and styles of di- and tri-morphic flowers, &c. These differences are
sometimes connected with the development of parts in succession, and not
simultaneously.
Teratological deviations of size differ from those of which mention has just
been made chiefly in this, that they are more limited in their manifestations.
It is not, as a rule, the whole plant, or the whole series of nutritive or of
reproductive organs, that are affected, but it is certain parts only; the
alteration in size is more a relative change than an absolute one.
For convenience sake the teratological alterations of size may be divided
into those which are the result of increased growth and those which arise
from diminished action. It will be seen, therefore, that in these instances it
is the bulk of the organs that is increased, not their number; moreover, their
development or metamorphosis is not necessarily altered. In connection
with increased size an alteration of consistence is so frequent that the two
phenomena are here taken together. It will be borne in mind that the
Page 519
changes of consistence from membranous to succulent or woody are very
frequent in the ordinary course of development. They may also occur as
accidental phenomena, or the normal conditions of any particular flower or
fruit may be exactly reversed, the usually succulent fruit becoming dry and
capsular, and so forth.
PART I.
HYPERTROPHY.
The term hypertrophy may serve as a general one to comprise all the
instances of excessive growth and increased size of organs, whether the
increase be general or in one direction merely. General hypertrophy is more
a variation than a deformity, unless indeed it be caused by insect puncture
or the presence of a fungus, in which case the excessive size results from a
diseased condition. For our present purpose hypertrophy may be considered
as it affects the axile or the foliar organs, and also according to the way in
which the increased size is manifested, as by increased thickness or
swelling—intumescence, or by augmented length-elongation, by expansion
or flattening, or, lastly, by the formation of excrescences or outgrowths,
which may be classed under the head of luxuriance or enation.
As size must be considered in this place relatively, it is not possible to lay
down any precise line separating what are considered to be the normal
dimensions from those which are abnormal.
In practice no inconvenience will be found to accrue from this inability to
establish a fixed rule, and we may say that an hypertrophied organ is one
which, from some cause or other, attains dimensions which are not habitual
to the plant in its usual, healthy, well-formed state.
frequent in the ordinary course of development. They may also occur as
accidental phenomena, or the normal conditions of any particular flower or
fruit may be exactly reversed, the usually succulent fruit becoming dry and
capsular, and so forth.
PART I.
HYPERTROPHY.
The term hypertrophy may serve as a general one to comprise all the
instances of excessive growth and increased size of organs, whether the
increase be general or in one direction merely. General hypertrophy is more
a variation than a deformity, unless indeed it be caused by insect puncture
or the presence of a fungus, in which case the excessive size results from a
diseased condition. For our present purpose hypertrophy may be considered
as it affects the axile or the foliar organs, and also according to the way in
which the increased size is manifested, as by increased thickness or
swelling—intumescence, or by augmented length-elongation, by expansion
or flattening, or, lastly, by the formation of excrescences or outgrowths,
which may be classed under the head of luxuriance or enation.
As size must be considered in this place relatively, it is not possible to lay
down any precise line separating what are considered to be the normal
dimensions from those which are abnormal.
In practice no inconvenience will be found to accrue from this inability to
establish a fixed rule, and we may say that an hypertrophied organ is one
which, from some cause or other, attains dimensions which are not habitual
to the plant in its usual, healthy, well-formed state.
Page 520
It will be seen that under this general head of hypertrophy, increase of size,
however brought about, is included; thus, not only increase in length, but
also in thickness; alterations of substance or consistence, no less than of
dimensions, are here grouped together. The alterations of consistence
resulting from an inordinate development of cellular, fibrous, or ligneous
tissue, are, of course, strictly homologous with the similar changes which
occur, under ordinary circumstances, during the ripening of fruits or
otherwise.
Hypertrophy, whatever form it may assume, may be so slight as not
perceptibly to interfere with the functions of the part affected, or it may
exist to such an extent as to impair the due exercise of its office. It may
affect any or all parts of the plant, and is generally coexistent with, if not
actually dependent on, some other malformation. Thus, the inordinate
growth of some parts is most generally attended by deficiency in the size
and number of others, as in the peripheral florets of Viburnum or
Hydrangea, where the corollas are relatively very large, and the stamens
and pistils abortive.
CHAPTER I.
ENLARGEMENT.
A swollen or thickened condition (renflement) is usually the result of a
disproportionate formation of the cellular tissue as contrasted with the
woody framework of the plant. We see marked instances of it in cultivated
carrots and turnips, the normal condition of the roots or root-stocks in these
plants being one of considerable hardness and toughness, and their form
slender, tapering, and more or less branched.
however brought about, is included; thus, not only increase in length, but
also in thickness; alterations of substance or consistence, no less than of
dimensions, are here grouped together. The alterations of consistence
resulting from an inordinate development of cellular, fibrous, or ligneous
tissue, are, of course, strictly homologous with the similar changes which
occur, under ordinary circumstances, during the ripening of fruits or
otherwise.
Hypertrophy, whatever form it may assume, may be so slight as not
perceptibly to interfere with the functions of the part affected, or it may
exist to such an extent as to impair the due exercise of its office. It may
affect any or all parts of the plant, and is generally coexistent with, if not
actually dependent on, some other malformation. Thus, the inordinate
growth of some parts is most generally attended by deficiency in the size
and number of others, as in the peripheral florets of Viburnum or
Hydrangea, where the corollas are relatively very large, and the stamens
and pistils abortive.
CHAPTER I.
ENLARGEMENT.
A swollen or thickened condition (renflement) is usually the result of a
disproportionate formation of the cellular tissue as contrasted with the
woody framework of the plant. We see marked instances of it in cultivated
carrots and turnips, the normal condition of the roots or root-stocks in these
plants being one of considerable hardness and toughness, and their form
slender, tapering, and more or less branched.
Page 521
The disproportionate development of cellular tissue is also seen in tubers
and bulbs, and in the swollen stems of such plants as Echinocactus,
Adenium obesum, some species of Vitis, &c. So, too, the upper portion of
the flower-stalk occasionally becomes much dilated, so as ultimately to
form a portion of the fruit. But it is not necessary to give farther illustrations
of this common tendency in some organs to become hypertrophied. As a
result of injury from insects or fungi, galls and excrescences of various
kinds are very common, but their consideration lies beyond the scope of the
present work.
Fig. 200.—Pelargonium, one branch of which was hypertrophied.
Enlargement of axile organs.—All the species of Pelargonium, Geranium,
Mirabilis, as well as those of Caryophylleæ and other orders, have tumid
nodes as a normal occurrence. In the genus Pelargonium this swelling is
sometimes not confined to the nodes, but extends to the interspaces between
them, e.g. P. spinosum. This condition, which happens as a natural feature
and bulbs, and in the swollen stems of such plants as Echinocactus,
Adenium obesum, some species of Vitis, &c. So, too, the upper portion of
the flower-stalk occasionally becomes much dilated, so as ultimately to
form a portion of the fruit. But it is not necessary to give farther illustrations
of this common tendency in some organs to become hypertrophied. As a
result of injury from insects or fungi, galls and excrescences of various
kinds are very common, but their consideration lies beyond the scope of the
present work.
Fig. 200.—Pelargonium, one branch of which was hypertrophied.
Enlargement of axile organs.—All the species of Pelargonium, Geranium,
Mirabilis, as well as those of Caryophylleæ and other orders, have tumid
nodes as a normal occurrence. In the genus Pelargonium this swelling is
sometimes not confined to the nodes, but extends to the interspaces between
them, e.g. P. spinosum. This condition, which happens as a natural feature
Page 522
in the species just named, may also occur as an exceptional thing in others.
The author is indebted to Dr. Sankey for a branch of Pelargonium which
was thus thickened, the remaining branches not being in any way affected.
The leaves on the swollen branch were smaller than the others, and their
stalks more flattened. There was, in this instance, no trace of fungus or
insect to account for the swelling of a single branch, which might, therefore,
be due to bud-variation, perhaps to reversion to some ancestral form. The
repeated cross fertilisations to which Pelargoniums have been subjected
render this hypothesis not an improbable one.
As an accompaniment to a spiral torsion of the woody fibres, this distension
of the stem is frequently met with, as in Valeriana, Dipsacus, &c. (See
Spiral Torsion.)
Knaurs.—On certain trees, such as the oak, the hornbeam, some species of
Cratægus, &c., hard woody lumps may occasionally be seen projecting,
varying greatly in size, from that of a pea to that of a cocoa-nut. They are
covered with bark, and consist in the interior of very hard layers of wood
disposed irregularly, so as to form objects of beauty for cabinet-makers'
purposes. From the frequent presence of small atrophied leaf-buds on their
surface, it would seem as if the structures in question were shortened
branches, in which the woody layers had become inordinately developed, as
if by compensation for the curtailment in length.[488] The cause of their
formation is not known, but it has been ascertained that they are not due to
insect agency. Knaurs may occasionally be used for purposes of
propagation, as in the case of the "uovoli of the olive" and the "burrs" that
are formed on some varieties of apple, from which both roots and leaf-
shoots are produced in abundance.
A distinction must be drawn between those instances in which the swelling
is solid throughout from the excessive formation of cellular tissue, and
The author is indebted to Dr. Sankey for a branch of Pelargonium which
was thus thickened, the remaining branches not being in any way affected.
The leaves on the swollen branch were smaller than the others, and their
stalks more flattened. There was, in this instance, no trace of fungus or
insect to account for the swelling of a single branch, which might, therefore,
be due to bud-variation, perhaps to reversion to some ancestral form. The
repeated cross fertilisations to which Pelargoniums have been subjected
render this hypothesis not an improbable one.
As an accompaniment to a spiral torsion of the woody fibres, this distension
of the stem is frequently met with, as in Valeriana, Dipsacus, &c. (See
Spiral Torsion.)
Knaurs.—On certain trees, such as the oak, the hornbeam, some species of
Cratægus, &c., hard woody lumps may occasionally be seen projecting,
varying greatly in size, from that of a pea to that of a cocoa-nut. They are
covered with bark, and consist in the interior of very hard layers of wood
disposed irregularly, so as to form objects of beauty for cabinet-makers'
purposes. From the frequent presence of small atrophied leaf-buds on their
surface, it would seem as if the structures in question were shortened
branches, in which the woody layers had become inordinately developed, as
if by compensation for the curtailment in length.[488] The cause of their
formation is not known, but it has been ascertained that they are not due to
insect agency. Knaurs may occasionally be used for purposes of
propagation, as in the case of the "uovoli of the olive" and the "burrs" that
are formed on some varieties of apple, from which both roots and leaf-
shoots are produced in abundance.
A distinction must be drawn between those instances in which the swelling
is solid throughout from the excessive formation of cellular tissue, and
Page 523
those wherein it is hollow from the more rapid growth of the outer as
contrasted with the inner portions. These latter cases might be classed under
the head of distension.
Fig. 201.—Formation of tubers or hypertrophied buds in the axils of leaves in the potato.
Enlargement of the buds may be seen in the case of bulbs and tubers.
Occasionally these organs are developed in the axils of leaves, when their
nature becomes apparent. A swollen bud or bulbil in this situation is not
uncommon in some cultivated tulips and lilies. The presence of small tubers
in the axils of the leaves in the potato, as shown in fig. 201, is also not
unfrequent.
contrasted with the inner portions. These latter cases might be classed under
the head of distension.
Fig. 201.—Formation of tubers or hypertrophied buds in the axils of leaves in the potato.
Enlargement of the buds may be seen in the case of bulbs and tubers.
Occasionally these organs are developed in the axils of leaves, when their
nature becomes apparent. A swollen bud or bulbil in this situation is not
uncommon in some cultivated tulips and lilies. The presence of small tubers
in the axils of the leaves in the potato, as shown in fig. 201, is also not
unfrequent.
Page 524
Fig. 202.—Inflorescence of ash (Fraxinus), with hypertrophied pedicels, flowers absent.
Enlargement of the flower-stalk.—The cauliflower and broccoli afford
familiar illustrations of hypertrophy of the flower-stalk, accompanied by a
corresponding defective development of the flowers. In the case of the ash
the terminal pedicels occasionally become swollen and distorted, while the
flowers are completely deficient, as shown in the adjacent cut (fig. 202).
In grapes a similar condition may occasionally be met with in which the
terminal pedicels become greatly swollen and fused into a solid mass. It
would seem probable that this change is due to insect puncture, or to the
effect of fungus growth at an early stage of development, but as to this point
there is at present no evidence.[489]
Enlargement of the flower-stalk.—The cauliflower and broccoli afford
familiar illustrations of hypertrophy of the flower-stalk, accompanied by a
corresponding defective development of the flowers. In the case of the ash
the terminal pedicels occasionally become swollen and distorted, while the
flowers are completely deficient, as shown in the adjacent cut (fig. 202).
In grapes a similar condition may occasionally be met with in which the
terminal pedicels become greatly swollen and fused into a solid mass. It
would seem probable that this change is due to insect puncture, or to the
effect of fungus growth at an early stage of development, but as to this point
there is at present no evidence.[489]
Page 525
Fig. 203.—Monstrous pear, showing extension and ramification of the succulent floral axis. The
bases of the sepals are also succulent.
In the apple a dilatation of the flower-stalk below the ordinary fruit may
occasionally be observed, thus giving rise to the appearance of two fruits
superposed and separated one from the other by a constriction. (See fig.
176, p. 327.) The lower swelling is entirely axial in these cases, as no trace
of carpels is to be seen. M. Carrière[490] mentions an instance wherein from
the base of one apple projected a second smaller one, destitute of carpels,
but surmounted by calyx-lobes as usual. The direction of this
supernumerary apple was the exact opposite of that of the primary fruit.
bases of the sepals are also succulent.
In the apple a dilatation of the flower-stalk below the ordinary fruit may
occasionally be observed, thus giving rise to the appearance of two fruits
superposed and separated one from the other by a constriction. (See fig.
176, p. 327.) The lower swelling is entirely axial in these cases, as no trace
of carpels is to be seen. M. Carrière[490] mentions an instance wherein from
the base of one apple projected a second smaller one, destitute of carpels,
but surmounted by calyx-lobes as usual. The direction of this
supernumerary apple was the exact opposite of that of the primary fruit.
Page 526
Fig. 204.—Monstrous pear, showing extension and swelling of axis, &c.
In pears, quinces, and apples, a not uncommon deviation is one in which the
axis is prolonged beyond the ordinary fruit, like which it is much swollen.
Occasionally the axis is not only prolonged, but even ramifies, the branches
partaking of the succulent character of the ordinary pome. Such instances
are frequently classed under the head of prolification, but they have in
general no claim to be considered in this light, for the reasons already given
in the chapter relating to that subject. (See p. 135.)[491]
A very curious illustration of hypertrophy of the flower-stalk is recorded
and figured by M. Carrière[492] in the cherry. The calyx in these fruits was
completely superior, the succulent portion of the fruit being made up of the
dilated extremity of the peduncle, and possibly in part of the base of the
calyx. The general appearance was thus that of a crab-apple. There was no
stone in the interior, but simply a rudimentary kernel or seed.[493]
In pears, quinces, and apples, a not uncommon deviation is one in which the
axis is prolonged beyond the ordinary fruit, like which it is much swollen.
Occasionally the axis is not only prolonged, but even ramifies, the branches
partaking of the succulent character of the ordinary pome. Such instances
are frequently classed under the head of prolification, but they have in
general no claim to be considered in this light, for the reasons already given
in the chapter relating to that subject. (See p. 135.)[491]
A very curious illustration of hypertrophy of the flower-stalk is recorded
and figured by M. Carrière[492] in the cherry. The calyx in these fruits was
completely superior, the succulent portion of the fruit being made up of the
dilated extremity of the peduncle, and possibly in part of the base of the
calyx. The general appearance was thus that of a crab-apple. There was no
stone in the interior, but simply a rudimentary kernel or seed.[493]
Page 527
Moquin-Tandon records an instance in which the stamens of each individual
flower in the inflorescence of a vine were hypertrophied, the sepals, petals,
and other organs of the flower, being proportionately diminished.[494]
In this place may also be mentioned the hypertrophied condition of the
placenta observed by Alphonse de Candolle in a species of Solanum, and
also in a species of Melastoma. Not only was the placenta unusually large in
these flowers, but it also protruded beyond the ovary.[495] A similar state of
things in Lobelia and Cuphea has already been alluded to under the head of
Alterations of Direction (p. 210).
The following singular growth in a tomato is described by the Rev. M. J.
Berkeley in the 'Gardeners' Chronicle' for 1866, p. 1217, and appears to
have been an extension of the placenta:—"On the first glance it seemed as
if an unusually large grape-stone had accidentally fallen on the upper
surface of the fruit, and was attached by the narrow base. The process was,
however, five lines long, and much narrowed below, besides which, though
it was pale green above, the base was coral-red, like the tomato itself. It
grew on a narrow and shallow crack on the surface of the fruit, and was
found below to communicate directly with a fibro-vascular bundle, which
entered into the composition of a portion of the placenta. On making a
vertical section, instead of being succulent, as I expected, it was white and
spongy within, with several lacunae, and one or two irregular fibro-vascular
bundles, with highly developed spiral vessels threading the centre. These
vessels, moreover, were tinged with brown, as in many cases of diseased
tissues. There was not the slightest appearance of placentæ or anything
indicating an abortive fruit. On closer examination the cuticle was found to
consist of thick-walled cells, exactly like those of the tomato, while the
spongy mass consisted of a similar tissue to the fleshy portion of the fruit,
but with far less wrinkled walls, and more indistinct intercellular spaces.
The most striking point, however, was the immense quantity of very
flower in the inflorescence of a vine were hypertrophied, the sepals, petals,
and other organs of the flower, being proportionately diminished.[494]
In this place may also be mentioned the hypertrophied condition of the
placenta observed by Alphonse de Candolle in a species of Solanum, and
also in a species of Melastoma. Not only was the placenta unusually large in
these flowers, but it also protruded beyond the ovary.[495] A similar state of
things in Lobelia and Cuphea has already been alluded to under the head of
Alterations of Direction (p. 210).
The following singular growth in a tomato is described by the Rev. M. J.
Berkeley in the 'Gardeners' Chronicle' for 1866, p. 1217, and appears to
have been an extension of the placenta:—"On the first glance it seemed as
if an unusually large grape-stone had accidentally fallen on the upper
surface of the fruit, and was attached by the narrow base. The process was,
however, five lines long, and much narrowed below, besides which, though
it was pale green above, the base was coral-red, like the tomato itself. It
grew on a narrow and shallow crack on the surface of the fruit, and was
found below to communicate directly with a fibro-vascular bundle, which
entered into the composition of a portion of the placenta. On making a
vertical section, instead of being succulent, as I expected, it was white and
spongy within, with several lacunae, and one or two irregular fibro-vascular
bundles, with highly developed spiral vessels threading the centre. These
vessels, moreover, were tinged with brown, as in many cases of diseased
tissues. There was not the slightest appearance of placentæ or anything
indicating an abortive fruit. On closer examination the cuticle was found to
consist of thick-walled cells, exactly like those of the tomato, while the
spongy mass consisted of a similar tissue to the fleshy portion of the fruit,
but with far less wrinkled walls, and more indistinct intercellular spaces.
The most striking point, however, was the immense quantity of very
Page 528
irregular and unequal starch-grains with which they were gorged, which
gave a peculiar sparkling appearance to them when seen en masse. I am
inclined to regard the body rather as an abortive axis than an undeveloped
fruit. In almost all, if not all, these cases of abnormal growth, whether from
leaves, petioles, fruit, or other portions of the plant, we find an immediate
connection with one or more spiral vessels, which if not existent at first are
developed sooner or later. In the present case the connection of the fibro-
vascular tissue of the fruit and abnormal growth was plain enough, but
whether it existed when the body was first given off I am unable to say, as it
was fully developed when the fruit was brought to me."
Enlargement of the leaves.—Increase in the size or substance of leaves
takes places in several ways, and affects the whole or only certain portions
of them. The simplest form of this malformation is met with in our
cabbages, which, by the art of the gardener, have been made to produce
leaves of greater size and thickness than those which are developed in the
wild form. In such instances the whole substance of the leaf is increased in
bulk, and the increase affects the fibrous framework of the leaves as well as
the cellular portions, though the exaggerated development of the latter is
out of proportion to that of the former.
In some species of Podocarpus there may occasionally be seen at the base
of the branchlets a dozen or more fleshy scales, of a rose colour, passing
gradually into the ordinary leaves of the plant, and evidently analogous to
the three fleshy confluent bracts which surround the ripe fruit.
In other instances, while the fibrous framework of the leaf retains its usual
degree of development, the cellular parenchyma is developed in excess,
and, if the increase is so arranged that the number of superposed layers of
the cellular tissue is not increased, or their thickness exaggerated, then we
get such leaves as those of the "kail," or of the "Savoys" leaves, which are
gave a peculiar sparkling appearance to them when seen en masse. I am
inclined to regard the body rather as an abortive axis than an undeveloped
fruit. In almost all, if not all, these cases of abnormal growth, whether from
leaves, petioles, fruit, or other portions of the plant, we find an immediate
connection with one or more spiral vessels, which if not existent at first are
developed sooner or later. In the present case the connection of the fibro-
vascular tissue of the fruit and abnormal growth was plain enough, but
whether it existed when the body was first given off I am unable to say, as it
was fully developed when the fruit was brought to me."
Enlargement of the leaves.—Increase in the size or substance of leaves
takes places in several ways, and affects the whole or only certain portions
of them. The simplest form of this malformation is met with in our
cabbages, which, by the art of the gardener, have been made to produce
leaves of greater size and thickness than those which are developed in the
wild form. In such instances the whole substance of the leaf is increased in
bulk, and the increase affects the fibrous framework of the leaves as well as
the cellular portions, though the exaggerated development of the latter is
out of proportion to that of the former.
In some species of Podocarpus there may occasionally be seen at the base
of the branchlets a dozen or more fleshy scales, of a rose colour, passing
gradually into the ordinary leaves of the plant, and evidently analogous to
the three fleshy confluent bracts which surround the ripe fruit.
In other instances, while the fibrous framework of the leaf retains its usual
degree of development, the cellular parenchyma is developed in excess,
and, if the increase is so arranged that the number of superposed layers of
the cellular tissue is not increased, or their thickness exaggerated, then we
get such leaves as those of the "kail," or of the "Savoys" leaves, which are
Page 529
technically called by descriptive botanists "folia bullata." In such leaves the
disc of the leaf, rather than the margin, is increased and its surface is thrown
up into little conical projections, which are hollow on the under side.
But leaves may increase beyond their usual size without such grave
alterations of form as those to which allusion has just been made. It is well
known that if a tree be cut down and new shoots be sent out from the stump,
the leaves formed on these shoots very often greatly exceed the ordinary
ones in dimensions. Such cases as this hardly come under the head of
malformations. But where one part only of the leaf is excessively
developed, the other portion remaining in its ordinary condition, there can
be no hesitation in ranking the phenomenon as teratological.
Thus, Moquin says that the median nerve may be prolonged beyond the
blade of the leaf in the form of a short strap or ribbon-like excrescence,
while, at other times, the lateral parts of the leaf are subjected to undue
development. He refers to a case cited by Schlotterbec[496] in which each
side of the leaves of a yellow "violier" (wallflower) was dilated into a kind
of projecting lobe on either side of the true apex of the leaf, thus rendering
it in appearance three-lobed. M. Delavaud[497] puts on record a case of
hypertrophy in the leaves of the common elm, resulting in the formation of
an additional lobe and a return to the tricostate type. A leaf so affected is
stated to have presented the appearance of a fusion of two leaves. (See also
Multiplication of leaves, p. 353.)
The hypertrophied and coloured leaf of Gesnera occupying the place of the
absent inflorescence has been previously alluded to under the head of
displacement (p. 88).
In some instances hypertrophy is the opposite of suppression; as in the case
previously mentioned, where the stipule in the inflorescence of a pea, which
disc of the leaf, rather than the margin, is increased and its surface is thrown
up into little conical projections, which are hollow on the under side.
But leaves may increase beyond their usual size without such grave
alterations of form as those to which allusion has just been made. It is well
known that if a tree be cut down and new shoots be sent out from the stump,
the leaves formed on these shoots very often greatly exceed the ordinary
ones in dimensions. Such cases as this hardly come under the head of
malformations. But where one part only of the leaf is excessively
developed, the other portion remaining in its ordinary condition, there can
be no hesitation in ranking the phenomenon as teratological.
Thus, Moquin says that the median nerve may be prolonged beyond the
blade of the leaf in the form of a short strap or ribbon-like excrescence,
while, at other times, the lateral parts of the leaf are subjected to undue
development. He refers to a case cited by Schlotterbec[496] in which each
side of the leaves of a yellow "violier" (wallflower) was dilated into a kind
of projecting lobe on either side of the true apex of the leaf, thus rendering
it in appearance three-lobed. M. Delavaud[497] puts on record a case of
hypertrophy in the leaves of the common elm, resulting in the formation of
an additional lobe and a return to the tricostate type. A leaf so affected is
stated to have presented the appearance of a fusion of two leaves. (See also
Multiplication of leaves, p. 353.)
The hypertrophied and coloured leaf of Gesnera occupying the place of the
absent inflorescence has been previously alluded to under the head of
displacement (p. 88).
In some instances hypertrophy is the opposite of suppression; as in the case
previously mentioned, where the stipule in the inflorescence of a pea, which
Page 530
is usually undeveloped and rudimentary, was developed in the form of a
leafy cup or pitcher.
Another instance of the development of parts usually suppressed, is
afforded by the bud-scales of Magnolia fuscata, which may sometimes be
found with small but perfect leaves projecting from them, the leaf in this
case being the lamina which is ordinarily abortive, while the scales are the
representatives of the stipules. This condition is said by Hooker and
Thomson ('Flora Indica,' p. 73) to be constant in Magnolia Campbelli.
Enlargement of the perianth, &c.—One or all the segments of the
perianth may be subjected to hypertrophy; thus, the utricle of Carex vulpina
may frequently be observed to attain four or five times its usual size, the
contained ovary remaining unaffected. This condition is generally the result
of insect puncture. The growth of parasitic fungi will produce a similar
result, as is often seen in the common shepherd's purse, Thlaspi bursa
pastoris, and other Cruciferæ. The perianth of Rumex aquaticus has been
also observed to be occasionally hypertrophied in conjunction with a
similar condition of the pistil and with atrophy of the ovules.
Moquin relates having found flowers of Salsola Kali and of Chenopodium
murale in which some of the segments of the perianth were five or six times
larger than they should be.
leafy cup or pitcher.
Another instance of the development of parts usually suppressed, is
afforded by the bud-scales of Magnolia fuscata, which may sometimes be
found with small but perfect leaves projecting from them, the leaf in this
case being the lamina which is ordinarily abortive, while the scales are the
representatives of the stipules. This condition is said by Hooker and
Thomson ('Flora Indica,' p. 73) to be constant in Magnolia Campbelli.
Enlargement of the perianth, &c.—One or all the segments of the
perianth may be subjected to hypertrophy; thus, the utricle of Carex vulpina
may frequently be observed to attain four or five times its usual size, the
contained ovary remaining unaffected. This condition is generally the result
of insect puncture. The growth of parasitic fungi will produce a similar
result, as is often seen in the common shepherd's purse, Thlaspi bursa
pastoris, and other Cruciferæ. The perianth of Rumex aquaticus has been
also observed to be occasionally hypertrophied in conjunction with a
similar condition of the pistil and with atrophy of the ovules.
Moquin relates having found flowers of Salsola Kali and of Chenopodium
murale in which some of the segments of the perianth were five or six times
larger than they should be.
Page 531
Fig. 205.—Hypertrophy of the perianth in Cocos nucifera.
The adjoining woodcut represents a singular condition of some cocoa-nuts
in the Kew Museum, the appearance of which is due apparently to an
hypertrophied condition of the segments of the perianth, which have not
only increased in length as the central nut has ripened, but have developed
in their tissues that fibrous tissue which ordinarily is found in the pericarp
only. This view of the structure of these nuts is borne out by the fact that,
under normal circumstances, the base of the perianth contains a
considerable amount of fibrous material. In the present case this has
increased to such an extent that the fruit appears surrounded by a double
husk, by an inner one as usual, and by an outer six-parted one.
It will be remembered that in some of the Cinchonaceæ, e.g. Mussænda,
Pinckneya, Calycophyllum, one or more of the calycine lobes are normally
dilated and petaloid, the others remaining small and comparatively
inconspicuous. Inequality in size is, indeed, a common occurrence in the
The adjoining woodcut represents a singular condition of some cocoa-nuts
in the Kew Museum, the appearance of which is due apparently to an
hypertrophied condition of the segments of the perianth, which have not
only increased in length as the central nut has ripened, but have developed
in their tissues that fibrous tissue which ordinarily is found in the pericarp
only. This view of the structure of these nuts is borne out by the fact that,
under normal circumstances, the base of the perianth contains a
considerable amount of fibrous material. In the present case this has
increased to such an extent that the fruit appears surrounded by a double
husk, by an inner one as usual, and by an outer six-parted one.
It will be remembered that in some of the Cinchonaceæ, e.g. Mussænda,
Pinckneya, Calycophyllum, one or more of the calycine lobes are normally
dilated and petaloid, the others remaining small and comparatively
inconspicuous. Inequality in size is, indeed, a common occurrence in the
Page 532
sepals of many natural orders—Polygalaceæ, Leguminosæ, Labiatæ, &c.
The flowers of a rose are mentioned by Moquin as having presented an
enlargement of the calyx without any other alterations in form.
Schlechtendal has noticed the same thing in Papaver Rhœas, Reichenbach
in Campanula persicifolia, and A. de Candolle in C. Rapunculus. M.
Brongniart also has recorded[498] a remarkable variety of Primula sinensis
cultivated in the Jardin des Plantes at Paris, wherein the calyx is
enormously developed. MM. Fournier and Bonnet have described flowers
of Rubus with hypertrophied calyx in conjunction with atrophy and
virescence of the petals and other changes.[499]
The corolla may be hypertrophied in some cases, though the change is more
rare than in most other organs. Moquin-Tandon mentions as subject to this
anomaly species of Galeopsis, Prunella, Scabiosa, and Dipsacus, and also
mentions a remarkable variety of Viola odorata cultivated in the
neighbourhood of Toulouse. The same learned author also alludes to the so-
called double Composites, viz. those in which the usually tubular florets of
the disc assume the form and proportions of those of the ray, but these are
hardly cases of hypertrophy.
Enlargement of the andrœcium.—Dunal[500] alludes to a curious instance
in a species of Verbascum, the lower flowers of which had hairy stamens as
usual, but the filaments of the topmost flower were quite destitute of hairs,
and dilated like a flat ribbon.
Moquin relates having found in the neighbourhood of Toulouse a plant of
Solanum Dulcamara in which all the upper flowers had two or three
stamens of larger dimensions than the others. This happens habitually in
Solanum tridynamum and S. Amazonicum, and to a less extent in S.
vespertilio and S. cornutum; also in some species of Hyoscyamus. These
The flowers of a rose are mentioned by Moquin as having presented an
enlargement of the calyx without any other alterations in form.
Schlechtendal has noticed the same thing in Papaver Rhœas, Reichenbach
in Campanula persicifolia, and A. de Candolle in C. Rapunculus. M.
Brongniart also has recorded[498] a remarkable variety of Primula sinensis
cultivated in the Jardin des Plantes at Paris, wherein the calyx is
enormously developed. MM. Fournier and Bonnet have described flowers
of Rubus with hypertrophied calyx in conjunction with atrophy and
virescence of the petals and other changes.[499]
The corolla may be hypertrophied in some cases, though the change is more
rare than in most other organs. Moquin-Tandon mentions as subject to this
anomaly species of Galeopsis, Prunella, Scabiosa, and Dipsacus, and also
mentions a remarkable variety of Viola odorata cultivated in the
neighbourhood of Toulouse. The same learned author also alludes to the so-
called double Composites, viz. those in which the usually tubular florets of
the disc assume the form and proportions of those of the ray, but these are
hardly cases of hypertrophy.
Enlargement of the andrœcium.—Dunal[500] alludes to a curious instance
in a species of Verbascum, the lower flowers of which had hairy stamens as
usual, but the filaments of the topmost flower were quite destitute of hairs,
and dilated like a flat ribbon.
Moquin relates having found in the neighbourhood of Toulouse a plant of
Solanum Dulcamara in which all the upper flowers had two or three
stamens of larger dimensions than the others. This happens habitually in
Solanum tridynamum and S. Amazonicum, and to a less extent in S.
vespertilio and S. cornutum; also in some species of Hyoscyamus. These
Page 533
cases show the close affinity between the Solanaceæ and the
Scrophulariaceæ.
Enlargement of the gynœcium.—In some flowers which have become
accidentally female the pistil becomes unusually large, and even to such an
extent as to prevent the passage of the pollen. Moquin remarks having seen
this enlargement in the pistils of Suæda fruticosa and Kochia scoparia. The
flowers of these Chenopods, under these circumstances, resemble the
female flowers of some nettles. The styles of Anemone are also much
enlarged as the result of cultivation, and from their petaloid appearance
resemble those of the Iris (Goethe). MM. Seringe and Heyland[501] have
figured some anomalous flowers of Diplotaxis tenuifolia in which the pistil,
more or less distended and deformed, was considerably elongated below, so
that it seemed to be borne upon a long stalk, analogous to that of fruits of
Capparids. Dr. Klinsman[502] mentions an instance of a similar kind
combined with hypertrophy of the sepals and pistils; indeed, the alteration
is not uncommon among Crucifers. Pyrethrum inodorum is very subject to
hypertrophy. The styles of its radial florets become elongated without any
other alteration; at the same time the small corollas become green, and
show a tendency to assume a foliaceous condition. Sometimes the
hypertrophy affects also the styles of the central florets, and these also
become enlarged to double or treble their usual dimensions.
Linné has remarked that the ovary of Tragopogon sometimes assumes very
large dimensions, as also does the pappus. He mentions a double-flowered
variety, the ovaries of which become ten or twelve times larger than
ordinary. M. Clos[503] records an instance in Rumex scutatus wherein the
pistil was hypertrophied or club-shaped, and open at the top, or in other
cases funnel-shaped, three-lobed at the summit, each lobe terminated by a
style. One of the most frequent causes tending to the hypertrophy of the
pistil is attributable to the puncture of insects; thus, when the ovary of
Scrophulariaceæ.
Enlargement of the gynœcium.—In some flowers which have become
accidentally female the pistil becomes unusually large, and even to such an
extent as to prevent the passage of the pollen. Moquin remarks having seen
this enlargement in the pistils of Suæda fruticosa and Kochia scoparia. The
flowers of these Chenopods, under these circumstances, resemble the
female flowers of some nettles. The styles of Anemone are also much
enlarged as the result of cultivation, and from their petaloid appearance
resemble those of the Iris (Goethe). MM. Seringe and Heyland[501] have
figured some anomalous flowers of Diplotaxis tenuifolia in which the pistil,
more or less distended and deformed, was considerably elongated below, so
that it seemed to be borne upon a long stalk, analogous to that of fruits of
Capparids. Dr. Klinsman[502] mentions an instance of a similar kind
combined with hypertrophy of the sepals and pistils; indeed, the alteration
is not uncommon among Crucifers. Pyrethrum inodorum is very subject to
hypertrophy. The styles of its radial florets become elongated without any
other alteration; at the same time the small corollas become green, and
show a tendency to assume a foliaceous condition. Sometimes the
hypertrophy affects also the styles of the central florets, and these also
become enlarged to double or treble their usual dimensions.
Linné has remarked that the ovary of Tragopogon sometimes assumes very
large dimensions, as also does the pappus. He mentions a double-flowered
variety, the ovaries of which become ten or twelve times larger than
ordinary. M. Clos[503] records an instance in Rumex scutatus wherein the
pistil was hypertrophied or club-shaped, and open at the top, or in other
cases funnel-shaped, three-lobed at the summit, each lobe terminated by a
style. One of the most frequent causes tending to the hypertrophy of the
pistil is attributable to the puncture of insects; thus, when the ovary of
Page 534
Juncus articulatus is thus punctured, it acquires a size two or three times
larger than ordinary, becoming at the same time sterile.[504]
Occasionally the enlargement may be due to a fusion or incorporation of
other elements; thus, M. Lemaire describes an instance in which the style of
Sinningia purpurea was much larger than ordinary, tubular, bearing three
small lobes, and altogether bearing much resemblance to the column or
"gynosteme" of Orchids. This appearance was due to the cohesion and
intimate union of the styles with three abortive stamens.[505]
Enlargement of the fruit.—Most cultivated fruits are in a state of true
hypertrophy. Girod de Chantrans, after many trials, succeeded in producing
a peculiar variety of pea with pods double the ordinary size.[506] M.
Clos[507] mentions a case wherein the carpels of Delphinium dictyocarpum
were hypertrophied. The change in size may or may not be attended by a
difference in form; thus, in certain Leguminosæ, as Medicago lupulina,
Melilotus leucantha, the carpels are sometimes hypertrophied and
elongated, so as to resemble a claw or hook.[508]
The fruit of the common groundsel (Senecio vulgaris) is in its normal
condition two or three times shorter than the involucre, and cylindrical for
its whole length, but it frequently happens that the fruits become as long as
the involucre itself, and taper from the base upwards, so as to become
beaked. Under this head may also be mentioned the fleshy bulbils that are
found in the capsules of Crinum, Amaryllis, and Agave. These are true seeds
enormously dilated.[509] In these seeds the outer coating becomes very thick
and fleshy, and is traversed by spiral vessels.
It is obvious that very important results in a practical point of view may be
and have been arrived at by cultivators availing themselves of this tendency
of plants to increase in dimensions under certain circumstances. It is
larger than ordinary, becoming at the same time sterile.[504]
Occasionally the enlargement may be due to a fusion or incorporation of
other elements; thus, M. Lemaire describes an instance in which the style of
Sinningia purpurea was much larger than ordinary, tubular, bearing three
small lobes, and altogether bearing much resemblance to the column or
"gynosteme" of Orchids. This appearance was due to the cohesion and
intimate union of the styles with three abortive stamens.[505]
Enlargement of the fruit.—Most cultivated fruits are in a state of true
hypertrophy. Girod de Chantrans, after many trials, succeeded in producing
a peculiar variety of pea with pods double the ordinary size.[506] M.
Clos[507] mentions a case wherein the carpels of Delphinium dictyocarpum
were hypertrophied. The change in size may or may not be attended by a
difference in form; thus, in certain Leguminosæ, as Medicago lupulina,
Melilotus leucantha, the carpels are sometimes hypertrophied and
elongated, so as to resemble a claw or hook.[508]
The fruit of the common groundsel (Senecio vulgaris) is in its normal
condition two or three times shorter than the involucre, and cylindrical for
its whole length, but it frequently happens that the fruits become as long as
the involucre itself, and taper from the base upwards, so as to become
beaked. Under this head may also be mentioned the fleshy bulbils that are
found in the capsules of Crinum, Amaryllis, and Agave. These are true seeds
enormously dilated.[509] In these seeds the outer coating becomes very thick
and fleshy, and is traversed by spiral vessels.
It is obvious that very important results in a practical point of view may be
and have been arrived at by cultivators availing themselves of this tendency
of plants to increase in dimensions under certain circumstances. It is
Page 535
needless to do more than refer to the many fruits, vegetables, and cereals,
which have thus become enlarged and improved by careful selection and
rearing.
Alterations of consistence often accompany changes in size. The change
may be one whereby the tissues become unusually hardened, by the
excessive formation of secondary woody deposits, or softer and more
succulent than ordinary, from the formation of an inordinate amount of
loose cellular tissue. Generally speaking, the appearances presented in such
cases are not sufficiently striking to demand notice other than as regards
their size. One illustration, however, may be cited from its singularity. This
was the case of a dahlia, in which the centre of the flower was occupied by
a projecting knob as large as a walnut, brown in colour, and very hard in
texture. This knob was nothing but the enlarged and indurated extremity of
the common receptacle, destitute of the scales and florets which usually
spring from it. No insect-puncture could be detected, and no other reason
for this peculiarity could be ascertained.
FOOTNOTES:
[488] On the subject of knaurs, the reader is referred to Trécul, 'Ann. Sc. Nat.,' 3 ser., vol. xx,
p. 65; Lindley, 'Theory of Horticulture;' Rev. M. J. Berkeley, 'Gardeners' Chronicle,' 1855, p.
756.
[489] Jaeger, 'Flora.' 1860. p. 49, tab. i.
[490] 'Revue Horticole,' 1868, p. 110, figs. 12, 13.
[491] The reader may also refer for further information on the subject of malformed pears to
Irmisch. 'Flora,' 1858, p. 38, tab. i; Lindley, 'Theory of Horticulture'; Caspary, 'Bull. Soc. Bot.
France,' vol. vi, 1859 (Rev. Bibl.), p. 235; Duhamel, 'Phys. Arbr.,' liv. iii, cap. 3. p. 393, fig.
308; Bonnet, 'Recherch. Us. feuilles,' tab. xxvi, fig. 2; Moquin-Tandon, 'El. Ter. Veg.,' p. 384,
&c. Some of the cases recorded are, however, instances of true prolification.
[492] 'Revue Horticole' 1868, p. 310.
which have thus become enlarged and improved by careful selection and
rearing.
Alterations of consistence often accompany changes in size. The change
may be one whereby the tissues become unusually hardened, by the
excessive formation of secondary woody deposits, or softer and more
succulent than ordinary, from the formation of an inordinate amount of
loose cellular tissue. Generally speaking, the appearances presented in such
cases are not sufficiently striking to demand notice other than as regards
their size. One illustration, however, may be cited from its singularity. This
was the case of a dahlia, in which the centre of the flower was occupied by
a projecting knob as large as a walnut, brown in colour, and very hard in
texture. This knob was nothing but the enlarged and indurated extremity of
the common receptacle, destitute of the scales and florets which usually
spring from it. No insect-puncture could be detected, and no other reason
for this peculiarity could be ascertained.
FOOTNOTES:
[488] On the subject of knaurs, the reader is referred to Trécul, 'Ann. Sc. Nat.,' 3 ser., vol. xx,
p. 65; Lindley, 'Theory of Horticulture;' Rev. M. J. Berkeley, 'Gardeners' Chronicle,' 1855, p.
756.
[489] Jaeger, 'Flora.' 1860. p. 49, tab. i.
[490] 'Revue Horticole,' 1868, p. 110, figs. 12, 13.
[491] The reader may also refer for further information on the subject of malformed pears to
Irmisch. 'Flora,' 1858, p. 38, tab. i; Lindley, 'Theory of Horticulture'; Caspary, 'Bull. Soc. Bot.
France,' vol. vi, 1859 (Rev. Bibl.), p. 235; Duhamel, 'Phys. Arbr.,' liv. iii, cap. 3. p. 393, fig.
308; Bonnet, 'Recherch. Us. feuilles,' tab. xxvi, fig. 2; Moquin-Tandon, 'El. Ter. Veg.,' p. 384,
&c. Some of the cases recorded are, however, instances of true prolification.
[492] 'Revue Horticole' 1868, p. 310.
Page 536
[493] The interest of this accident is great, as showing how an habitually superior ovary may
become inferior—a change so rare in its occurrence that its existence has been denied, and
thus forming a marked contrast with the frequency with which the converse change of an
inferior ovary to a superior one, from want of union with the calyx or from imperfect
development of the peduncle, may be observed. It is also interesting as showing how the
peduncle may become swollen, and at the same time how the woody deposit of the endocarp
may, as if by compensation, be deficient. And, again, the malformation is not without
significance in regard to the relationship between the drupaceous and the pomaceous
subdivisions of Rosaceæ. The case would fitly be included under alterations of position, but
the sheets relating to that subject were printed off before the publication of M. Carrière's
notice.
[494] 'Bull. Soc. Bot. France,' 1860, vol. vii, p. 881.
[495] "Monstr. Veget.," in 'Neue Denkschrift.'
[496] "Sched. de Monst. Plant." in 'Act. Helvet.,' t. ii, pl. ii, f. 14.
[497] 'Bull. Soc. Bot. France,' vol. viii, 1861, p. 144.
[498] 'Ann. Sc. Nat.,' sér. 2, t. i, p. 308, pl. ix c, fig. 1.
[499] 'Bull. Soc. Bot. France,' 1862, t. ix, p. 37.
[500] 'Consid. org. Fleur.,' Montpell., 1829, 25, 26, pl. ii, f. 18 and 19.
[501] 'Bull. Bot.,' t. i, p. 7, tab. 1.
[502] 'Linnæa,' vol. x, p. 604, tab. 5.
[503] 'Mém. Acad. Sc. Toulouse,' 5 ser., vol. iii.
[504] 'Ré. nosol. Végét.,' pp. 342.
[505] 'Illustr. Hortic.,' 1868, Misc., p. 62.
[506] 'Ann. Soc. Linn.,' Paris, t. i, p. 139.
[507] 'Mém. Acad. Toulouse,' t. 6, 1862.
[508] 'D. C. Prod.,' ii, pp. 172, 187.
become inferior—a change so rare in its occurrence that its existence has been denied, and
thus forming a marked contrast with the frequency with which the converse change of an
inferior ovary to a superior one, from want of union with the calyx or from imperfect
development of the peduncle, may be observed. It is also interesting as showing how the
peduncle may become swollen, and at the same time how the woody deposit of the endocarp
may, as if by compensation, be deficient. And, again, the malformation is not without
significance in regard to the relationship between the drupaceous and the pomaceous
subdivisions of Rosaceæ. The case would fitly be included under alterations of position, but
the sheets relating to that subject were printed off before the publication of M. Carrière's
notice.
[494] 'Bull. Soc. Bot. France,' 1860, vol. vii, p. 881.
[495] "Monstr. Veget.," in 'Neue Denkschrift.'
[496] "Sched. de Monst. Plant." in 'Act. Helvet.,' t. ii, pl. ii, f. 14.
[497] 'Bull. Soc. Bot. France,' vol. viii, 1861, p. 144.
[498] 'Ann. Sc. Nat.,' sér. 2, t. i, p. 308, pl. ix c, fig. 1.
[499] 'Bull. Soc. Bot. France,' 1862, t. ix, p. 37.
[500] 'Consid. org. Fleur.,' Montpell., 1829, 25, 26, pl. ii, f. 18 and 19.
[501] 'Bull. Bot.,' t. i, p. 7, tab. 1.
[502] 'Linnæa,' vol. x, p. 604, tab. 5.
[503] 'Mém. Acad. Sc. Toulouse,' 5 ser., vol. iii.
[504] 'Ré. nosol. Végét.,' pp. 342.
[505] 'Illustr. Hortic.,' 1868, Misc., p. 62.
[506] 'Ann. Soc. Linn.,' Paris, t. i, p. 139.
[507] 'Mém. Acad. Toulouse,' t. 6, 1862.
[508] 'D. C. Prod.,' ii, pp. 172, 187.
Page 537
[509] Richard, "Obs. sur les bulbilles des Crinum;" 'Ann. Sc. Nat.,' t. ii, p. 12. pl. i, fig. 1, 2.
See also A. Braun, "Mémoire sur les graines charnues des Amaryllidèes," &c.; 'Ann. Sc. Nat.,'
1860, vol. xiv, p. 1, tab. 1.
CHAPTER II.
ELONGATION.
The class of cases coming under this head are sufficiently indicated by the
name. There are many instances of this phenomenon occurring under
different conditions, which, though unusual, can hardly be called abnormal,
such, for instance, as the great lengthening of roots in their search for water,
the excessive elongation that takes place in plants when grown at a distance
from the light, in their endeavour to attain to which they become, as
gardeners phrase it, "drawn." A similar result is brought about in forests or
plantations, where long spars are required, by allowing the trees to grow
very close to each other, so as to prevent the lateral extension of the
branches. When plants grow in running water their roots, stems, and
sometimes their leaves, become excessively elongated, as in Ranunculus
fluitans, the flower-stalks of Valisneria spiralis, &c. These are cases of
variation rather than of malformation, but are none the less curious, or
sometimes perplexing; thus, Lapeyrouse described, in his 'Supplement à la
flore des Pyrenées,' p. 27, under the name Potamogeton bifolium, a plant
which Mr. Bentham subsequently discovered to be nothing but a flowerless
variety of Vicia Faba distorted by its growth in water.[510]
Elongation of the root.—This, as already remarked, is more often a
variation than a malformation, and is usually due to the presence of water at
a distance necessitating growth at the extremities of the root, or to the
presence of some obstacle, such as a stone, to avoid which the root
See also A. Braun, "Mémoire sur les graines charnues des Amaryllidèes," &c.; 'Ann. Sc. Nat.,'
1860, vol. xiv, p. 1, tab. 1.
CHAPTER II.
ELONGATION.
The class of cases coming under this head are sufficiently indicated by the
name. There are many instances of this phenomenon occurring under
different conditions, which, though unusual, can hardly be called abnormal,
such, for instance, as the great lengthening of roots in their search for water,
the excessive elongation that takes place in plants when grown at a distance
from the light, in their endeavour to attain to which they become, as
gardeners phrase it, "drawn." A similar result is brought about in forests or
plantations, where long spars are required, by allowing the trees to grow
very close to each other, so as to prevent the lateral extension of the
branches. When plants grow in running water their roots, stems, and
sometimes their leaves, become excessively elongated, as in Ranunculus
fluitans, the flower-stalks of Valisneria spiralis, &c. These are cases of
variation rather than of malformation, but are none the less curious, or
sometimes perplexing; thus, Lapeyrouse described, in his 'Supplement à la
flore des Pyrenées,' p. 27, under the name Potamogeton bifolium, a plant
which Mr. Bentham subsequently discovered to be nothing but a flowerless
variety of Vicia Faba distorted by its growth in water.[510]
Elongation of the root.—This, as already remarked, is more often a
variation than a malformation, and is usually due to the presence of water at
a distance necessitating growth at the extremities of the root, or to the
presence of some obstacle, such as a stone, to avoid which the root
Page 538
elongates till it has passed the obstruction. Occasionally in Crocus corms
some of the fibrils may be met with much lengthened and thickened, and
invested with a fleshy sheath. It is not certain, however, that these structures
are roots; possibly, nay probably, they may be processes from the stem
thrust downwards into the soil, similar to the formations already described
in the tulip (p. 85, fig. 39).
Elongation of the inflorescence.—Under this heading it is necessary to
consider lengthening of the common rachis in the case of an aggregate
inflorescence, and lengthening of the individual flower-stalks, whether they
be solitary or portions of a multiple inflorescence. The two phenomena may
occur together, but they are quite as often independent one of the other.
Thus, among Umbelliferæ the umbels are occasionally met with supported
on unusually long stalks, while the pedicels of the individual flowers may
or not be increased in length; so also with some of the Composites, or the
heads of flowers of some Leguminosæ, Trifolium repens, &c. &c.
Another illustration of the sort is that recorded by M. Fournier, wherein the
usually umbellate inflorescence of Pelargonium was, through the
lengthening of the main stalk, transformed into a raceme. Among
Composites a similar change may sometimes be met with.
MM. Clos and De Schönefeld have recorded the existence of a variety of
the sweet chestnut (Castanea)in which the female catkins were as long, and
bore nearly as many flowers, as the male spikes. This is stated to be of
constant occurrence in some localities, and to be accompanied by a
diminished size of the fruits. A similar elongation has been observed in the
case of the walnut, catkins of which have been seen bearing thirty to thirty-
five large nuts.[511]
some of the fibrils may be met with much lengthened and thickened, and
invested with a fleshy sheath. It is not certain, however, that these structures
are roots; possibly, nay probably, they may be processes from the stem
thrust downwards into the soil, similar to the formations already described
in the tulip (p. 85, fig. 39).
Elongation of the inflorescence.—Under this heading it is necessary to
consider lengthening of the common rachis in the case of an aggregate
inflorescence, and lengthening of the individual flower-stalks, whether they
be solitary or portions of a multiple inflorescence. The two phenomena may
occur together, but they are quite as often independent one of the other.
Thus, among Umbelliferæ the umbels are occasionally met with supported
on unusually long stalks, while the pedicels of the individual flowers may
or not be increased in length; so also with some of the Composites, or the
heads of flowers of some Leguminosæ, Trifolium repens, &c. &c.
Another illustration of the sort is that recorded by M. Fournier, wherein the
usually umbellate inflorescence of Pelargonium was, through the
lengthening of the main stalk, transformed into a raceme. Among
Composites a similar change may sometimes be met with.
MM. Clos and De Schönefeld have recorded the existence of a variety of
the sweet chestnut (Castanea)in which the female catkins were as long, and
bore nearly as many flowers, as the male spikes. This is stated to be of
constant occurrence in some localities, and to be accompanied by a
diminished size of the fruits. A similar elongation has been observed in the
case of the walnut, catkins of which have been seen bearing thirty to thirty-
five large nuts.[511]
Page 539
In the strobile of the hop, Humulus Lupulus, a like elongation may
sometimes be met with, generally in association with a more or less leafy
condition of some of the scales.
Of a similar character, but complicated with extrusion or eversion of an
ordinarily concave axis, is the fig described by Zuccarini,[512] and from the
appearances presented by which that author draws the inference that the
peculiar appearance of the fig is due to the formation of a large number of
small bracts blended together for the greater part of their length, and
accompanied by the suppression of the internodes, and consequent
shortening of the axis. In the monstrous fruit the axis is prolonged, and
forms a kind of raceme or catkin, surrounded at the base by numerous
bracts, as in many Amentaceæ. (See p. 204, figs. 105, 106.)
A lengthening of the axis of the female strobiles of Coniferæ is not of
infrequent occurrence in Cryptomeria japonica, Larie europæa, &c., and
this is usually associated, as has been before stated, with a leaf-like
condition of the bracts, and sometimes even with the development of leaf-
bearing shoots in place of the scales. (See under Prolification of
Inflorescence and Phyllomorphy, and for references, p. 115.)
Elongation of the secondary flower-stalks.—In the previous section the
effect of elongation of the main rachis has been considered. A
corresponding deviation occurs in the peduncles or pedicels, and sometimes
alters the general character of the inflorescence very considerably,
converting a spike into a raceme, a raceme into a corymb, a capitulum into
an umbel, and so forth. A few such cases may here be alluded to. Fig. 206
represents a specimen of Ranunculus acris, in which the lower and lateral
flower-stalks were not only increased in number, but so much lengthened as
to form a flat-topped inflorescence—a corymbose cyme. In many
leguminous plants, as in Trifolium repens, Lotus corniculatus, &c., what is
sometimes be met with, generally in association with a more or less leafy
condition of some of the scales.
Of a similar character, but complicated with extrusion or eversion of an
ordinarily concave axis, is the fig described by Zuccarini,[512] and from the
appearances presented by which that author draws the inference that the
peculiar appearance of the fig is due to the formation of a large number of
small bracts blended together for the greater part of their length, and
accompanied by the suppression of the internodes, and consequent
shortening of the axis. In the monstrous fruit the axis is prolonged, and
forms a kind of raceme or catkin, surrounded at the base by numerous
bracts, as in many Amentaceæ. (See p. 204, figs. 105, 106.)
A lengthening of the axis of the female strobiles of Coniferæ is not of
infrequent occurrence in Cryptomeria japonica, Larie europæa, &c., and
this is usually associated, as has been before stated, with a leaf-like
condition of the bracts, and sometimes even with the development of leaf-
bearing shoots in place of the scales. (See under Prolification of
Inflorescence and Phyllomorphy, and for references, p. 115.)
Elongation of the secondary flower-stalks.—In the previous section the
effect of elongation of the main rachis has been considered. A
corresponding deviation occurs in the peduncles or pedicels, and sometimes
alters the general character of the inflorescence very considerably,
converting a spike into a raceme, a raceme into a corymb, a capitulum into
an umbel, and so forth. A few such cases may here be alluded to. Fig. 206
represents a specimen of Ranunculus acris, in which the lower and lateral
flower-stalks were not only increased in number, but so much lengthened as
to form a flat-topped inflorescence—a corymbose cyme. In many
leguminous plants, as in Trifolium repens, Lotus corniculatus, &c., what is
Page 540
usually a compact spike, or head of flowers, becomes a raceme from the
elongation of the pedicels. In Umbelliferæ a similar change occurs, by
virtue of which sometimes the umbels themselves, and at other times the
florets, are raised on unusually long stalks, as in Angelica Razoulzii, Carum
Carui, Thysselinum palustre.[513] In Compositæ, when affected by an
analogous change, the capitulum assumes the appearance of a simple
umbel, as in Hypochæris radicata, Senecio vulgaris, and other plants.
Fig. 206.—Inflorescence of Ranunculus acris, with secondary peduncles lengthened.
In some of the double-flowered apples which have been previously alluded
to, the flower-stalk is inordinately long when compared with the adjacent
ones. Possibly in some of these cases the absence of the usual swelling of
the upper part of the peduncle may be connected with its increased length.
One of the most striking instances of lengthened flower-stalk occurred in an
elongation of the pedicels. In Umbelliferæ a similar change occurs, by
virtue of which sometimes the umbels themselves, and at other times the
florets, are raised on unusually long stalks, as in Angelica Razoulzii, Carum
Carui, Thysselinum palustre.[513] In Compositæ, when affected by an
analogous change, the capitulum assumes the appearance of a simple
umbel, as in Hypochæris radicata, Senecio vulgaris, and other plants.
Fig. 206.—Inflorescence of Ranunculus acris, with secondary peduncles lengthened.
In some of the double-flowered apples which have been previously alluded
to, the flower-stalk is inordinately long when compared with the adjacent
ones. Possibly in some of these cases the absence of the usual swelling of
the upper part of the peduncle may be connected with its increased length.
One of the most striking instances of lengthened flower-stalk occurred in an
Page 541
apple flower, wherein there was no swelling beneath the calyx, while the
latter was represented by five perfect stalked leaves.
Elongation of the leaves.—In the case of water plants this change keeps
pace with the corresponding growth of the stem, e.g. Ranunculus fluitans,
and in terrestrial plants there are varieties termed longifoliar, from the
unusual length of the leaves. A similar lengthening occurs in the involucral
leaves of Umbelliferæ and Compositæ, changing very materially the general
aspect of the inflorescence. Occasionally, also, the leaf-lobes of parsley
(Apium Petroselinum) and other crested-leaved plants may be observed to
lose their ordinary wavy form, and to be lengthened into flat riband-like
segments, as shown in fig. 207.
The only further illustrations that it is requisite to give of such changes in
this place are those occurring in lobed or compounded leaves, which, from
a lengthening of the midrib or central stalk, convert a digitate or palmate
leaf into a pinnate one. In these instances the lobes or leaflets become
separated one from another by a kind of apostasis. This change may be
frequently seen in the horse-chestnut, particularly in the young shoots
formed after the trees have been pruned or pollarded. In the adjoining cut
the intermediate stages between a palmate or digitate leaf to a pinnate one
may be seen. The specimens from which the drawing was made were taken
from the same tree at the same time.
latter was represented by five perfect stalked leaves.
Elongation of the leaves.—In the case of water plants this change keeps
pace with the corresponding growth of the stem, e.g. Ranunculus fluitans,
and in terrestrial plants there are varieties termed longifoliar, from the
unusual length of the leaves. A similar lengthening occurs in the involucral
leaves of Umbelliferæ and Compositæ, changing very materially the general
aspect of the inflorescence. Occasionally, also, the leaf-lobes of parsley
(Apium Petroselinum) and other crested-leaved plants may be observed to
lose their ordinary wavy form, and to be lengthened into flat riband-like
segments, as shown in fig. 207.
The only further illustrations that it is requisite to give of such changes in
this place are those occurring in lobed or compounded leaves, which, from
a lengthening of the midrib or central stalk, convert a digitate or palmate
leaf into a pinnate one. In these instances the lobes or leaflets become
separated one from another by a kind of apostasis. This change may be
frequently seen in the horse-chestnut, particularly in the young shoots
formed after the trees have been pruned or pollarded. In the adjoining cut
the intermediate stages between a palmate or digitate leaf to a pinnate one
may be seen. The specimens from which the drawing was made were taken
from the same tree at the same time.
Page 542
Fig. 207.—Portion of leaf of parsley, showing the change from short wavy to long flat leaf-lobes.
In the white clover, Trifolium repens, a similar transition may often be
observed, as also in some species of Potentilla.[514]
Elongation of the parts of the flower.—The only circumstance that needs
especial mention under this section is the great lengthening that sometimes
takes place in the carpels, sometimes as a result of injury from insects or
fungus, at other times without assignable cause.
In the white clover, Trifolium repens, a similar transition may often be
observed, as also in some species of Potentilla.[514]
Elongation of the parts of the flower.—The only circumstance that needs
especial mention under this section is the great lengthening that sometimes
takes place in the carpels, sometimes as a result of injury from insects or
fungus, at other times without assignable cause.
Page 543
Fig. 208.—Leaves of horse-chestnut, Æsculus, showing passage from digitate to pinnate leaves.
In the case of inferior ovaries this lengthening is, perhaps, even more
common, as in Umbelliferæ, Compositæ, &c. The common groundsel
(Senecio vulgaris) is especially liable to this form of enlargement of the
pistil, either in association with a leafy condition of the pappus or without
any such change.
Elongation of the thalamus, placenta, &c.—In some plants, as in
Magnolia or Myosurus, the thalamus becomes much elongated, and bears
the carpels disposed spirally around it. A similar lengthening occurs in
malformed flowers, usually in association with a similar change in the
lower or outer part of the flower, by virtue of which the whorls become
separated from each other (Apostasis). Elongation and protrusion of the
placenta have been already alluded to at p. 119, and also at p. 125. In some
In the case of inferior ovaries this lengthening is, perhaps, even more
common, as in Umbelliferæ, Compositæ, &c. The common groundsel
(Senecio vulgaris) is especially liable to this form of enlargement of the
pistil, either in association with a leafy condition of the pappus or without
any such change.
Elongation of the thalamus, placenta, &c.—In some plants, as in
Magnolia or Myosurus, the thalamus becomes much elongated, and bears
the carpels disposed spirally around it. A similar lengthening occurs in
malformed flowers, usually in association with a similar change in the
lower or outer part of the flower, by virtue of which the whorls become
separated from each other (Apostasis). Elongation and protrusion of the
placenta have been already alluded to at p. 119, and also at p. 125. In some
Page 544
of these cases the elongated placenta has taken the form of a leaf-bearing
shoot.[515]
Apostasis.—Engelmann made use of this term to express the separation of
parts one from another by the unusual elongation of the internodes.[516] He
drew a distinction between the separation of individual organs one from the
other, and the corresponding displacement of whorls. The subject has
already been, to a considerable degree, treated of in these pages under the
head of dialysis, displacement, and prolification, and but little need here be
added. With reference to the distance between one whorl and another, it will
be remembered that, although in the majority of cases the floral whorls are
packed closely together, yet in other instances the floral axis becomes
elongated, and thus separates the whorls one from another, by structures
such as the gynophores, androphores, &c., of Passifloreæ, Caryophylleæ,
Capparideæ, &c. &c.
A similar elongation of the thalamus, bringing about the separation of the
floral whorls, or of their constituent parts, is very commonly met with in
association with median prolification. Where the individual floral elements
are thus thrown out of their usual verticillate arrangement, they naturally
assume a spiral disposition, and are, in some cases, united by their margins,
so that a spiral sheet or tube is formed, surrounding the axis. This frequently
occurs in double flowers of the Chinese primrose, Primula sinensis.
Engelmann[517] figures a case wherein the calyx of Anagallis phœnicea was
separated by a rather long internode from the corolla, and a like illustration
in Torilis Anthriscus.
shoot.[515]
Apostasis.—Engelmann made use of this term to express the separation of
parts one from another by the unusual elongation of the internodes.[516] He
drew a distinction between the separation of individual organs one from the
other, and the corresponding displacement of whorls. The subject has
already been, to a considerable degree, treated of in these pages under the
head of dialysis, displacement, and prolification, and but little need here be
added. With reference to the distance between one whorl and another, it will
be remembered that, although in the majority of cases the floral whorls are
packed closely together, yet in other instances the floral axis becomes
elongated, and thus separates the whorls one from another, by structures
such as the gynophores, androphores, &c., of Passifloreæ, Caryophylleæ,
Capparideæ, &c. &c.
A similar elongation of the thalamus, bringing about the separation of the
floral whorls, or of their constituent parts, is very commonly met with in
association with median prolification. Where the individual floral elements
are thus thrown out of their usual verticillate arrangement, they naturally
assume a spiral disposition, and are, in some cases, united by their margins,
so that a spiral sheet or tube is formed, surrounding the axis. This frequently
occurs in double flowers of the Chinese primrose, Primula sinensis.
Engelmann[517] figures a case wherein the calyx of Anagallis phœnicea was
separated by a rather long internode from the corolla, and a like illustration
in Torilis Anthriscus.
Page 545
Fig. 209.—Flower of Delphinium, showing apostasis of carpels, from lengthening of the thalamus,
&c. (Cramer.)
A frequent change in Crucifers is due to the formation of a long stalk
bearing the pod, and thus giving rise to the appearance met as a constant
occurrence in Capparids.
In Tropæolum majus a similar elevation of the pistil may occasionally be
seen.
The adjacent figure of a monstrous Delphinium taken from Cramer
illustrates well the elongation of the floral axis and the apostasis of the
carpels. In this instance the axis is terminated by a second flower (median
prolification).
One of the best-marked illustrations of these changes occurs in a permanent
malformation of Epilobium hirsutum, specimens of which were originally
obtained from the late Professor Henslow. The several floral parts are here,
&c. (Cramer.)
A frequent change in Crucifers is due to the formation of a long stalk
bearing the pod, and thus giving rise to the appearance met as a constant
occurrence in Capparids.
In Tropæolum majus a similar elevation of the pistil may occasionally be
seen.
The adjacent figure of a monstrous Delphinium taken from Cramer
illustrates well the elongation of the floral axis and the apostasis of the
carpels. In this instance the axis is terminated by a second flower (median
prolification).
One of the best-marked illustrations of these changes occurs in a permanent
malformation of Epilobium hirsutum, specimens of which were originally
obtained from the late Professor Henslow. The several floral parts are here,
Page 546
some virescent, others truly foliaceous, and each whorl is separated from its
neighbour by a rather long internode. In Fuchsia and Campanula a like
change may occasionally be observed.
Engelmann, in addition to those previously mentioned, cites the following
plants as having manifested this change:
*Convallaria majalis!, *Tulipa Gesneriana!, Veronica Chamædrys,
Orobanche gracilis, Solanum Lycopersicum, Gentiana campestris,
Hypericum, Helleborus fetidus, Caltha palustris, Brassica oleracea! and
many Rosaceæ, Caryophylleæ, Cruciferæ, and Ranunculaceæ. (See
Dialysis, Median Prolification, &c.)
Apostasis of the sub-floral or involucral leaves is not of infrequent
occurrence in malformations affecting Compositæ and Umbelliferæ. In the
following genera it has been observed with especial frequency:—Torilis
Anthriscus, Eryngium, Athamanta Cervaria, Leontodon, Tragopogon
pratense!, Wedelia perfoliata! In garden anemones, also, it is a common
deviation.
FOOTNOTES:
[510] 'Cat. Plant.,' Lang., p. 113.
[511] 'Bull. Soc. Bot. France,' t. i, 1854, p. 173, and t. xiii, p. 96.
[512] 'Abhandl. Math. Phys. Class.,' Band. iv, Abhandl. i, tab. i.
[513] See Cramer, 'Bildungsabweich,' pp. 62–79, and Fleischer, 'Missbild, der
Culturpflanzen.'
[514] Schlechtendal, 'Bot. Zeit.,' 1844, p. 457; 'Linnæa,' xi, p. 301, xiv, p. 363; 'Bot. Zeit.,'
1856, p. 72; Masters, 'Rep. Brit. Assoc.,' Manchester, 1861; Coultas, 'What may be learnt from
a tree,' p. 118.
[515] For further details refer to the chapter on Displacements, p. 86.
neighbour by a rather long internode. In Fuchsia and Campanula a like
change may occasionally be observed.
Engelmann, in addition to those previously mentioned, cites the following
plants as having manifested this change:
*Convallaria majalis!, *Tulipa Gesneriana!, Veronica Chamædrys,
Orobanche gracilis, Solanum Lycopersicum, Gentiana campestris,
Hypericum, Helleborus fetidus, Caltha palustris, Brassica oleracea! and
many Rosaceæ, Caryophylleæ, Cruciferæ, and Ranunculaceæ. (See
Dialysis, Median Prolification, &c.)
Apostasis of the sub-floral or involucral leaves is not of infrequent
occurrence in malformations affecting Compositæ and Umbelliferæ. In the
following genera it has been observed with especial frequency:—Torilis
Anthriscus, Eryngium, Athamanta Cervaria, Leontodon, Tragopogon
pratense!, Wedelia perfoliata! In garden anemones, also, it is a common
deviation.
FOOTNOTES:
[510] 'Cat. Plant.,' Lang., p. 113.
[511] 'Bull. Soc. Bot. France,' t. i, 1854, p. 173, and t. xiii, p. 96.
[512] 'Abhandl. Math. Phys. Class.,' Band. iv, Abhandl. i, tab. i.
[513] See Cramer, 'Bildungsabweich,' pp. 62–79, and Fleischer, 'Missbild, der
Culturpflanzen.'
[514] Schlechtendal, 'Bot. Zeit.,' 1844, p. 457; 'Linnæa,' xi, p. 301, xiv, p. 363; 'Bot. Zeit.,'
1856, p. 72; Masters, 'Rep. Brit. Assoc.,' Manchester, 1861; Coultas, 'What may be learnt from
a tree,' p. 118.
[515] For further details refer to the chapter on Displacements, p. 86.
Page 547
[516] 'De Antholysi,' p. 42, § 49.
[517] Loc. cit., tab. 2, f. 6.
CHAPTER III.
ENATION.
Under the above heading are included certain forms arising from excess not
of growth, but of development, and consisting in the formation of
supplementary lobes or excrescences from various organs.
The new formations are not due either to a repetition or to a partition of any
organ, but are out-growths from others previously formed.
In prolification and in multiplication the adventitious structures are of
independent origin. In fission the new developments grow simultaneously
with the older ones, of which, indeed, they are mere repetitions. Moreover,
in fission the supplementary lobes do not, in general, project a plan different
from that of the original structure, at least in the first instance, though their
direction may ultimately become changed.
In enation the new growth projects from a previously formed organ after it
has attained to considerable size, or even after its ordinary proportions have
been attained, and it sprouts out from the beginning in a plane which is at a
considerable angle to that of the parent organ, and it is sometimes of a
different structure from it, and has different functions to fulfil.
Many of the instances that occur of scales projecting from petals, as in
Caryophylleæ, Sapindaceæ, &c., the coronal filaments of passion-flowers,
the cup of Narcissus, the appendages that beset the segments of the perianth
[517] Loc. cit., tab. 2, f. 6.
CHAPTER III.
ENATION.
Under the above heading are included certain forms arising from excess not
of growth, but of development, and consisting in the formation of
supplementary lobes or excrescences from various organs.
The new formations are not due either to a repetition or to a partition of any
organ, but are out-growths from others previously formed.
In prolification and in multiplication the adventitious structures are of
independent origin. In fission the new developments grow simultaneously
with the older ones, of which, indeed, they are mere repetitions. Moreover,
in fission the supplementary lobes do not, in general, project a plan different
from that of the original structure, at least in the first instance, though their
direction may ultimately become changed.
In enation the new growth projects from a previously formed organ after it
has attained to considerable size, or even after its ordinary proportions have
been attained, and it sprouts out from the beginning in a plane which is at a
considerable angle to that of the parent organ, and it is sometimes of a
different structure from it, and has different functions to fulfil.
Many of the instances that occur of scales projecting from petals, as in
Caryophylleæ, Sapindaceæ, &c., the coronal filaments of passion-flowers,
the cup of Narcissus, the appendages that beset the segments of the perianth
Page 548
in Lilium lancifolium, and other similar growths, may be referred to a like
process. In many cases this has been proved by a study of the development
of the flower, from which it appears that the growths in question are
developed subsequently to the formation of the ordinary floral whorls. It is
requisite, however, to be cautious in pronouncing upon the exact nature of
these bodies, in the absence of a knowledge of their period and mode of
formation. They may be mere outgrowths from one or other of the
customary whorls, or they may represent abortive stamens or petals, &c.
Where circumstances prevent the course of development from being traced,
something may be inferred as to their real nature from their position in
regard to the other parts of the flower, from their anatomical structure, and
from analogy or comparison with like organs in other plants. The period of
their formation is, perhaps, of less importance than was at one time
supposed, since it is well ascertained that, in some cases, the formation of
the parts of the flower, e.g. the stamens of mallows, follows a centrifugal
rather than a centripetal order.
In the case of monstrous developments of this nature too much care can
hardly be exercised, and the observer should rarely venture on an
explanation of the nature of the case from the evidence afforded by the
monstrous growth apart from that to be derived from the study of the
development and organization of the normal flower and from analogous
formations in allied plants.
Excrescences from axile organs—Warts.—In a preceding paragraph the
formation of gnaurs has been alluded to. There are other outgrowths, called
warts, occasionally met with in trees, and which are more closely connected
with the central tissues of the stem, while at the same time they are not
provided with buds, in which two particulars warts differ from gnaurs.
process. In many cases this has been proved by a study of the development
of the flower, from which it appears that the growths in question are
developed subsequently to the formation of the ordinary floral whorls. It is
requisite, however, to be cautious in pronouncing upon the exact nature of
these bodies, in the absence of a knowledge of their period and mode of
formation. They may be mere outgrowths from one or other of the
customary whorls, or they may represent abortive stamens or petals, &c.
Where circumstances prevent the course of development from being traced,
something may be inferred as to their real nature from their position in
regard to the other parts of the flower, from their anatomical structure, and
from analogy or comparison with like organs in other plants. The period of
their formation is, perhaps, of less importance than was at one time
supposed, since it is well ascertained that, in some cases, the formation of
the parts of the flower, e.g. the stamens of mallows, follows a centrifugal
rather than a centripetal order.
In the case of monstrous developments of this nature too much care can
hardly be exercised, and the observer should rarely venture on an
explanation of the nature of the case from the evidence afforded by the
monstrous growth apart from that to be derived from the study of the
development and organization of the normal flower and from analogous
formations in allied plants.
Excrescences from axile organs—Warts.—In a preceding paragraph the
formation of gnaurs has been alluded to. There are other outgrowths, called
warts, occasionally met with in trees, and which are more closely connected
with the central tissues of the stem, while at the same time they are not
provided with buds, in which two particulars warts differ from gnaurs.
Page 549
Excrescences of this kind often attain a very large size, and may be seen on
old elms and other trees, but, as their formation is probably more
pathological than teratological, no further notice of these structures need
here be given. No special notice need here be taken of the tubercles on the
roots of so many Leguminosæ, nor of the peculiar excrescences on the roots
of Taxodium distichum, as these appear to be normal formations. But it may
be well to mention in this place an anomalous development which occurs
occasionally in Ruscus aculeatus, and in which, from the upper surface of
the ordinary flattened leaf-like branch, projects at right angles a second
similar branch, so that in section the appearance would be like that of the
inverted letter t; thus, [symbol: Inverted upper-case T].
Enation from foliar organs—Leaves.—The development of adventitious
lobes from leaves may take place either from their surfaces or their margins.
A few illustrations may be given of each. In cabbage leaves a formation of
adventitious laminæ projecting at right angles from the primary one may
frequently be observed. In the instance figured (fig. 210) the new growths
proceeded almost exclusively from the thick midrib, which, in the figure, is
shown cut through just above the base. Not only is the ordinary semilunar
band of vascular tissue to be here seen, but a similar broken line of vessels
exists on the upper side of the leaf-stalk; thus the whole structure resembles
that of a stem or a branch as much as that of a true leaf.
old elms and other trees, but, as their formation is probably more
pathological than teratological, no further notice of these structures need
here be given. No special notice need here be taken of the tubercles on the
roots of so many Leguminosæ, nor of the peculiar excrescences on the roots
of Taxodium distichum, as these appear to be normal formations. But it may
be well to mention in this place an anomalous development which occurs
occasionally in Ruscus aculeatus, and in which, from the upper surface of
the ordinary flattened leaf-like branch, projects at right angles a second
similar branch, so that in section the appearance would be like that of the
inverted letter t; thus, [symbol: Inverted upper-case T].
Enation from foliar organs—Leaves.—The development of adventitious
lobes from leaves may take place either from their surfaces or their margins.
A few illustrations may be given of each. In cabbage leaves a formation of
adventitious laminæ projecting at right angles from the primary one may
frequently be observed. In the instance figured (fig. 210) the new growths
proceeded almost exclusively from the thick midrib, which, in the figure, is
shown cut through just above the base. Not only is the ordinary semilunar
band of vascular tissue to be here seen, but a similar broken line of vessels
exists on the upper side of the leaf-stalk; thus the whole structure resembles
that of a stem or a branch as much as that of a true leaf.
Page 550
Fig. 210.—Section through base of midrib of cabbage leaf, showing supplementary laminæ, &c.
The development of secondary leaves from the surfaces of primary ones
(phyllomania, autophyllogeny) has already been alluded to at p. 355.
Some of the cases wherein a leaf seems to have a double lamina may be
alluded to here, though possibly they would more properly be referred to
fission. The appearance presented is as if four wings projected from the
midrib, so that a cross section would be nearly in the form of [Symbol: )O(
turned 90 degrees.]. In an orange leaf presenting this appearance the lower
surface of one lamina was, as usual, dull in colour, while the upper surface
of the subjacent lamina was likewise dull; hence the impression might arise
that this was an instance of the adhesion of two leaves back to back, but the
petioles were not twisted, as they must have been had two leaves thus been
united, and neither in the petiole nor in the midrib was there the slightest
indication of fusion, the vascular bundles being arranged in a circular
manner, not in a horseshoe-like arrangement, as would have been the case
had adhesion taken place.[518] (See p. 33.)
The development of secondary leaves from the surfaces of primary ones
(phyllomania, autophyllogeny) has already been alluded to at p. 355.
Some of the cases wherein a leaf seems to have a double lamina may be
alluded to here, though possibly they would more properly be referred to
fission. The appearance presented is as if four wings projected from the
midrib, so that a cross section would be nearly in the form of [Symbol: )O(
turned 90 degrees.]. In an orange leaf presenting this appearance the lower
surface of one lamina was, as usual, dull in colour, while the upper surface
of the subjacent lamina was likewise dull; hence the impression might arise
that this was an instance of the adhesion of two leaves back to back, but the
petioles were not twisted, as they must have been had two leaves thus been
united, and neither in the petiole nor in the midrib was there the slightest
indication of fusion, the vascular bundles being arranged in a circular
manner, not in a horseshoe-like arrangement, as would have been the case
had adhesion taken place.[518] (See p. 33.)
Page 551
Such leaves as those of the hedgehog holly, Ilex Aquifolium, var. feroæ, and,
to a less extent, bullate leaves, may also be mentioned here as illustrations
of hypertrophy or enation.
Fig. 211.—Nephrodium molle. Ordinary frond and forked and crested varieties of the same, the crest
arising from the inordinate development of the margins of the pinnules.
When the increased development occurs at the margin of the leaves,
especially, the result is a wavy or crisped appearance, "folia undulata, vel
crispa."[519] These conditions occur normally in such leaves as those of
Rumex crispus, Malva crispa, &c., and are developed to an extreme degree
in garden varieties of parsley, some kails, &c., as well as in many ferns, but
these are probably cases rather of fission than enation as here understood.
[520]
Enation from the sepals.—The basal lobes of the calyx in Campanula
Medium, under normal circumstances, may be referred to in illustration of
to a less extent, bullate leaves, may also be mentioned here as illustrations
of hypertrophy or enation.
Fig. 211.—Nephrodium molle. Ordinary frond and forked and crested varieties of the same, the crest
arising from the inordinate development of the margins of the pinnules.
When the increased development occurs at the margin of the leaves,
especially, the result is a wavy or crisped appearance, "folia undulata, vel
crispa."[519] These conditions occur normally in such leaves as those of
Rumex crispus, Malva crispa, &c., and are developed to an extreme degree
in garden varieties of parsley, some kails, &c., as well as in many ferns, but
these are probably cases rather of fission than enation as here understood.
[520]
Enation from the sepals.—The basal lobes of the calyx in Campanula
Medium, under normal circumstances, may be referred to in illustration of
Page 552
this occurrence, while the adventitious spurs on the calyx of some
monstrous flowers seem due also to a like cause. These have already been
alluded to at p. 315.
Enation from the corolla.—The instances of this are more frequent than in
the case of the calyx, and admit of classification according as they occur in
polypetalous or gamopetalous flowers, on the outer or inner surface of the
petals, &c. Under natural circumstances the formation of scales, lobes, &c.,
from the petals, as in some Caryophylleæ, Sapindaceæ, &c. &c., may be
explained, as already remarked, by this process, rather than by fission,
chorisis, or by substitution of petals for stamens, &c. Each case must,
however, be examined on its own merits, as it is not safe to decide upon the
arrangement of parts in one flower by simply referring to the analogy of
others. In the following illustrations the course of development has not, in
all cases, been observed, and hence the explanation here given must be
taken with some reserve; for should it prove that the adventitious lobes,
&c., are formed simultaneously with the ordinary petals, the case will be
one of chorisis rather than of enation, as here understood. Again, it may be
that the supernumerary organs really represent petals or stamens in disguise,
though this hypothesis demands the further assumption (in order to account
for the interference with the law of alternation) that suppression of certain
organs has taken place.
Taking first those instances in which the supplementary petals appear on the
inner surface of the corolla, as being at once the most frequent, and as
presenting the closest analogy, with similar conformations, under natural
circumstances, certain double-flowered varieties of the Chinese primrose,
Primula sinensis, may be mentioned. In these flowers the calyx is normal,
the tube of the corolla is traversed by ten vascular bundles, and the limb is
divided into ten fimbriated lobes. About halfway up the tube, on the inner
surface, are given off five supernumerary petals, opposite to as many lobes
monstrous flowers seem due also to a like cause. These have already been
alluded to at p. 315.
Enation from the corolla.—The instances of this are more frequent than in
the case of the calyx, and admit of classification according as they occur in
polypetalous or gamopetalous flowers, on the outer or inner surface of the
petals, &c. Under natural circumstances the formation of scales, lobes, &c.,
from the petals, as in some Caryophylleæ, Sapindaceæ, &c. &c., may be
explained, as already remarked, by this process, rather than by fission,
chorisis, or by substitution of petals for stamens, &c. Each case must,
however, be examined on its own merits, as it is not safe to decide upon the
arrangement of parts in one flower by simply referring to the analogy of
others. In the following illustrations the course of development has not, in
all cases, been observed, and hence the explanation here given must be
taken with some reserve; for should it prove that the adventitious lobes,
&c., are formed simultaneously with the ordinary petals, the case will be
one of chorisis rather than of enation, as here understood. Again, it may be
that the supernumerary organs really represent petals or stamens in disguise,
though this hypothesis demands the further assumption (in order to account
for the interference with the law of alternation) that suppression of certain
organs has taken place.
Taking first those instances in which the supplementary petals appear on the
inner surface of the corolla, as being at once the most frequent, and as
presenting the closest analogy, with similar conformations, under natural
circumstances, certain double-flowered varieties of the Chinese primrose,
Primula sinensis, may be mentioned. In these flowers the calyx is normal,
the tube of the corolla is traversed by ten vascular bundles, and the limb is
divided into ten fimbriated lobes. About halfway up the tube, on the inner
surface, are given off five supernumerary petals, opposite to as many lobes
Page 553
of the corolla. Some of the supplementary petals have a stamen in front of
them, in the same relative position as in the normal flower. In some cases
the back or outer surface of the supplementary petal is turned towards the
inner or upper surface of the primary corolla, thus [Symbol: ((turned 90
degrees cw]; while, in other instances, the front of the adventitious lobe is
directed towards the corresponding surface of the original petal, thus
[Symbol: () turned 90 degrees]. Whether these supernumerary petals are
formed by chorisis or by enation cannot, with certainty, be determined
without examining the early stages of development.
Fig. 212.—Datura fastuosa. True corolla turned back to show the supernumerary corolla with the
petal-like segments attached to its outer surface (reduced).
Of more interest are those instances where the adventitious growth is on the
outside of the corolla; thus in a garden azalea there was intermediate
between the calyx and the corolla, both of which were normal, a series of
five petalodes, alternating with the sepals, and, therefore, opposite to the
them, in the same relative position as in the normal flower. In some cases
the back or outer surface of the supplementary petal is turned towards the
inner or upper surface of the primary corolla, thus [Symbol: ((turned 90
degrees cw]; while, in other instances, the front of the adventitious lobe is
directed towards the corresponding surface of the original petal, thus
[Symbol: () turned 90 degrees]. Whether these supernumerary petals are
formed by chorisis or by enation cannot, with certainty, be determined
without examining the early stages of development.
Fig. 212.—Datura fastuosa. True corolla turned back to show the supernumerary corolla with the
petal-like segments attached to its outer surface (reduced).
Of more interest are those instances where the adventitious growth is on the
outside of the corolla; thus in a garden azalea there was intermediate
between the calyx and the corolla, both of which were normal, a series of
five petalodes, alternating with the sepals, and, therefore, opposite to the
Page 554
lobes of the corolla, and adherent with them at the very base, though
elsewhere detached. These petalodes were concave on the surface looking
towards the calyx, and were there brightly coloured, while the tint of the
opposite surface looking towards the corolla was of a duller hue,
corresponding with that of the outside of the corolla-tube. This arrangement
of the colour was thus precisely similar to that which occurred in the four-
winged leaves already referred to at p. 446. In some flowers of Datura
fastuosa a similar series of excrescences was observed; the calyx and the
corolla were normal within the latter, intervening between it and the
stamens was a second corolla produced by duplication, and adherent to the
inner surface of this latter were five stamens. So far there was nothing very
peculiar; it remains to say, however, that on the outer surface of the second
corolla were five petal-like lobes closely adherent to it below, but partially
detached above. The colour of the adventitious segments was paler on the
outside than on the inner surface, as in the corolla itself. The position of the
several parts was such that they were opposite one to the other; hence,
while the lobes of the inner corolla were opposite to those of the outer one,
the intermediate petalodes were opposite to both; thus:
S S S S S
--------------------------
P P P P P
| X X X X X
|--------------------------
| P P P P P
|st st st st st
The X indicating the position of the petalodes.
elsewhere detached. These petalodes were concave on the surface looking
towards the calyx, and were there brightly coloured, while the tint of the
opposite surface looking towards the corolla was of a duller hue,
corresponding with that of the outside of the corolla-tube. This arrangement
of the colour was thus precisely similar to that which occurred in the four-
winged leaves already referred to at p. 446. In some flowers of Datura
fastuosa a similar series of excrescences was observed; the calyx and the
corolla were normal within the latter, intervening between it and the
stamens was a second corolla produced by duplication, and adherent to the
inner surface of this latter were five stamens. So far there was nothing very
peculiar; it remains to say, however, that on the outer surface of the second
corolla were five petal-like lobes closely adherent to it below, but partially
detached above. The colour of the adventitious segments was paler on the
outside than on the inner surface, as in the corolla itself. The position of the
several parts was such that they were opposite one to the other; hence,
while the lobes of the inner corolla were opposite to those of the outer one,
the intermediate petalodes were opposite to both; thus:
S S S S S
--------------------------
P P P P P
| X X X X X
|--------------------------
| P P P P P
|st st st st st
The X indicating the position of the petalodes.
Page 555
Fig. 213.—Gloxinia, with supernumerary segments on the outside of the true corolla.
A still more singular case is that of a variety of the Gloxinia, described
originally by Professor Edouard Morren,[521] but which is now becoming
common in English gardens. When first observed these flowers were
observed to produce petaloid segments outside the ordinary corolla, and
partially adherent to (or rather, not completely separated from it) much as in
the azalea before mentioned, the outer surface being brightly coloured, like
the inner surface of the corolla in ordinary gloxinias. Being encouraged and
tended by gardeners, in course of time, instead of a series of petalodes,
more or less distinct from one another, a second corolla or "catacorolla" was
formed outside the primary one, so that a hose in hose flower was produced,
but, in this case, the supplementary flower was formed on the outside and
not within the ordinary corolla. Moreover, the disposition of the colour was
reversed, for in the outermost corolla the richest hues were on the outer
surface, while in the inner or true corolla they were on the inside.
A still more singular case is that of a variety of the Gloxinia, described
originally by Professor Edouard Morren,[521] but which is now becoming
common in English gardens. When first observed these flowers were
observed to produce petaloid segments outside the ordinary corolla, and
partially adherent to (or rather, not completely separated from it) much as in
the azalea before mentioned, the outer surface being brightly coloured, like
the inner surface of the corolla in ordinary gloxinias. Being encouraged and
tended by gardeners, in course of time, instead of a series of petalodes,
more or less distinct from one another, a second corolla or "catacorolla" was
formed outside the primary one, so that a hose in hose flower was produced,
but, in this case, the supplementary flower was formed on the outside and
not within the ordinary corolla. Moreover, the disposition of the colour was
reversed, for in the outermost corolla the richest hues were on the outer
surface, while in the inner or true corolla they were on the inside.
Page 556
Professor Morren considers the adventitious petalodes as rudiments of so
many supplementary flowers, axillary to the calyx, and adnate to the
corolla; each lobe then would, in this view, represent an imperfect flower,
and the completed catacorolla would be formed of a series of confluent
flowers of this description. But this view involves the assumption of the
suppression of all the parts of the flower, except the lobes in question.
Fig. 214.—"Catacorolla" of Gloxinia, formed from the union of adventitious petalodes on the outside
of the true corolla (after Morren).
The view here propounded that the lobes in question are enations from the
true petals, which become confluent, so as to form the catacorolla, is surely
more simple, involves no assumptions of suppression of parts; and
moreover, is borne out by the examination of some flowers, where the
production of these adventitious lobes from the outside of the minute
partially developed petals could be distinctly seen.
many supplementary flowers, axillary to the calyx, and adnate to the
corolla; each lobe then would, in this view, represent an imperfect flower,
and the completed catacorolla would be formed of a series of confluent
flowers of this description. But this view involves the assumption of the
suppression of all the parts of the flower, except the lobes in question.
Fig. 214.—"Catacorolla" of Gloxinia, formed from the union of adventitious petalodes on the outside
of the true corolla (after Morren).
The view here propounded that the lobes in question are enations from the
true petals, which become confluent, so as to form the catacorolla, is surely
more simple, involves no assumptions of suppression of parts; and
moreover, is borne out by the examination of some flowers, where the
production of these adventitious lobes from the outside of the minute
partially developed petals could be distinctly seen.
Page 557
Enation from the stamens.—An illustration of this process occurred in
some double-flowered rhododendrons, which presented the following
arrangement of parts:—calyx and corolla normal; within the latter eight
petal-like stamens, forming a pseudo-corolla. The appearance presented by
the petaloid filaments and anthers was as if they were adnate to the centre
of the petals, but, on closer examination, it appeared that the petaloid
expansion to which the dilated filament was apparently attached, was
equally a part of the stamens; in other words, that the filament was provided
with four petal-like wings, two on each side [Symbol: 00 topped by (turned
cw 90°, an o above and another ( turned cw above that]. This disposition
was well seen in the anther, half of which was, in some cases, petaloid like
the filament; in fact, the inner wing of the latter was directly continuous
with the petal-like expansion from the anther. A section through the latter
showed, going from within outwards, the cut edges of two perfect
polliniferous lobes in the centre; and on either side the petaloid wing
representing the remaining anther-lobe; outside these were the edges of the
remaining wings, one on each side. (See p. 290, fig. 155.)
Enation from the carpels.—The only instances of this that need be
referred to are such cases as those in which spur-like projections, horns,
tubercles, or winged expansions, are formed from the surface of the ovary
during the course of its development. The extraordinary cornute oranges
described and figured by Ferrari, Gallesio, and other writers on the genus
Citrus, may be mentioned under this head. A similar formation occurs in the
fruit of some species of Solanum. (See p. 316.)
FOOTNOTES:
[518] It is desirable in this place to allude to a singular case of fissiparous division of a leaf
of Prunus Laurocerasus described by Prof. Alexander Dickson ('Seemann's Journ. Botany,'
vol. v, 1867, p. 323), and which did not come under the writer's notice till after the sheet
relating to fission, p. 61, had been sent to press. Dr. Dickson thus speaks of this abnormal leaf:
—"The petiole (unchanged) supported two laminæ, placed back to back, and united by their
some double-flowered rhododendrons, which presented the following
arrangement of parts:—calyx and corolla normal; within the latter eight
petal-like stamens, forming a pseudo-corolla. The appearance presented by
the petaloid filaments and anthers was as if they were adnate to the centre
of the petals, but, on closer examination, it appeared that the petaloid
expansion to which the dilated filament was apparently attached, was
equally a part of the stamens; in other words, that the filament was provided
with four petal-like wings, two on each side [Symbol: 00 topped by (turned
cw 90°, an o above and another ( turned cw above that]. This disposition
was well seen in the anther, half of which was, in some cases, petaloid like
the filament; in fact, the inner wing of the latter was directly continuous
with the petal-like expansion from the anther. A section through the latter
showed, going from within outwards, the cut edges of two perfect
polliniferous lobes in the centre; and on either side the petaloid wing
representing the remaining anther-lobe; outside these were the edges of the
remaining wings, one on each side. (See p. 290, fig. 155.)
Enation from the carpels.—The only instances of this that need be
referred to are such cases as those in which spur-like projections, horns,
tubercles, or winged expansions, are formed from the surface of the ovary
during the course of its development. The extraordinary cornute oranges
described and figured by Ferrari, Gallesio, and other writers on the genus
Citrus, may be mentioned under this head. A similar formation occurs in the
fruit of some species of Solanum. (See p. 316.)
FOOTNOTES:
[518] It is desirable in this place to allude to a singular case of fissiparous division of a leaf
of Prunus Laurocerasus described by Prof. Alexander Dickson ('Seemann's Journ. Botany,'
vol. v, 1867, p. 323), and which did not come under the writer's notice till after the sheet
relating to fission, p. 61, had been sent to press. Dr. Dickson thus speaks of this abnormal leaf:
—"The petiole (unchanged) supported two laminæ, placed back to back, and united by their
Page 558
midribs (i.e. not separated) to within about an inch from their extremities, which were
perfectly free from each other. These laminæ stood vertically, their edges being directed
towards and away from the axis; and as they were placed back to back, the shining surfaces,
corresponding in structure to the normal upper leaf-surface, were directed laterally outwards.
In the axil of this abnormal leaf were two axillary buds. The existence of two leaf-apices and
two axillary buds shows that this was not due to an accidental exuberance of development, but
to fissiparous division, which, had it been complete, would have resulted in the replacement of
a single leaf by two leaves. The arrangement in Prof. Dickson's leaf may be thus represented:
[symbol:)OO(with X above]. The nature of the case may be even better seen by comparison
with the normal arrangement, which would be [symbol: (OX turned 90 degrees ccw], while in
those cases where the fission of the leaf occurs in the same plane as that of the primary lamina,
as where a leaf splits into two lobes at the apex, with a midrib to each, the arrangement is as
follows: [symbol: OX turned 90 degrees ccw, with 2 arcs below], the X in all cases
representing the position of the axis, the O that of the axillary bud, and the [symbol: (turned 90
degrees ccw] that of the laminæ."
[519] Linn., 'Phil. Bot.,' § 274. The term "crispa" is surely preferable to that of Ré,
"phyllorhyseme."
[520] See C. Morren, "Consid. sur les déformations," &c., in 'Bull. Acad. Belg.,' 1852, tom,
xix, part 3, p. 444; and as to ferns, see Moore, 'Nature-Printed British Ferns,' 8vo ed., where
numerous illustrations are given.
[521] 'Bull. Acad. Belg.,' t. xix, p. 224, tab. i; and 'Gardeners' Chronicle,' 1865, p. 865.
PART II.
ATROPHY.
Under the head of atrophy are included those cases wherein the organs
affected are actually present, but in a dwarfed and stunted condition as
compared with surrounding parts.
The diminished size is, in such instances, obviously due to a partial
development and to an arrest of growth at a certain stage, from the
perfectly free from each other. These laminæ stood vertically, their edges being directed
towards and away from the axis; and as they were placed back to back, the shining surfaces,
corresponding in structure to the normal upper leaf-surface, were directed laterally outwards.
In the axil of this abnormal leaf were two axillary buds. The existence of two leaf-apices and
two axillary buds shows that this was not due to an accidental exuberance of development, but
to fissiparous division, which, had it been complete, would have resulted in the replacement of
a single leaf by two leaves. The arrangement in Prof. Dickson's leaf may be thus represented:
[symbol:)OO(with X above]. The nature of the case may be even better seen by comparison
with the normal arrangement, which would be [symbol: (OX turned 90 degrees ccw], while in
those cases where the fission of the leaf occurs in the same plane as that of the primary lamina,
as where a leaf splits into two lobes at the apex, with a midrib to each, the arrangement is as
follows: [symbol: OX turned 90 degrees ccw, with 2 arcs below], the X in all cases
representing the position of the axis, the O that of the axillary bud, and the [symbol: (turned 90
degrees ccw] that of the laminæ."
[519] Linn., 'Phil. Bot.,' § 274. The term "crispa" is surely preferable to that of Ré,
"phyllorhyseme."
[520] See C. Morren, "Consid. sur les déformations," &c., in 'Bull. Acad. Belg.,' 1852, tom,
xix, part 3, p. 444; and as to ferns, see Moore, 'Nature-Printed British Ferns,' 8vo ed., where
numerous illustrations are given.
[521] 'Bull. Acad. Belg.,' t. xix, p. 224, tab. i; and 'Gardeners' Chronicle,' 1865, p. 865.
PART II.
ATROPHY.
Under the head of atrophy are included those cases wherein the organs
affected are actually present, but in a dwarfed and stunted condition as
compared with surrounding parts.
The diminished size is, in such instances, obviously due to a partial
development and to an arrest of growth at a certain stage, from the
Page 559
operation of various causes, either external or inherent to the organization
itself. It may affect any part of the plant, and exists, in very varying degree,
in different instances, being sometimes so slight in amount as not to
preclude the exercise of the functions of the part; while in others, the
structure is so incomplete that the office cannot be performed. These
differences depend, of course, upon the stage of development which the
organ had reached when its growth was checked. For practical purposes
atrophy may be distinguished from suppression by the fact that in the latter
case a certain element of the flower or plant which, under ordinary
circumstances, is present, is entirely wanting, while, in the former class, it
exists but in a rudimentary condition.
Again, atrophy is to be separated from that general diminution in the size of
the whole plant or of distinct parts of that plant which is comprised under
the term "nanism." Thus the several dwarf varieties of plants (var. nanæ), or
those in which the leaves or flowers are smaller than usual (var. parvifoliæ,
v. parvifloræ), are truly regarded as variations, and not as malformations
properly so called.
Atrophy is partial and special in its operation, nanism is general.
Under ordinary circumstances atrophy is exemplified by the presence of
rudimentary or imperfect organs, as, for instance, in Pentstemon,
Scrophularia, &c., where one stamen is atrophied.
For convenience sake atrophy may be divided into abortion and
degeneration, the first including cases where, from arrest of development
occurring at an early stage, organs are present; but in a much smaller and
more rudimentary condition than usual, their form and general appearance,
except so far as regards their dimensions, not being materially altered. On
the other hand, in cases of degeneration, development is not entirely
itself. It may affect any part of the plant, and exists, in very varying degree,
in different instances, being sometimes so slight in amount as not to
preclude the exercise of the functions of the part; while in others, the
structure is so incomplete that the office cannot be performed. These
differences depend, of course, upon the stage of development which the
organ had reached when its growth was checked. For practical purposes
atrophy may be distinguished from suppression by the fact that in the latter
case a certain element of the flower or plant which, under ordinary
circumstances, is present, is entirely wanting, while, in the former class, it
exists but in a rudimentary condition.
Again, atrophy is to be separated from that general diminution in the size of
the whole plant or of distinct parts of that plant which is comprised under
the term "nanism." Thus the several dwarf varieties of plants (var. nanæ), or
those in which the leaves or flowers are smaller than usual (var. parvifoliæ,
v. parvifloræ), are truly regarded as variations, and not as malformations
properly so called.
Atrophy is partial and special in its operation, nanism is general.
Under ordinary circumstances atrophy is exemplified by the presence of
rudimentary or imperfect organs, as, for instance, in Pentstemon,
Scrophularia, &c., where one stamen is atrophied.
For convenience sake atrophy may be divided into abortion and
degeneration, the first including cases where, from arrest of development
occurring at an early stage, organs are present; but in a much smaller and
more rudimentary condition than usual, their form and general appearance,
except so far as regards their dimensions, not being materially altered. On
the other hand, in cases of degeneration, development is not entirely
Page 560
checked, but rather perverted, so that not only the dimensions are lessened,
but the form is altered.
CHAPTER I.
ABORTION.
The sense in which this term is here understood has been explained in the
preceding paragraph. It is only necessary to say further, that cases of
abortion are to be distinguished from those of suppression, on the one hand,
and those of degeneration on the other. In suppression there is from the first
an absolute deficiency of a particular organ. In degeneration the part is
present, but in a diminished and perverted condition. In abortion it exists,
but in a stunted and dwarfed, but not otherwise permuted state.
Abortion of axile organs.—When the main stem is arrested in its growth,
the habit and general appearance of the plant are materially altered, as in the
so-called stemless plants, plantæ acaules. In these the internodes are so
slightly developed that the leaves are closely crowded in tufts or rosettes.
When this shortening of the stem (acaulosia) occurs, without other
considerable change in other organs, the deviation is classed under the head
of variation rather than of monstrosity; and, indeed, in very many plants,
this arrested growth of the axis is the rule rather than the exception. When
occurring in an abnormal manner, atrophy of the stem is most frequently
attended by other more or less grave alterations in other structures; thus
Moquin-Tandon[522] cites an instance of Camphorosma monspeliaca,
wherein the stems presented the form of very short, hard, woody tubercles,
thickly clothed with deformed leaves, and invested by a vast number of
hairs, longer and more dense than usual. A similar deformity sometimes
occurs in an Indian species of Artabotrys; in these specimens the branchlets
but the form is altered.
CHAPTER I.
ABORTION.
The sense in which this term is here understood has been explained in the
preceding paragraph. It is only necessary to say further, that cases of
abortion are to be distinguished from those of suppression, on the one hand,
and those of degeneration on the other. In suppression there is from the first
an absolute deficiency of a particular organ. In degeneration the part is
present, but in a diminished and perverted condition. In abortion it exists,
but in a stunted and dwarfed, but not otherwise permuted state.
Abortion of axile organs.—When the main stem is arrested in its growth,
the habit and general appearance of the plant are materially altered, as in the
so-called stemless plants, plantæ acaules. In these the internodes are so
slightly developed that the leaves are closely crowded in tufts or rosettes.
When this shortening of the stem (acaulosia) occurs, without other
considerable change in other organs, the deviation is classed under the head
of variation rather than of monstrosity; and, indeed, in very many plants,
this arrested growth of the axis is the rule rather than the exception. When
occurring in an abnormal manner, atrophy of the stem is most frequently
attended by other more or less grave alterations in other structures; thus
Moquin-Tandon[522] cites an instance of Camphorosma monspeliaca,
wherein the stems presented the form of very short, hard, woody tubercles,
thickly clothed with deformed leaves, and invested by a vast number of
hairs, longer and more dense than usual. A similar deformity sometimes
occurs in an Indian species of Artabotrys; in these specimens the branchlets
Page 561
are contracted in length, and bear numerous closely packed scaly leaves,
densely hairy, and much smaller than ordinary.
Spines and thorns may he looked on as atrophied branches, and seem to
result from poorness of soil, as the same plants, which, in hungry land,
produce spines, develop their branches to the full extent when grown under
more favorable conditions.[523]
In the birch an arrest of development in some of the branches is of common
occurrence. The branch suddenly ceases to grow in length; at the same time
it thickens at the end into a large bulbous knob, from which are developed a
profusion of small twigs, whose direction is sometimes exactly the reverse
of that of the main branch. (See p. 347.)
The branches of the common spruce fir, especially the lateral ones, when
attacked by a particular species of aphis, are very apt to be developed into a
cone-like excrescence.[524]
A shortened condition of the flower-stalks occurs occasionally, greatly
altering the general character of the inflorescence. This has been observed
in pelargoniums and in the Chinese primrose, in both of which the effect
was to replace the umbellate form of inflorescence by a capitate one.
Abortion of the receptacle.—Here may be mentioned those cases of
flowers with habitually inferior ovary (real or apparent), in which the
receptacle fails, from some cause or other, to dilate as usual. This has
already been alluded to under the head of Prolification, Displacements, &c.
(pp. 78, 130, &c., figs. 35–37, 64, &c.), and hence requires only incidental
comment in this place. There are, however, certain other cases of a similar
nature which may here be referred to; such as the abortive condition of the
inferior ovary, or rather of the receptacle, that usually encircles the ovary in
Compositæ and Umbelliferæ. In the former natural order the following
densely hairy, and much smaller than ordinary.
Spines and thorns may he looked on as atrophied branches, and seem to
result from poorness of soil, as the same plants, which, in hungry land,
produce spines, develop their branches to the full extent when grown under
more favorable conditions.[523]
In the birch an arrest of development in some of the branches is of common
occurrence. The branch suddenly ceases to grow in length; at the same time
it thickens at the end into a large bulbous knob, from which are developed a
profusion of small twigs, whose direction is sometimes exactly the reverse
of that of the main branch. (See p. 347.)
The branches of the common spruce fir, especially the lateral ones, when
attacked by a particular species of aphis, are very apt to be developed into a
cone-like excrescence.[524]
A shortened condition of the flower-stalks occurs occasionally, greatly
altering the general character of the inflorescence. This has been observed
in pelargoniums and in the Chinese primrose, in both of which the effect
was to replace the umbellate form of inflorescence by a capitate one.
Abortion of the receptacle.—Here may be mentioned those cases of
flowers with habitually inferior ovary (real or apparent), in which the
receptacle fails, from some cause or other, to dilate as usual. This has
already been alluded to under the head of Prolification, Displacements, &c.
(pp. 78, 130, &c., figs. 35–37, 64, &c.), and hence requires only incidental
comment in this place. There are, however, certain other cases of a similar
nature which may here be referred to; such as the abortive condition of the
inferior ovary, or rather of the receptacle, that usually encircles the ovary in
Compositæ and Umbelliferæ. In the former natural order the following
Page 562
plants have been met with in this condition:—*Tragopogon pratense!,
*Cirsium arvense, Hypochæris radicata, Senecio vulgaris!, Coreopsis
Drummondi. In the latter order, Daucus Carota! Œnanthe crocata! and
Thysselinum palustre, seem most frequently to have been observed in this
state.[525] In some gourds the receptacle may be seen partially developed
only, and forming a kind of cup, from which the true carpels protrude.
Abortion of the leaves.—Arrest of growth in the leaves occurs in different
ways; sometimes the whole leaf is smaller than usual; at other times certain
parts only are reduced in size; while, in a third class of cases, portions of the
leaf are entirely suppressed.
Moquin[526] mentions having seen the leaves of Chenopodium vulvaria, and
of Diplotaxis muralis reduced to a fourth of their natural size; and he
alludes to other cases of the same nature, seen by other observers, in
Hypericum perforatum and Blitum polymorphum.
Nicandra physaloides[527] has also been met with in a similar condition,
which, indeed, is a common result of insect-puncture, and of fungous
growth in plants. Those instances in which the leaf is diminished in size,
without any attendant malformation in other organs, may be regarded rather
as variations than as monstrosities, as in the case of the entire-leaved
varieties of those plants which ordinarily have cut or divided leaves, e.g.
Plantago Coronopus, var. integrifolia, Papaver Rhœas integrifolia, &c. &c.
The same remark may be made of those specimens in which one part of the
leaf is developed to a less extent than another, as happens in the submerged
leaves of such plants as Ranunculus aquatilis, Cabomba aquatica, the
spiney leaves of Berberis, the fenestrated leaves of Ouvirandra, &c. In the
illustrations last cited the relative deficiency of one portion, as contrasted
with another, takes place as a constant occurrence, and is uniform and
regular throughout the whole leaf. When, on the other hand, the deficiency
*Cirsium arvense, Hypochæris radicata, Senecio vulgaris!, Coreopsis
Drummondi. In the latter order, Daucus Carota! Œnanthe crocata! and
Thysselinum palustre, seem most frequently to have been observed in this
state.[525] In some gourds the receptacle may be seen partially developed
only, and forming a kind of cup, from which the true carpels protrude.
Abortion of the leaves.—Arrest of growth in the leaves occurs in different
ways; sometimes the whole leaf is smaller than usual; at other times certain
parts only are reduced in size; while, in a third class of cases, portions of the
leaf are entirely suppressed.
Moquin[526] mentions having seen the leaves of Chenopodium vulvaria, and
of Diplotaxis muralis reduced to a fourth of their natural size; and he
alludes to other cases of the same nature, seen by other observers, in
Hypericum perforatum and Blitum polymorphum.
Nicandra physaloides[527] has also been met with in a similar condition,
which, indeed, is a common result of insect-puncture, and of fungous
growth in plants. Those instances in which the leaf is diminished in size,
without any attendant malformation in other organs, may be regarded rather
as variations than as monstrosities, as in the case of the entire-leaved
varieties of those plants which ordinarily have cut or divided leaves, e.g.
Plantago Coronopus, var. integrifolia, Papaver Rhœas integrifolia, &c. &c.
The same remark may be made of those specimens in which one part of the
leaf is developed to a less extent than another, as happens in the submerged
leaves of such plants as Ranunculus aquatilis, Cabomba aquatica, the
spiney leaves of Berberis, the fenestrated leaves of Ouvirandra, &c. In the
illustrations last cited the relative deficiency of one portion, as contrasted
with another, takes place as a constant occurrence, and is uniform and
regular throughout the whole leaf. When, on the other hand, the deficiency
Page 563
in question happens accidentally and irregularly, the change may be
considered as a malformation. One side of the blade of the leaf is frequently
affected in this manner, the other portions remaining unaffected. It would
appear as if any plant might be thus altered, but the following species
appear to be particularly subject to this change: Æesculus Hippocastanum,
Digitalis purpurea, Morus alba, Fagus silvatica contracta (hort.), Codiæum
variegatum var. erosum (hort.), Broussonettia papyrifera, Scolopendrium
vulgare, &c.
Frequently this irregular diminution in proportion is coexistent with an
unusual degree of cleavage or laciniation of the margin, as in Acer
platanoides laciniatum, Tilia asplenifolia, Alnus imperialis (hort.), Fagus
silvatica var. (hort.), &c.
In the case of what are sometimes termed interrupted leaves, the laminar
portions of the leaf are here and there deficient on both sides of the midrib,
leaving small portions of the latter, as it were, denuded and connecting the
segments of the laminæ one with the other. This has been observed amongst
other plants in Veronica latifolia, Broussonettia papyrifer, Codiæum
variegatum var. interruptum (hort.), Scolopendrium vulgare, &c.[528] (See p.
328.)
In some of the leaves which have been already referred to in illustration of
the inordinate growth of the cellular portions, the increased development of
parenchyma is associated with a contracted state of the midrib and its
branches, producing a puckered appearance of the leaf, an exaggerated
degree of that change which produces what are termed "folia bullata." In
illustration may be cited various species of Mentha, Perilla, Coleus, Fagus
silvatica crispa, Cytisus, Laburnum var., and other forms, cultivated in
gardens for their singularity.
considered as a malformation. One side of the blade of the leaf is frequently
affected in this manner, the other portions remaining unaffected. It would
appear as if any plant might be thus altered, but the following species
appear to be particularly subject to this change: Æesculus Hippocastanum,
Digitalis purpurea, Morus alba, Fagus silvatica contracta (hort.), Codiæum
variegatum var. erosum (hort.), Broussonettia papyrifera, Scolopendrium
vulgare, &c.
Frequently this irregular diminution in proportion is coexistent with an
unusual degree of cleavage or laciniation of the margin, as in Acer
platanoides laciniatum, Tilia asplenifolia, Alnus imperialis (hort.), Fagus
silvatica var. (hort.), &c.
In the case of what are sometimes termed interrupted leaves, the laminar
portions of the leaf are here and there deficient on both sides of the midrib,
leaving small portions of the latter, as it were, denuded and connecting the
segments of the laminæ one with the other. This has been observed amongst
other plants in Veronica latifolia, Broussonettia papyrifer, Codiæum
variegatum var. interruptum (hort.), Scolopendrium vulgare, &c.[528] (See p.
328.)
In some of the leaves which have been already referred to in illustration of
the inordinate growth of the cellular portions, the increased development of
parenchyma is associated with a contracted state of the midrib and its
branches, producing a puckered appearance of the leaf, an exaggerated
degree of that change which produces what are termed "folia bullata." In
illustration may be cited various species of Mentha, Perilla, Coleus, Fagus
silvatica crispa, Cytisus, Laburnum var., and other forms, cultivated in
gardens for their singularity.
Page 564
Entire absence of the stalk of the leaf occurs normally in sessile leaves; on
the other hand the blade of the leaf is only occasionally developed in the
phyllodineous Acacias, in some species of Oxalis, Indigofera, Lebeckia,
Ranunculus, Bupleurum, &c.
De Candolle,[529] from a consideration of Strelitzia juncea, in which the
petiole alone is developed, was led to the inference that in many
monocotyledonous plants the blade of the leaf was never developed, the
portion present being the sheath or stalk, unprovided with limb. The
correctness of this inference is shown, amongst other things, by the
occasional presence of a leaf-blade in Strelitzia juncea itself.
Occasionally the laminar portions of the leaf are completely wanting,
leaving only the main ribs, as in the case of Berberis, while the adjoining
figure (fig. 215) represents an instance of a cabbage wherein the innermost
leaves are represented by thick fleshy cylindrical bodies corresponding to
the midribs of the ordinary leaves. There is in cultivation a variety of the
cabbage which constantly presents this peculiarity.
the other hand the blade of the leaf is only occasionally developed in the
phyllodineous Acacias, in some species of Oxalis, Indigofera, Lebeckia,
Ranunculus, Bupleurum, &c.
De Candolle,[529] from a consideration of Strelitzia juncea, in which the
petiole alone is developed, was led to the inference that in many
monocotyledonous plants the blade of the leaf was never developed, the
portion present being the sheath or stalk, unprovided with limb. The
correctness of this inference is shown, amongst other things, by the
occasional presence of a leaf-blade in Strelitzia juncea itself.
Occasionally the laminar portions of the leaf are completely wanting,
leaving only the main ribs, as in the case of Berberis, while the adjoining
figure (fig. 215) represents an instance of a cabbage wherein the innermost
leaves are represented by thick fleshy cylindrical bodies corresponding to
the midribs of the ordinary leaves. There is in cultivation a variety of the
cabbage which constantly presents this peculiarity.
Page 565
Fig. 215.—Inner leaves of cabbage reduced to their midribs.
The suppression of one or more leaflets of a compound leaf has already
been referred to at p. 396.
Abortion of the perianth, calyx, and corolla.—Illustrations of partial
development in these organs are not rare, under ordinary circumstances, as
for instance the "obsolete" calyx of Umbellifers. In the cauliflower the
branches of the inflorescence are contracted in length, while their
succulence is much increased; at their extremities they bear crowds of
imperfect flowers, in which the calyx only is visible, and that only in a
rudimentary and partially developed condition. Imperfect development of
the whole or of some of the constituent parts is more common in the case of
the corolla than in that of the calyx. In Arenaria serpyllifolia the petals,
especially in autumn, are only one fourth the length of the sepals. Anagallis
phœnicea, Honckenya peploides, Arabis alpina, Ranunculus auricomus,
The suppression of one or more leaflets of a compound leaf has already
been referred to at p. 396.
Abortion of the perianth, calyx, and corolla.—Illustrations of partial
development in these organs are not rare, under ordinary circumstances, as
for instance the "obsolete" calyx of Umbellifers. In the cauliflower the
branches of the inflorescence are contracted in length, while their
succulence is much increased; at their extremities they bear crowds of
imperfect flowers, in which the calyx only is visible, and that only in a
rudimentary and partially developed condition. Imperfect development of
the whole or of some of the constituent parts is more common in the case of
the corolla than in that of the calyx. In Arenaria serpyllifolia the petals,
especially in autumn, are only one fourth the length of the sepals. Anagallis
phœnicea, Honckenya peploides, Arabis alpina, Ranunculus auricomus,
Page 566
Rubus fruticosus, and Geranium columbinum, also frequently afford
illustrations of this circumstance.
Fig. 216.—Abortion of four out of five petals, Viola tricolor, side and front views.
At fig. 216 is represented a pansy in which four of the five petals were very
small and colourless, while the lower spurred petal was of the usual size
and colour. In this flower the stamens and pistils were wholly suppressed,
and the flower-stalk, instead of being bent near the flower, retained its
primary straight direction. Similar atrophic conditions of the corolla occur
habitually among Violaceæ.
The diminished size of the petals sometimes coexists with an increase in
their number, as in a flower of Streptocarpus Rexii, mentioned by Bureau.
[530]
illustrations of this circumstance.
Fig. 216.—Abortion of four out of five petals, Viola tricolor, side and front views.
At fig. 216 is represented a pansy in which four of the five petals were very
small and colourless, while the lower spurred petal was of the usual size
and colour. In this flower the stamens and pistils were wholly suppressed,
and the flower-stalk, instead of being bent near the flower, retained its
primary straight direction. Similar atrophic conditions of the corolla occur
habitually among Violaceæ.
The diminished size of the petals sometimes coexists with an increase in
their number, as in a flower of Streptocarpus Rexii, mentioned by Bureau.
[530]
Page 567
Among monocotyledons this partial development seems to be even more
frequent than in dicotyledons. In addition to the well-known cases of certain
species of Bellevalia and Muscari, wherein the uppermost flowers of the
raceme are more or less atrophied (see p. 347, fig. 179), a few less common
illustrations may be cited. In crocuses it is not a very uncommon
circumstance to find the three inner segments of the perianth smaller than
natural, and generally unequal in size. This occurs without any other
perceptible change in the flower.
Schlechtendal[531] mentions a flower of Fritillaria imperialis in which the
perianthial leaves were relatively very small, and destitute of the usual
nectary, while the stamens, on the other hand, were of their natural size and
appearance. Fresenius[532] records a similar occurrence in the same plant.
Morren[533] gives details of like appearances in Hymenocallis americana,
and Delavaud[534] in Tigridia pavonia.
In certain orchids an arrested development of the perianth is habitual, as in
Oncidium abortivum (fig. 217), where, on a large branching panicle,
numerous abortive, but few perfect, flowers are produced. In a similar way
the petals and labellum of Odontoglossum Uro-Skinneri have been found
reduced to filamentous processes.
frequent than in dicotyledons. In addition to the well-known cases of certain
species of Bellevalia and Muscari, wherein the uppermost flowers of the
raceme are more or less atrophied (see p. 347, fig. 179), a few less common
illustrations may be cited. In crocuses it is not a very uncommon
circumstance to find the three inner segments of the perianth smaller than
natural, and generally unequal in size. This occurs without any other
perceptible change in the flower.
Schlechtendal[531] mentions a flower of Fritillaria imperialis in which the
perianthial leaves were relatively very small, and destitute of the usual
nectary, while the stamens, on the other hand, were of their natural size and
appearance. Fresenius[532] records a similar occurrence in the same plant.
Morren[533] gives details of like appearances in Hymenocallis americana,
and Delavaud[534] in Tigridia pavonia.
In certain orchids an arrested development of the perianth is habitual, as in
Oncidium abortivum (fig. 217), where, on a large branching panicle,
numerous abortive, but few perfect, flowers are produced. In a similar way
the petals and labellum of Odontoglossum Uro-Skinneri have been found
reduced to filamentous processes.
Page 568
Fig. 217.—Flower of Oncidium abortivum, magnified.
Abortion of the stamens.—Atrophy of one or more stamens is of very
common occurrence, as a general rule, in many genera of plants, e.g.
Scrophularia, Erodium, many Restiaceæ, &c. &c. As a strictly teratological
condition atrophy of the stamens is more rare than complete suppression. It
has been noticed in Arabis alpina, Cerastium glomeratum, C. tetrandrum,
Rhamnus catharticus, Anemone, Hepatica, &c. It happens frequently
among Orchids both wild and cultivated. In the Hymenocallis flowers
described by the elder Morren, four out of five stamens were atrophied. In
other flowers, otherwise perfectly formed, one abortive stamen was found
bearing a spherical indehiscent anther. All these atrophied anthers of
Hymenocallis were found to contain pollen, differing at first sight but little
from what is usual, but presenting this important peculiarity, that while the
normal pollen does not burst until it comes into contact with the stigma, in
the abnormal flowers the outer coat of the pollen-grains split while still
Abortion of the stamens.—Atrophy of one or more stamens is of very
common occurrence, as a general rule, in many genera of plants, e.g.
Scrophularia, Erodium, many Restiaceæ, &c. &c. As a strictly teratological
condition atrophy of the stamens is more rare than complete suppression. It
has been noticed in Arabis alpina, Cerastium glomeratum, C. tetrandrum,
Rhamnus catharticus, Anemone, Hepatica, &c. It happens frequently
among Orchids both wild and cultivated. In the Hymenocallis flowers
described by the elder Morren, four out of five stamens were atrophied. In
other flowers, otherwise perfectly formed, one abortive stamen was found
bearing a spherical indehiscent anther. All these atrophied anthers of
Hymenocallis were found to contain pollen, differing at first sight but little
from what is usual, but presenting this important peculiarity, that while the
normal pollen does not burst until it comes into contact with the stigma, in
the abnormal flowers the outer coat of the pollen-grains split while still
Page 569
within the anther, from which latter, indeed, they could not escape, owing to
the indehiscent nature of the latter. Again, the pollen-tube of the abnormal
grains cracked, in its turn, on mere exposure to the air, and liberated the
fovilla, so that the pollen of these atrophied anthers was necessarily
impotent, because it opened before it could be applied to the stigma, even
had that been rendered possible by the opening of the anther.
An abortive condition of the stamens and of the pollen, is of very common
occurrence among hybridised plants. Gaertner and other writers have
spoken of this defective condition as contabescence.[535] It forms one reason
for the sterility so frequently observed in the case of true hybrids. In some
hybrid passion-flowers, while all other parts of the flower were apparently
perfect, even to the ovules, the stamens were atrophied, and distorted, and
contained little or no pollen; the few grains of the latter being smaller than
usual. (See under Heterogamy, pp. 193–196, and p. 398.)
Abortion of the pistil, fruit, &c.—Traces of the carpels occur in many
male flowers of unisexual plants, e.g. Sterculiaceæ, Euphorbiaceæ,
Restiaceæ, &c. &c., and in some natural orders there appears to be a
tendency towards a diœcious condition, e.g. Caryophylleæ, as in Lychnis
dioica, Silene otites, Arenaria tetraquetra, &c. The last-named plant is
stated to have, in some cases, imperfect pistils; in others, rudimentary
stamens; while a third set of flowers are hermaphrodite.[536] The ovary of
aconites, according to Moquin, is very subject to atrophy.
the indehiscent nature of the latter. Again, the pollen-tube of the abnormal
grains cracked, in its turn, on mere exposure to the air, and liberated the
fovilla, so that the pollen of these atrophied anthers was necessarily
impotent, because it opened before it could be applied to the stigma, even
had that been rendered possible by the opening of the anther.
An abortive condition of the stamens and of the pollen, is of very common
occurrence among hybridised plants. Gaertner and other writers have
spoken of this defective condition as contabescence.[535] It forms one reason
for the sterility so frequently observed in the case of true hybrids. In some
hybrid passion-flowers, while all other parts of the flower were apparently
perfect, even to the ovules, the stamens were atrophied, and distorted, and
contained little or no pollen; the few grains of the latter being smaller than
usual. (See under Heterogamy, pp. 193–196, and p. 398.)
Abortion of the pistil, fruit, &c.—Traces of the carpels occur in many
male flowers of unisexual plants, e.g. Sterculiaceæ, Euphorbiaceæ,
Restiaceæ, &c. &c., and in some natural orders there appears to be a
tendency towards a diœcious condition, e.g. Caryophylleæ, as in Lychnis
dioica, Silene otites, Arenaria tetraquetra, &c. The last-named plant is
stated to have, in some cases, imperfect pistils; in others, rudimentary
stamens; while a third set of flowers are hermaphrodite.[536] The ovary of
aconites, according to Moquin, is very subject to atrophy.
Page 570
Fig. 218.—Bladder plum.
During the maturation of the pistil, and its passage to the fruit, great
changes of consistence frequently take place, owing to the development of
cellular tissue, or of woody matter, according as the fruit is succulent or
woody. It sometimes happens that, owing to some disturbing causes, the
changes that usually occur fail to do so; thus, the stone of plums is
occasionally deficient, as in what are termed bladder-plums (fig. 218); some
of these, consisting merely of a thin bladder, are curiously like the pods of
Colutea.[537]
MM. Fournier and Bonnet[538] describe a fruit of a Rubus, with perfectly
dry fruits, like those of a Geum, and this form was considered by Steudel to
form a distinct species. It is, however, merely a variety in which the fruits
have not become succulent.[539]
During the maturation of the pistil, and its passage to the fruit, great
changes of consistence frequently take place, owing to the development of
cellular tissue, or of woody matter, according as the fruit is succulent or
woody. It sometimes happens that, owing to some disturbing causes, the
changes that usually occur fail to do so; thus, the stone of plums is
occasionally deficient, as in what are termed bladder-plums (fig. 218); some
of these, consisting merely of a thin bladder, are curiously like the pods of
Colutea.[537]
MM. Fournier and Bonnet[538] describe a fruit of a Rubus, with perfectly
dry fruits, like those of a Geum, and this form was considered by Steudel to
form a distinct species. It is, however, merely a variety in which the fruits
have not become succulent.[539]
Page 571
Schlechtendal describes[540] the ordinarily baccate fruit of a vine as
becoming dry, and even dehiscing by valves like a capsule.
In maize it occasionally happens that one or two of the longitudinal series
of fruits become abortive, leaving a smooth furrow, at first of a greenish
colour, but ultimately of a reddish yellow. Often a second row of fruits,
opposite to the first, is also atrophied, so that the whole spike changes its
cylindrical form for a flattened one.[541] See also under Heterogamy,
Meiophylly, &c.
Abortion of the ovules.—In the case of a pluri-ovulate ovary it rarely
happens that all the ovules attain to maturity, some never get fertilised,
others, pressed on by their neighbours on either side, become impeded in
their development, and finally disappear, or remain as rudiments.[542] This
is the case, under ordinary circumstances, and still more so in the case of
hybrid plants, or of monsters. Where the outer coats of the ovule become
more or less leafy in appearance (see p. 262), the inner investments become
more or less atrophied, or are even more frequently entirely suppressed, as
is also the nucleus.
In other cases, a simple arrest of development takes place; the ovule, for
instance, which should be anatropal, remains straight, while the
integuments, checked in their development, form imperfect sheaths from
which the shrivelled nucleus protrudes.
Depauperated Ferns.—The preceding illustrations have been taken from
flowering plants chiefly, but a similar defective development is manifested
in cryptogamous plants. The contraction and imperfect development of the
fronds of some varieties of ferns, hence called depauperated, may receive
passing notice, as also the cases in which the sori or clusters of spore cases
are denuded of their usual covering, owing to the abortion or imperfect
becoming dry, and even dehiscing by valves like a capsule.
In maize it occasionally happens that one or two of the longitudinal series
of fruits become abortive, leaving a smooth furrow, at first of a greenish
colour, but ultimately of a reddish yellow. Often a second row of fruits,
opposite to the first, is also atrophied, so that the whole spike changes its
cylindrical form for a flattened one.[541] See also under Heterogamy,
Meiophylly, &c.
Abortion of the ovules.—In the case of a pluri-ovulate ovary it rarely
happens that all the ovules attain to maturity, some never get fertilised,
others, pressed on by their neighbours on either side, become impeded in
their development, and finally disappear, or remain as rudiments.[542] This
is the case, under ordinary circumstances, and still more so in the case of
hybrid plants, or of monsters. Where the outer coats of the ovule become
more or less leafy in appearance (see p. 262), the inner investments become
more or less atrophied, or are even more frequently entirely suppressed, as
is also the nucleus.
In other cases, a simple arrest of development takes place; the ovule, for
instance, which should be anatropal, remains straight, while the
integuments, checked in their development, form imperfect sheaths from
which the shrivelled nucleus protrudes.
Depauperated Ferns.—The preceding illustrations have been taken from
flowering plants chiefly, but a similar defective development is manifested
in cryptogamous plants. The contraction and imperfect development of the
fronds of some varieties of ferns, hence called depauperated, may receive
passing notice, as also the cases in which the sori or clusters of spore cases
are denuded of their usual covering, owing to the abortion or imperfect
Page 572
development of the indusium, as in what are termed exindusiate varieties.
[543]
General remarks on abortion, coincident changes, &c.—Reference has
already been made, while treating of hypertrophy, suppression, &c., to
certain other changes affecting the flower at the same time. Atrophy of one
organ or set of organs, for instance, is frequently accompanied by a
compensating hypertrophy or by an increased number of other parts. In the
feather-hyacinth, Muscari comosum, var., monstrosum, the absence of
flowers is compensated for by the inordinate formation of brightly coloured
threads which appear to be modified pedicels (see pp. 347, 348); so also in
the wig plant, Rhus Cotinus. So the atrophy of the stamens, in some
flowers, is coincident with the hypertrophy of the pistil. Thus, Unger,
'Denkschr. d. Kais. Acad. der Wissensch. Math. Nat. Classe,' Mai 25, 1848,
p. 103, tab. ix, describes a case wherein the corolla and stamens of
Desmodium marylandicum were atrophied, while the calyx and legume, on
the other hand, were hypertrophied.
Fusion of the members of one whorl with one another, or with the
components of an adjacent series, often entails atrophy or suppression,
either in the united organs themselves, or in adjacent ones. A foliaceous
condition of the outer portions of a flower is very generally attended by
atrophy or complete suppression of the inner portions.
From this point of view the observations of Morren[544] on the different
degrees of atrophy up to complete suppression, observable in the flowers of
Bellevalia comosa, are of importance. According to this observer, the most
highly differentiated parts, such as the stigmas, the ovules, and the anthers,
are the first to disappear, the filaments often being developed without
anthers. Ultimately a deformed and empty perianth alone remains. In the
ordinary course of things the mouth of the perianth is open, but in some of
[543]
General remarks on abortion, coincident changes, &c.—Reference has
already been made, while treating of hypertrophy, suppression, &c., to
certain other changes affecting the flower at the same time. Atrophy of one
organ or set of organs, for instance, is frequently accompanied by a
compensating hypertrophy or by an increased number of other parts. In the
feather-hyacinth, Muscari comosum, var., monstrosum, the absence of
flowers is compensated for by the inordinate formation of brightly coloured
threads which appear to be modified pedicels (see pp. 347, 348); so also in
the wig plant, Rhus Cotinus. So the atrophy of the stamens, in some
flowers, is coincident with the hypertrophy of the pistil. Thus, Unger,
'Denkschr. d. Kais. Acad. der Wissensch. Math. Nat. Classe,' Mai 25, 1848,
p. 103, tab. ix, describes a case wherein the corolla and stamens of
Desmodium marylandicum were atrophied, while the calyx and legume, on
the other hand, were hypertrophied.
Fusion of the members of one whorl with one another, or with the
components of an adjacent series, often entails atrophy or suppression,
either in the united organs themselves, or in adjacent ones. A foliaceous
condition of the outer portions of a flower is very generally attended by
atrophy or complete suppression of the inner portions.
From this point of view the observations of Morren[544] on the different
degrees of atrophy up to complete suppression, observable in the flowers of
Bellevalia comosa, are of importance. According to this observer, the most
highly differentiated parts, such as the stigmas, the ovules, and the anthers,
are the first to disappear, the filaments often being developed without
anthers. Ultimately a deformed and empty perianth alone remains. In the
ordinary course of things the mouth of the perianth is open, but in some of
Page 573
these malformations it is closed, and when that happens, the effects of
atrophy are the more observable in the stamens and pistils.
The impotence of the pollen in certain atrophied flowers, as noticed by the
same observer, is of much interest, especially in reference to the sexual
relationship between the different forms in polymorphic flowers as studied
by Mr. Darwin.
A change in direction may also be noted as a common accompaniment of
atrophy or suppression; thus, in a capsule of Veronica Beccabunga, which
was one-celled by the abortion of one carpel, the style was lateral instead of
terminal.
As to the causes of these structural deviations but little is known; certain of
them have been already alluded to. In some cases atrophy and suppression
maybe regarded as permanent states of a condition usually transitory, but
this is clearly not always the case. Among external causes anything bringing
about an enfeebled condition might be supposed to lead to atrophy, or
suppression of some parts.
Gaertner[545] attributes the arrested development and fall of flowers to some
among the following causes:—1. non-application of the pollen of the same
variety, and consequent imperfect fertilisation; 2. any considerable injury to
the calyx, &c.; 3. destruction of the style or stigma before the fertilisation of
the ovary; 4. application to the stigma of imperfect or heterogeneous pollen
or indifferent pulverulent matter; 5. defective conceptive power in the
ovary.
Abortion of the ovules is considered by the same authority to be due to—1.
deficiency of heat; 2. excess of moisture; 3. peculiar formation of the ovary;
4. over-luxuriant development of roots or buds; 5. peculiar conditions of
cultivation; thus, cuttings and layers produce sterile and abortive seeds
atrophy are the more observable in the stamens and pistils.
The impotence of the pollen in certain atrophied flowers, as noticed by the
same observer, is of much interest, especially in reference to the sexual
relationship between the different forms in polymorphic flowers as studied
by Mr. Darwin.
A change in direction may also be noted as a common accompaniment of
atrophy or suppression; thus, in a capsule of Veronica Beccabunga, which
was one-celled by the abortion of one carpel, the style was lateral instead of
terminal.
As to the causes of these structural deviations but little is known; certain of
them have been already alluded to. In some cases atrophy and suppression
maybe regarded as permanent states of a condition usually transitory, but
this is clearly not always the case. Among external causes anything bringing
about an enfeebled condition might be supposed to lead to atrophy, or
suppression of some parts.
Gaertner[545] attributes the arrested development and fall of flowers to some
among the following causes:—1. non-application of the pollen of the same
variety, and consequent imperfect fertilisation; 2. any considerable injury to
the calyx, &c.; 3. destruction of the style or stigma before the fertilisation of
the ovary; 4. application to the stigma of imperfect or heterogeneous pollen
or indifferent pulverulent matter; 5. defective conceptive power in the
ovary.
Abortion of the ovules is considered by the same authority to be due to—1.
deficiency of heat; 2. excess of moisture; 3. peculiar formation of the ovary;
4. over-luxuriant development of roots or buds; 5. peculiar conditions of
cultivation; thus, cuttings and layers produce sterile and abortive seeds
Page 574
much more frequently than plants of the same species raised from seed; 6.
abortion of the seed is often combined with luxuriant development of the
walls of the fruit.
Temperature and climatal changes in general seem not to be without effect,
as has been already mentioned in the case of Arenaria tetraquetra, which is
polygamous when growing in mountain districts. Other illustrations of a
similar character are mentioned under the head of Heterogamy (p. 196).
Pressure has been already alluded to as one of the most obvious of the
inducing causes of atrophy and suppression.
In the case of Ranunculus auricomus before cited, in which the petals are
rarely perfect, M. de Rochebrune considers that the deficiencies in question
depend, in great measure, on the amount of moisture in the localities where
the plant grows. In most places the flowers and carpels are apt to become
more or less abortive, while the leaves are luxuriant; while, in dry places,
the foliage is small, but the flowers are more perfect. This is quite
consonant with other facts relating to the development of flowers or of
leaves in general.
But while external agencies undoubtedly play some part in bringing about
these changes, it is almost certain that internal causes inherent to the
organization of the plant are more important. Mr. Darwin[546] accounts for
the existence of rudimentary organs by the operation of the general rule of
inheritance, and explains their stunted condition as the effect of disuse, not
so much, of course, in the particular flower as in its predecessors. This
disuse may be the result of the superior efficacy of foreign pollen as
contrasted with that formed in the individual flower itself. In this way many
hermaphrodite flowers tend to become diœcious, as in Caryophyllaceæ,
Orchidaceæ, Plantaginaceæ, Primulaceæ and other orders.
abortion of the seed is often combined with luxuriant development of the
walls of the fruit.
Temperature and climatal changes in general seem not to be without effect,
as has been already mentioned in the case of Arenaria tetraquetra, which is
polygamous when growing in mountain districts. Other illustrations of a
similar character are mentioned under the head of Heterogamy (p. 196).
Pressure has been already alluded to as one of the most obvious of the
inducing causes of atrophy and suppression.
In the case of Ranunculus auricomus before cited, in which the petals are
rarely perfect, M. de Rochebrune considers that the deficiencies in question
depend, in great measure, on the amount of moisture in the localities where
the plant grows. In most places the flowers and carpels are apt to become
more or less abortive, while the leaves are luxuriant; while, in dry places,
the foliage is small, but the flowers are more perfect. This is quite
consonant with other facts relating to the development of flowers or of
leaves in general.
But while external agencies undoubtedly play some part in bringing about
these changes, it is almost certain that internal causes inherent to the
organization of the plant are more important. Mr. Darwin[546] accounts for
the existence of rudimentary organs by the operation of the general rule of
inheritance, and explains their stunted condition as the effect of disuse, not
so much, of course, in the particular flower as in its predecessors. This
disuse may be the result of the superior efficacy of foreign pollen as
contrasted with that formed in the individual flower itself. In this way many
hermaphrodite flowers tend to become diœcious, as in Caryophyllaceæ,
Orchidaceæ, Plantaginaceæ, Primulaceæ and other orders.
Page 575
Although many of the circumstances above mentioned apply to plants
whose structure is habitually rudimentary, there is no reason why they may
not, under due restrictions, be applied to plants whose organs are only
occasionally defective.
For further remarks on the subject of Abortion, the reader is referred to the sections relating
to suppression, etc., also to Moquin-Tandon, 'El. Terat. Veget.,' p. 120; C. Morren, "De
l'atrophie en général," in 'Bull. Acad. Belg.,' t. xviii, 1851, part i, p. 275.
FOOTNOTES:
[522] 'El. Ter. Veg.,' p. 132.
[523] Spinosæ arbores cultura sæpius deponunt spinas in hortis, 'Linn. Phil. Bot.,' § 272.
[524] Mr. Selby, in his 'History of British Forest Trees,' p. 465, gives the following account
of the formation of this peculiar growth:—"In the autumn the parent aphis deposits her eggs at
the base of the embryo leaves, within the bud destined to produce the shoots of the following
year. When these begin to burst and expand in spring, the leaves, at whose bases the eggs have
been deposited, instead of increasing in length, enlarge at the base, and form a cell or cyst
whose mouth is at first closed by a red velvety-looking substance. If opened in this state a nest
of small greenish aphides is distinctly visible, and at a certain period, or when they have
acquired maturity, which is towards the end of the summer, the mouth of the cell opens and the
insects fly off to inflict a similar injury upon the nascent buds of the year. In some instances
the leaves of only a portion of the circumference of a shoot are affected, in which case, though
a slight distortion may take place, the branch is not prevented from elongating; but in others,
where the whole of the leaves around the shoot are converted into nidi, elongation is prevented
and distortion to a great extent takes place."
[525] See Cramer, 'Bildungsabweich.,' pp. 53, 64, for further references.
[526] 'El. Ter. Veg.,' p. 124.
[527] Schlechtendal, 'Bot. Zeit.,' 1857, vol. xv, p. 67.
[528] On the subject of this paragraph the reader may consult A. Braun, "Ueber abnorme
Blattbildung," &c., in 'Verhandl.,' d. 35, 'Naturforscherversammlung;' Jaeger, 'Flora,' 1850. p.
481, tab. 4, Digitalis.
[529] 'Org. Veget.,' i. p. 286.
[530] 'Bull. Soc. Bot. France, vol. viii, 1861, p. 710.
whose structure is habitually rudimentary, there is no reason why they may
not, under due restrictions, be applied to plants whose organs are only
occasionally defective.
For further remarks on the subject of Abortion, the reader is referred to the sections relating
to suppression, etc., also to Moquin-Tandon, 'El. Terat. Veget.,' p. 120; C. Morren, "De
l'atrophie en général," in 'Bull. Acad. Belg.,' t. xviii, 1851, part i, p. 275.
FOOTNOTES:
[522] 'El. Ter. Veg.,' p. 132.
[523] Spinosæ arbores cultura sæpius deponunt spinas in hortis, 'Linn. Phil. Bot.,' § 272.
[524] Mr. Selby, in his 'History of British Forest Trees,' p. 465, gives the following account
of the formation of this peculiar growth:—"In the autumn the parent aphis deposits her eggs at
the base of the embryo leaves, within the bud destined to produce the shoots of the following
year. When these begin to burst and expand in spring, the leaves, at whose bases the eggs have
been deposited, instead of increasing in length, enlarge at the base, and form a cell or cyst
whose mouth is at first closed by a red velvety-looking substance. If opened in this state a nest
of small greenish aphides is distinctly visible, and at a certain period, or when they have
acquired maturity, which is towards the end of the summer, the mouth of the cell opens and the
insects fly off to inflict a similar injury upon the nascent buds of the year. In some instances
the leaves of only a portion of the circumference of a shoot are affected, in which case, though
a slight distortion may take place, the branch is not prevented from elongating; but in others,
where the whole of the leaves around the shoot are converted into nidi, elongation is prevented
and distortion to a great extent takes place."
[525] See Cramer, 'Bildungsabweich.,' pp. 53, 64, for further references.
[526] 'El. Ter. Veg.,' p. 124.
[527] Schlechtendal, 'Bot. Zeit.,' 1857, vol. xv, p. 67.
[528] On the subject of this paragraph the reader may consult A. Braun, "Ueber abnorme
Blattbildung," &c., in 'Verhandl.,' d. 35, 'Naturforscherversammlung;' Jaeger, 'Flora,' 1850. p.
481, tab. 4, Digitalis.
[529] 'Org. Veget.,' i. p. 286.
[530] 'Bull. Soc. Bot. France, vol. viii, 1861, p. 710.
Page 576
[531] 'Linnæa,' 1830, vol. v, p. 492.
[532] 'Mus. Senkenb.,' ii. p. 45.
[533] 'Bull. Acad. Roy. Belg.,' 1851, t. xviii. part i, p. 275.
[534] 'Bull. Soc. Bot. France,' vol. viii, 1861, p. 147.
[535] See Darwin, 'Variation of Domest. Anim. and Plants,' ii, 165.
[536] Gay, 'Ann. Sc. Nat.,' ser. i, 1824, t. iii, p. 44.
[537] See De Candolle, 'Mem. Legum.,' tab. 3, f. 1; Wyville Thomson, 'Trans. Bot. Soc.
Edinb.,' 1851, July 10th; Berkeley, 'Gardeners' Chronicle,' June 22nd, 1867, p. 654. A similar
case is described by Dr. Robb, in Sir W. Hooker's 'Journal of Botany,' 1841, vol. iii, p. 99, with
illustrative figures. The specimens there described were produced at New Brunswick, where
plum trees flower very freely, but seldom produce ripe fruit. Dr. Robb's account is as follows:
—"In the summer of 1839 I had an opportunity of watching the process of destruction among
the plums, and it was as follows—Before or soon after the segments of the corolla had fallen
off, the ovarium had become greenish yellow, soft, and flabby. As the fruit continued to
increase in magnitude, its colour grew darker and of a more ruddy yellow, and at the end of a
fortnight or three weeks the size of the abortive fruit rather exceeded that of a ripe walnut. In
fact, an observer might imagine himself to be walking amongst trees laden with ripe apricots,
but, like the fabled fruit on the banks of the Dead Sea, these plums, though tempting to the
eye, when examined, were found to be hollow, containing air, and consisting only of a
distended skin, insipid, and tasteless. By-and-bye a greenish mould is developed on the surface
of the blighted fruit; then the surface becomes black and shrivelled, and at the expiration of a
month from the time of flowering the whole are rotten and decomposed. The flower appears
about the beginning of June, and before August there is hardly a plum to be seen. It is curious
that where two flower-stalks arise from one point of the branch, one will often go on to ripen
in the normal way, while the other will become abortive, as above described."
In a specimen described by Mr. Berkeley there were two distinct ovules of equal size close to
the apex of the fruit, connected with the base by vessels running down the walls. It should be
observed that there is a worthless variety of plum, Kirke's stoneless, or Sans Noyau, in which
the kernel is not surrounded by any bony deposit.
[538] 'Bull. Soc. Bot. Fr.,' 1862, vol. ix, pp. 37 et 291.
[539] Carl Schimp, 'Fl. Friburg,' vii, p. 745; Hook, fil., 'Journ. Linn. Soc.,' vi, p. 9.
[540] 'Linnæa,' vol. v, 1830, p. 493.
[541] Moquin-Tandon, 'El. Ter. Veg.,' p. 325.
[532] 'Mus. Senkenb.,' ii. p. 45.
[533] 'Bull. Acad. Roy. Belg.,' 1851, t. xviii. part i, p. 275.
[534] 'Bull. Soc. Bot. France,' vol. viii, 1861, p. 147.
[535] See Darwin, 'Variation of Domest. Anim. and Plants,' ii, 165.
[536] Gay, 'Ann. Sc. Nat.,' ser. i, 1824, t. iii, p. 44.
[537] See De Candolle, 'Mem. Legum.,' tab. 3, f. 1; Wyville Thomson, 'Trans. Bot. Soc.
Edinb.,' 1851, July 10th; Berkeley, 'Gardeners' Chronicle,' June 22nd, 1867, p. 654. A similar
case is described by Dr. Robb, in Sir W. Hooker's 'Journal of Botany,' 1841, vol. iii, p. 99, with
illustrative figures. The specimens there described were produced at New Brunswick, where
plum trees flower very freely, but seldom produce ripe fruit. Dr. Robb's account is as follows:
—"In the summer of 1839 I had an opportunity of watching the process of destruction among
the plums, and it was as follows—Before or soon after the segments of the corolla had fallen
off, the ovarium had become greenish yellow, soft, and flabby. As the fruit continued to
increase in magnitude, its colour grew darker and of a more ruddy yellow, and at the end of a
fortnight or three weeks the size of the abortive fruit rather exceeded that of a ripe walnut. In
fact, an observer might imagine himself to be walking amongst trees laden with ripe apricots,
but, like the fabled fruit on the banks of the Dead Sea, these plums, though tempting to the
eye, when examined, were found to be hollow, containing air, and consisting only of a
distended skin, insipid, and tasteless. By-and-bye a greenish mould is developed on the surface
of the blighted fruit; then the surface becomes black and shrivelled, and at the expiration of a
month from the time of flowering the whole are rotten and decomposed. The flower appears
about the beginning of June, and before August there is hardly a plum to be seen. It is curious
that where two flower-stalks arise from one point of the branch, one will often go on to ripen
in the normal way, while the other will become abortive, as above described."
In a specimen described by Mr. Berkeley there were two distinct ovules of equal size close to
the apex of the fruit, connected with the base by vessels running down the walls. It should be
observed that there is a worthless variety of plum, Kirke's stoneless, or Sans Noyau, in which
the kernel is not surrounded by any bony deposit.
[538] 'Bull. Soc. Bot. Fr.,' 1862, vol. ix, pp. 37 et 291.
[539] Carl Schimp, 'Fl. Friburg,' vii, p. 745; Hook, fil., 'Journ. Linn. Soc.,' vi, p. 9.
[540] 'Linnæa,' vol. v, 1830, p. 493.
[541] Moquin-Tandon, 'El. Ter. Veg.,' p. 325.
Page 577
[542] Alph. De Candolle states that the position of the abortive ovules affords a good
character for discriminating between certain species of Quercus, 'Bibl. Univ. Genev.,' 1862, t.
xv, p. 929.
[543] See Moore, 'Nature-Printed Ferns,' 8vo, for numerous illustrations both of depauperate
and exindusiate ferns. Scolopendrium vulgare seems to be one of the ferns most commonly
affected in this way. Moore, loc. cit., vol. ii, pp. 135, 147, 159, 165, &c.
[544] 'Bull. Acad. Belg.,' t. xvii, p. 38, t. 1; Lobelia, p. 85.
[545] Cited in 'Henfrey's Botanical Gazette,' i, p. 179.
[546] 'Origin of Species,' p. 450.
CHAPTER II.
DEGENERATION.
While the terms atrophy and abortion apply in the main to a mere
diminution of size, as contrasted with the ordinary standard, degeneration
may be understood to apply to those cases in which not only is the absolute
bulk diminished, but the whole form is altered and depauperated.
Degeneration, thus, is the result not so much of a deficiency in growth as of
a perversion of development.
Under natural, i.e. habitual circumstances, the formation of pappus in place
of a leafy calyx may be considered as an illustration of degeneration. It is
evident, however, that no very decided line of demarcation can be drawn
between cases of perversion and of arrest of development.
Formation of scales.—These may be mere epidermal excrescences, or they
may be the abortive rudiments of leaves. Of this latter nature are the
"cataphyllary" leaves which invest the root stocks of so many perennial
plants, the perulæ of leaf-buds, or the paleæ on the common receptacle of
character for discriminating between certain species of Quercus, 'Bibl. Univ. Genev.,' 1862, t.
xv, p. 929.
[543] See Moore, 'Nature-Printed Ferns,' 8vo, for numerous illustrations both of depauperate
and exindusiate ferns. Scolopendrium vulgare seems to be one of the ferns most commonly
affected in this way. Moore, loc. cit., vol. ii, pp. 135, 147, 159, 165, &c.
[544] 'Bull. Acad. Belg.,' t. xvii, p. 38, t. 1; Lobelia, p. 85.
[545] Cited in 'Henfrey's Botanical Gazette,' i, p. 179.
[546] 'Origin of Species,' p. 450.
CHAPTER II.
DEGENERATION.
While the terms atrophy and abortion apply in the main to a mere
diminution of size, as contrasted with the ordinary standard, degeneration
may be understood to apply to those cases in which not only is the absolute
bulk diminished, but the whole form is altered and depauperated.
Degeneration, thus, is the result not so much of a deficiency in growth as of
a perversion of development.
Under natural, i.e. habitual circumstances, the formation of pappus in place
of a leafy calyx may be considered as an illustration of degeneration. It is
evident, however, that no very decided line of demarcation can be drawn
between cases of perversion and of arrest of development.
Formation of scales.—These may be mere epidermal excrescences, or they
may be the abortive rudiments of leaves. Of this latter nature are the
"cataphyllary" leaves which invest the root stocks of so many perennial
plants, the perulæ of leaf-buds, or the paleæ on the common receptacle of
Page 578
composite flowers. Other illustrations of a like character are to be met with
in the membranous scales that represent leaves in Ruscus, Asparagus,
Pinus, &c. Similar productions are met with within the flower, where they
may occur as the representatives of sepals, petals, stamens, or pistils, or as
mere excrescences. (See Enation.) Whole families of plants, e.g.
Sapindaceæ, are characterised by the presence of these organs, which are
often of great interest to the morphologist as indicating the true symmetry
of the flower, while they have acquired fresh importance since the
publication of Mr. Darwin's work on the 'Origin of Species,' wherein we are
taught to regard these rudiments as, in many cases, vestiges of organs that
were more completely developed in the progenitors of the present race of
plants, and the exercise of whose functions, from some cause or other,
having been rendered impossible, the structures become, in process of time,
proportionately stunted.
Thus, in diœcious plants we frequently find traces of stamens in the female
flowers, and rudiments of the pistil in the male flower, indicating, according
to the Darwinian hypothesis, that the ancestors of these plants were
hermaphrodite (see Heterogamy).
Mr. Darwin has also shown that, in some cases, the utmost degree of
fertility is attained, not from the action of the pollen on the stigma of the
same flower, but on the influence of the male element of one blossom upon
the female organs of another flower on another individual plant.
Hence, in such plants there is a tendency to a separation of the sexes, while,
from what has been before stated, it might be expected that rudiments of the
male or female organs would be found, and also as a result of the operation
of the law of inheritance. On the same principles it is easy to understand the
occasional presence of the perfect in place of the rudimentary organs, as in
Dianthus.
in the membranous scales that represent leaves in Ruscus, Asparagus,
Pinus, &c. Similar productions are met with within the flower, where they
may occur as the representatives of sepals, petals, stamens, or pistils, or as
mere excrescences. (See Enation.) Whole families of plants, e.g.
Sapindaceæ, are characterised by the presence of these organs, which are
often of great interest to the morphologist as indicating the true symmetry
of the flower, while they have acquired fresh importance since the
publication of Mr. Darwin's work on the 'Origin of Species,' wherein we are
taught to regard these rudiments as, in many cases, vestiges of organs that
were more completely developed in the progenitors of the present race of
plants, and the exercise of whose functions, from some cause or other,
having been rendered impossible, the structures become, in process of time,
proportionately stunted.
Thus, in diœcious plants we frequently find traces of stamens in the female
flowers, and rudiments of the pistil in the male flower, indicating, according
to the Darwinian hypothesis, that the ancestors of these plants were
hermaphrodite (see Heterogamy).
Mr. Darwin has also shown that, in some cases, the utmost degree of
fertility is attained, not from the action of the pollen on the stigma of the
same flower, but on the influence of the male element of one blossom upon
the female organs of another flower on another individual plant.
Hence, in such plants there is a tendency to a separation of the sexes, while,
from what has been before stated, it might be expected that rudiments of the
male or female organs would be found, and also as a result of the operation
of the law of inheritance. On the same principles it is easy to understand the
occasional presence of the perfect in place of the rudimentary organs, as in
Dianthus.
Page 579
In some instances the assumption of a scale-like form by any organ is
attended by a change in texture, the organs becoming dry and scarious, or
fleshy. Moquin cites in illustration of the first phenomenon the flower of a
Vicia, in which the petals were thick and fleshy, like the scales of a bulb;
and of the second the leaves of a Chrysanthemum, which were replaced by
small, glossy scales, like those which invest ordinary leaf-buds. Sometimes
the entire flower is replaced by accumulations of small, acute, green scales.
Cases of this kind, wherein the flowers of a pea and of the foxglove were
replaced by collections of small ovate green scales packed one over the
other till they resembled the strobile of a hop, have been already alluded to.
Most of these scales are represented as having had other accumulations of
scales in their axils.
Similar collections of scales may frequently be met with in the birch and in
the oak, and probably represent abortive leaf-buds. Other cases of a like
kind in Gentiana Amarella, where the scales are coloured, are mentioned
elsewhere.
In some kinds of Campanula a similar change is not uncommon.
Formation of hairs, spines, &c.—The adventitious production of hairs is
likewise frequently due to an arrested growth, in some cases arising from
pressure impeding the proper development of the organ. In other cases the
formation of hair seems to accompany the diminished development of some
organ, as on the barren pedicels of the wig plant, Rhus Cotinus. A similar
production of hair may be noticed in many cases where the development of
a branch or of a flower is arrested, and this occurs with especial frequency
where the arrest in growth is due to the puncture of an insect, or to the
formation of a gall. In such cases the hairs are mere excrescences from the
epidermis.
attended by a change in texture, the organs becoming dry and scarious, or
fleshy. Moquin cites in illustration of the first phenomenon the flower of a
Vicia, in which the petals were thick and fleshy, like the scales of a bulb;
and of the second the leaves of a Chrysanthemum, which were replaced by
small, glossy scales, like those which invest ordinary leaf-buds. Sometimes
the entire flower is replaced by accumulations of small, acute, green scales.
Cases of this kind, wherein the flowers of a pea and of the foxglove were
replaced by collections of small ovate green scales packed one over the
other till they resembled the strobile of a hop, have been already alluded to.
Most of these scales are represented as having had other accumulations of
scales in their axils.
Similar collections of scales may frequently be met with in the birch and in
the oak, and probably represent abortive leaf-buds. Other cases of a like
kind in Gentiana Amarella, where the scales are coloured, are mentioned
elsewhere.
In some kinds of Campanula a similar change is not uncommon.
Formation of hairs, spines, &c.—The adventitious production of hairs is
likewise frequently due to an arrested growth, in some cases arising from
pressure impeding the proper development of the organ. In other cases the
formation of hair seems to accompany the diminished development of some
organ, as on the barren pedicels of the wig plant, Rhus Cotinus. A similar
production of hair may be noticed in many cases where the development of
a branch or of a flower is arrested, and this occurs with especial frequency
where the arrest in growth is due to the puncture of an insect, or to the
formation of a gall. In such cases the hairs are mere excrescences from the
epidermis.
Page 580
Prickles differ but little from hairs save in their more woody texture, but
true spines or thorns are modifications either of a leaf or of a branch. Their
presence seems often dependent on the soil in which the plants grow, or on
other external circumstances.
They occur normally in the sepals of Paronychia serpyllifolia and other
plants.
Formation of glands.—Under this name are associated a number of
(generally) rudimentary organs very different in their morphological nature
and significance, and also in their functions. Some are truly glandular or
secreting organs, while others have no visible office. Anything like a
complete account of these structures would be out of place, and reference is
only made to them here on account of the occasional existence of
intermediate forms, which throw light on the morphological significance of
these structures. Thus, in Passiflora and Viburnum Opulus, the so-called
glands on the sides of the petiole appear to represent leaflets, and are not
unfrequently developed as such.
M. Dunal observed a flower of Cistus vaginatus in which some of the
stamens were replaced by an hypogynous disc.[547] Moquin has seen similar
instances in the flowers of a Rose, Hypericum, and Poppy.
M. Planchon[548] gives an account of some very curious malformations in
Drosera intermedia, which go to show that the ovules are homologous with
the glandular hairs on the margins of the leaves of these plants, an opinion
corroborated by the researches of MM. Grönland and Trécul.[549]
Dr. Hooker shows that the pitcher of Nepenthes is due to a modification of a
gland placed at the extremity of the midrib.[550]
true spines or thorns are modifications either of a leaf or of a branch. Their
presence seems often dependent on the soil in which the plants grow, or on
other external circumstances.
They occur normally in the sepals of Paronychia serpyllifolia and other
plants.
Formation of glands.—Under this name are associated a number of
(generally) rudimentary organs very different in their morphological nature
and significance, and also in their functions. Some are truly glandular or
secreting organs, while others have no visible office. Anything like a
complete account of these structures would be out of place, and reference is
only made to them here on account of the occasional existence of
intermediate forms, which throw light on the morphological significance of
these structures. Thus, in Passiflora and Viburnum Opulus, the so-called
glands on the sides of the petiole appear to represent leaflets, and are not
unfrequently developed as such.
M. Dunal observed a flower of Cistus vaginatus in which some of the
stamens were replaced by an hypogynous disc.[547] Moquin has seen similar
instances in the flowers of a Rose, Hypericum, and Poppy.
M. Planchon[548] gives an account of some very curious malformations in
Drosera intermedia, which go to show that the ovules are homologous with
the glandular hairs on the margins of the leaves of these plants, an opinion
corroborated by the researches of MM. Grönland and Trécul.[549]
Dr. Hooker shows that the pitcher of Nepenthes is due to a modification of a
gland placed at the extremity of the midrib.[550]
Page 581
Formation of tendrils.—These are of very varied morphological import;
sometimes they are degenerated peduncles, as in passion-flowers, or vines;
at other times they are of foliar origin; or, again, they may proceed from the
segments of the perianth, as in Hodgsonia and some other cucurbitaceous
plants. From their very different origin in different plants it is necessary to
study the development in each case, and not apply to the generality what
may be peculiar to one. In any case this formation in question generally
belongs more to general morphology than to teratology.[551]
Kirschleger, however, has recorded the existence of a cirrhose sepal in
Cucurbita Pepo.[552]
FOOTNOTES:
[547] 'Consid. Org. Fleur.,' p. 44, pl. ii, fig. 23.
[548] 'Ann. Sc. Nat.,' 3 ser., Bot. ix, pl. 6, ff. 1, 2.
[549] 'Ann. Sc. Nat.,' 3 ser., Bot. 1855, pp. 297, et 303.
[550] 'Trans. Linn. Soc.,' xxii, p. 415.
[551] See Darwin, "On Climbing Plants," 'Journal of Linnean Society,' vol. ix, p. 1.
[552] 'Flora,' 1845, p. 615.
sometimes they are degenerated peduncles, as in passion-flowers, or vines;
at other times they are of foliar origin; or, again, they may proceed from the
segments of the perianth, as in Hodgsonia and some other cucurbitaceous
plants. From their very different origin in different plants it is necessary to
study the development in each case, and not apply to the generality what
may be peculiar to one. In any case this formation in question generally
belongs more to general morphology than to teratology.[551]
Kirschleger, however, has recorded the existence of a cirrhose sepal in
Cucurbita Pepo.[552]
FOOTNOTES:
[547] 'Consid. Org. Fleur.,' p. 44, pl. ii, fig. 23.
[548] 'Ann. Sc. Nat.,' 3 ser., Bot. ix, pl. 6, ff. 1, 2.
[549] 'Ann. Sc. Nat.,' 3 ser., Bot. 1855, pp. 297, et 303.
[550] 'Trans. Linn. Soc.,' xxii, p. 415.
[551] See Darwin, "On Climbing Plants," 'Journal of Linnean Society,' vol. ix, p. 1.
[552] 'Flora,' 1845, p. 615.
Page 582
GENERAL CONCLUSIONS.
At the end of many of the preceding sections, and whenever the
requirements of the case demanded it, a brief summary of the main facts
and of the inferences to be derived from them has been given. It may be
useful to give in conclusion a few general remarks on the whole subject.
It will be seen from the numerous facts herein cited, that the so-called
monstrous formations (excluding morbid growths the result of disease or
injury) present no peculiarities absolutely foreign to the normal organisation
of plants. The difference between the natural and monstrous development is
one of degree and frequency of occurrence, not of kind.
Deviations from the customary form have been shown to arise from
excessive or diminished growth, or from arrested or exalted development.
Even in those instances where, for convenience' sake, the term perverted
development has been used, it must be understood as applying only to the
particular plant or organ under consideration, as the form assumed is
perfectly in accordance with the ordinary conformation of some other plant
or group of plants.
The period at which malformations occur is a matter of some importance;
this is, indeed, implied in the term arrest of development; evolution goes on
with growth up to a certain point and is then stopped, and thus changes are
At the end of many of the preceding sections, and whenever the
requirements of the case demanded it, a brief summary of the main facts
and of the inferences to be derived from them has been given. It may be
useful to give in conclusion a few general remarks on the whole subject.
It will be seen from the numerous facts herein cited, that the so-called
monstrous formations (excluding morbid growths the result of disease or
injury) present no peculiarities absolutely foreign to the normal organisation
of plants. The difference between the natural and monstrous development is
one of degree and frequency of occurrence, not of kind.
Deviations from the customary form have been shown to arise from
excessive or diminished growth, or from arrested or exalted development.
Even in those instances where, for convenience' sake, the term perverted
development has been used, it must be understood as applying only to the
particular plant or organ under consideration, as the form assumed is
perfectly in accordance with the ordinary conformation of some other plant
or group of plants.
The period at which malformations occur is a matter of some importance;
this is, indeed, implied in the term arrest of development; evolution goes on
with growth up to a certain point and is then stopped, and thus changes are
Page 583
brought about in the part affected of a different nature from those dependent
on non-development or suppression.
Some malformations are congenital, therefore, while others are acquired—
in the former instance the disturbance is coeval in origin, and
contemporaneous in its growth and development, with those of the affected
part; in the latter case the organ may have attained its ordinary degree of
perfection, or at least may have advanced some way towards it, before any
deviation shows itself. True chorisis or fission, for instance, is usually a
congenital affection, arising at a very early period of development, while
enation takes place from structures which are all but complete as to their
organisation, even though they may not have attained their full dimensions.
The date of appearance is also of consequence in determining the true
nature of some changes; it does not always follow, for instance, that
because one organ occupies the position of another, it is of the same nature
as the one whose place it fills. The presence of anthers on petals or on such
organs as the corona of Narcissus does not necessarily constitute those parts
actual stamens, but rather staminodes. The true stamens are either wanting,
or if present, they are in advance of their imitators as regards their
development.
General morphology of the leaf and axis. Homology. Since the time
when Goethe's generalisations were adopted by A. P. De Caudolle, special
attention has been given to the form and mode of development of the leaf-
organ; for as it was well said by Wolff, if once the course of evolution and
the structure of the leaf were known, those of the parts of the flower would
follow as a matter of course.
It is not necessary, in this place, to pursue the subject of the development
and construction of the leaf further than they are illustrated by ordinary
teratological phenomena.
on non-development or suppression.
Some malformations are congenital, therefore, while others are acquired—
in the former instance the disturbance is coeval in origin, and
contemporaneous in its growth and development, with those of the affected
part; in the latter case the organ may have attained its ordinary degree of
perfection, or at least may have advanced some way towards it, before any
deviation shows itself. True chorisis or fission, for instance, is usually a
congenital affection, arising at a very early period of development, while
enation takes place from structures which are all but complete as to their
organisation, even though they may not have attained their full dimensions.
The date of appearance is also of consequence in determining the true
nature of some changes; it does not always follow, for instance, that
because one organ occupies the position of another, it is of the same nature
as the one whose place it fills. The presence of anthers on petals or on such
organs as the corona of Narcissus does not necessarily constitute those parts
actual stamens, but rather staminodes. The true stamens are either wanting,
or if present, they are in advance of their imitators as regards their
development.
General morphology of the leaf and axis. Homology. Since the time
when Goethe's generalisations were adopted by A. P. De Caudolle, special
attention has been given to the form and mode of development of the leaf-
organ; for as it was well said by Wolff, if once the course of evolution and
the structure of the leaf were known, those of the parts of the flower would
follow as a matter of course.
It is not necessary, in this place, to pursue the subject of the development
and construction of the leaf further than they are illustrated by ordinary
teratological phenomena.
Page 584
From this point of view perhaps the most interesting circumstance is the
part that the sheath of the leaf plays.[553] In many cases of so-called
metamorphosis, it is the sheath of the leaf that is represented and not the
blade. In normal anatomy the sepals, petals, carpels, and even the stamens,
as a general rule, correspond to the sheath rather than to the blade of the
leaf, as may be seen by the arrangement of the veins. The blade of the leaf
seems to be set apart for special respiratory and absorbent offices, while the
sheath is in structure, if not in office, more akin to the stem. It would not be
easy apart from their position to distinguish between a tubular sheathing
leaf and a hollow stem. The development of adventitious growths by
chorisis or enation has been frequently alluded to in the foregoing pages,
and many illustrations have been given of the power that leaves have of
branching in more than one plane, owing to the projection of secondary
growing-points from the primary organ. These new centres of development
are closely connected with the fibro-vascular system of the leaf, so that no
sooner does a new growing point originate, than vessels are formed to
connect the new growth with the general fibrous cord, see pp. 355, 445.
This leads M. Casimir De Candollo to consider the entire leaf as a
composite structure. The morphological unit, says he, is the cellular
protrusion or growing point (saillie) and its corresponding fibro-vascular
bundle.[554]
The identity, in a morphological point of view, of the leaves and the lateral
parts of the flower is so thoroughly recognised that little need be said on
that score, save to repeat that the homology of the floral organs is usually
not so much with the entire leaf as with its sheath.
The most singular instances of morphological identity are those relating to
the sexual organs. We have seen the gradual transition of stamens to pistils,
and of pistils to stamens, the development of ovules on the edges of the
anther, the co-existence of pollen with ovules on an antheroid body, and,
part that the sheath of the leaf plays.[553] In many cases of so-called
metamorphosis, it is the sheath of the leaf that is represented and not the
blade. In normal anatomy the sepals, petals, carpels, and even the stamens,
as a general rule, correspond to the sheath rather than to the blade of the
leaf, as may be seen by the arrangement of the veins. The blade of the leaf
seems to be set apart for special respiratory and absorbent offices, while the
sheath is in structure, if not in office, more akin to the stem. It would not be
easy apart from their position to distinguish between a tubular sheathing
leaf and a hollow stem. The development of adventitious growths by
chorisis or enation has been frequently alluded to in the foregoing pages,
and many illustrations have been given of the power that leaves have of
branching in more than one plane, owing to the projection of secondary
growing-points from the primary organ. These new centres of development
are closely connected with the fibro-vascular system of the leaf, so that no
sooner does a new growing point originate, than vessels are formed to
connect the new growth with the general fibrous cord, see pp. 355, 445.
This leads M. Casimir De Candollo to consider the entire leaf as a
composite structure. The morphological unit, says he, is the cellular
protrusion or growing point (saillie) and its corresponding fibro-vascular
bundle.[554]
The identity, in a morphological point of view, of the leaves and the lateral
parts of the flower is so thoroughly recognised that little need be said on
that score, save to repeat that the homology of the floral organs is usually
not so much with the entire leaf as with its sheath.
The most singular instances of morphological identity are those relating to
the sexual organs. We have seen the gradual transition of stamens to pistils,
and of pistils to stamens, the development of ovules on the edges of the
anther, the co-existence of pollen with ovules on an antheroid body, and,
Page 585
stranger still, the actual development of pollen within the tissues of the
ovule itself! From such facts, in addition to what we know of the relative
position, internal structure, and mode of development of the organs, it is
impossible to avoid coming to the conclusion that, however distinctly these
parts may, under ordinary circumstances, be set apart for the performance of
distinct functions, morphologically they are homologous.
These ideas may be carried yet farther—the same sort of evidence, which is
adduced in support of the morphological identity of leaves with the parts of
the flower, may be advanced in confirmation of the opinion, that,
morphologically, there is no distinction between axis and leaf. The leaf,
according to this view, is a specialised portion of the axis set apart to do
certain work, just as the petals, stamens, &c., are leaves told off for distinct
uses. It is unnecessary to refer to the intermediate productions linking the
leaf-form to that of the axis, all that is requisite here is to point out the facts
that teratology lends in support of these views. These may be summed up
by the statement that almost all those attributes which morphologists
recognise as peculiar to one or the other organ respectively, may be and are
manifested by both. We have the stem acquiring the characters of the leaf,
and the leaf those of the stem. Thus we have seen leaves, leaf-buds,
branches, and flower-buds springing from leaves or leaf-organs;[555] see pp.
174, 177, 445, &c. The structure that we are apt to associate exclusively
with one is found to pertain to the other. The arrangement of the vascular
cords in the leaf-organ finds its counterpart in the axis, generally, it is true,
modified to suit altered circumstances or diverse purposes. In some cases
the disposition is absolutely indistinguishable in the two organs. It may then
be said that the distinctions usually drawn between axis and leaf are not
absolute, and that, however necessary such a separation may be for
descriptive or physiological purposes, morphologically the two organs are
identical. Again, it may be said that leaf and axis are two phases of the same
organ,—an organ capable of existing in its undifferentiated state in the form
ovule itself! From such facts, in addition to what we know of the relative
position, internal structure, and mode of development of the organs, it is
impossible to avoid coming to the conclusion that, however distinctly these
parts may, under ordinary circumstances, be set apart for the performance of
distinct functions, morphologically they are homologous.
These ideas may be carried yet farther—the same sort of evidence, which is
adduced in support of the morphological identity of leaves with the parts of
the flower, may be advanced in confirmation of the opinion, that,
morphologically, there is no distinction between axis and leaf. The leaf,
according to this view, is a specialised portion of the axis set apart to do
certain work, just as the petals, stamens, &c., are leaves told off for distinct
uses. It is unnecessary to refer to the intermediate productions linking the
leaf-form to that of the axis, all that is requisite here is to point out the facts
that teratology lends in support of these views. These may be summed up
by the statement that almost all those attributes which morphologists
recognise as peculiar to one or the other organ respectively, may be and are
manifested by both. We have the stem acquiring the characters of the leaf,
and the leaf those of the stem. Thus we have seen leaves, leaf-buds,
branches, and flower-buds springing from leaves or leaf-organs;[555] see pp.
174, 177, 445, &c. The structure that we are apt to associate exclusively
with one is found to pertain to the other. The arrangement of the vascular
cords in the leaf-organ finds its counterpart in the axis, generally, it is true,
modified to suit altered circumstances or diverse purposes. In some cases
the disposition is absolutely indistinguishable in the two organs. It may then
be said that the distinctions usually drawn between axis and leaf are not
absolute, and that, however necessary such a separation may be for
descriptive or physiological purposes, morphologically the two organs are
identical. Again, it may be said that leaf and axis are two phases of the same
organ,—an organ capable of existing in its undifferentiated state in the form
Page 586
of a thallus among Cryptogams, but which in the higher groups of plants
becomes marked out into separate portions, each portion having its own
distinct functions to fulfil for the common benefit of the whole organisation.
[556]
Special morphology.—Under this heading brief reference may be made to
some of the organs whose morphological nature has been, and still is, much
contested. It is clear that for the due elucidation of these matters,
development and the comparative investigation of similar structures in
different plants must be studied. Teratological data by themselves can no
more be trusted to give a correct solution of any particular question, than
the evidence furnished by other departments of botanical science taken
separately. With this statement by way of caution, allusion may be made to
some of the organs whose morphological construction is illustrated by the
facts recorded in the present volume.
Calyx-tube.—In descriptive botany it is the common practice to speak of a
calyx-tube, by which is meant a tubular or sheathing portion at the base of
the flower, below the sepals or calyx-lobes, and distinct or inseparable from
the ovary. The question morphology has to solve is whether this tubular
structure is to be considered as a portion of the axis, or whether it is to be
regarded as composed of the confluent bases of the sepals.
Mr. Bentham, who has recently reviewed the evidence as to the nature of
the calyx-tube in his paper on Myrtaceæ,[557] still holds to the notion that
the "calyx-tube" or "hypanthium" is formed from the concretion of the basal
portions of the sepals. He founds his conclusions upon such facts as the
following: the circumstance that the point of origin of the leaf is not always
the same as the point of disarticulation or separation from the axis,
inasmuch as the basal portion of the leaf is often adherent to the stem for
some distance, though still recognisable as foliar not axial in its nature. In
becomes marked out into separate portions, each portion having its own
distinct functions to fulfil for the common benefit of the whole organisation.
[556]
Special morphology.—Under this heading brief reference may be made to
some of the organs whose morphological nature has been, and still is, much
contested. It is clear that for the due elucidation of these matters,
development and the comparative investigation of similar structures in
different plants must be studied. Teratological data by themselves can no
more be trusted to give a correct solution of any particular question, than
the evidence furnished by other departments of botanical science taken
separately. With this statement by way of caution, allusion may be made to
some of the organs whose morphological construction is illustrated by the
facts recorded in the present volume.
Calyx-tube.—In descriptive botany it is the common practice to speak of a
calyx-tube, by which is meant a tubular or sheathing portion at the base of
the flower, below the sepals or calyx-lobes, and distinct or inseparable from
the ovary. The question morphology has to solve is whether this tubular
structure is to be considered as a portion of the axis, or whether it is to be
regarded as composed of the confluent bases of the sepals.
Mr. Bentham, who has recently reviewed the evidence as to the nature of
the calyx-tube in his paper on Myrtaceæ,[557] still holds to the notion that
the "calyx-tube" or "hypanthium" is formed from the concretion of the basal
portions of the sepals. He founds his conclusions upon such facts as the
following: the circumstance that the point of origin of the leaf is not always
the same as the point of disarticulation or separation from the axis,
inasmuch as the basal portion of the leaf is often adherent to the stem for
some distance, though still recognisable as foliar not axial in its nature. In
Page 587
the same manner, the corolla and andrœcium may be concrete at the base,
so that the stamens are for convenience' sake described as inserted into the
tube of the corolla, though it is generally admitted that both stamens and
petals are really hypogynous, and it is not usual to consider the corolla-tube
up to the divergence of the stamens as part of the receptacle. A similar
remark applies to the carpels and placentas. Mr. Bentham further considers
that the gradual disconnection of the various whorls, that may be traced in
many plants, is a further proof of concretion, rather than of expansion of the
axis, but this argument may fairly be met by the consideration that the
several whorls emerge at different heights.[558]
Organs originally free and distinct become ultimately combined at the base
by the gradual protrusion from the receptacle of a ring or tube under them,
as in the stamens of Leguminosæ; yet, says Mr. Bentham, no one would
propose to describe the staminal tube of monadelphous Leguminosæ as part
of the receptacle and not of the stamens. Perhaps not, for descriptive
purposes, but morphologically it would not be easy to separate such a tube
from the receptacle. The principal kinds of malformation which have a
bearing on this subject are mentioned at pp. 77–81 and 247, from which it
may be seen that the evidence furnished by teratology is conflicting. It
would seem, indeed, that while in some families of plants there may be a
real calyx-tube, in others the tubular portion is a sheath-like prolongation of
the axis. In Primula or Pedicularis, where the venation is clearly laminar,
the tubular portion is distinctly calycine. In other cases the so-called calyx-
tube seems as certainly to be an expansion of the receptacle, as in Rosaceæ,
Myrtaceæ, Melastomaceæ, Passiflora,[559] &c.
Where the petals and stamens are described as being inserted into the throat
of the calyx, or are perigynous, it may be assumed as a general rule, subject
to but few exceptions, that the so-called calyx-tube is really a portion of the
receptacle.[560] After all, this is very much a question of words, and for the
so that the stamens are for convenience' sake described as inserted into the
tube of the corolla, though it is generally admitted that both stamens and
petals are really hypogynous, and it is not usual to consider the corolla-tube
up to the divergence of the stamens as part of the receptacle. A similar
remark applies to the carpels and placentas. Mr. Bentham further considers
that the gradual disconnection of the various whorls, that may be traced in
many plants, is a further proof of concretion, rather than of expansion of the
axis, but this argument may fairly be met by the consideration that the
several whorls emerge at different heights.[558]
Organs originally free and distinct become ultimately combined at the base
by the gradual protrusion from the receptacle of a ring or tube under them,
as in the stamens of Leguminosæ; yet, says Mr. Bentham, no one would
propose to describe the staminal tube of monadelphous Leguminosæ as part
of the receptacle and not of the stamens. Perhaps not, for descriptive
purposes, but morphologically it would not be easy to separate such a tube
from the receptacle. The principal kinds of malformation which have a
bearing on this subject are mentioned at pp. 77–81 and 247, from which it
may be seen that the evidence furnished by teratology is conflicting. It
would seem, indeed, that while in some families of plants there may be a
real calyx-tube, in others the tubular portion is a sheath-like prolongation of
the axis. In Primula or Pedicularis, where the venation is clearly laminar,
the tubular portion is distinctly calycine. In other cases the so-called calyx-
tube seems as certainly to be an expansion of the receptacle, as in Rosaceæ,
Myrtaceæ, Melastomaceæ, Passiflora,[559] &c.
Where the petals and stamens are described as being inserted into the throat
of the calyx, or are perigynous, it may be assumed as a general rule, subject
to but few exceptions, that the so-called calyx-tube is really a portion of the
receptacle.[560] After all, this is very much a question of words, and for the
Page 588
following reasons,—very often the base of the calyx does evidently form a
tube, and no one can say where the calyx ends and the receptacle begins.
Again, many leaves are known to originate in the form of a ring-like
protrusion from the axis, and from this primary ring originate secondary
developments. Thus the asserted difference between a leaf, with such a
history of development, and an axial structure becomes obliterated. From
this point of view, peltate leaves like those of Tropæolum or Nelumbium
become very significant. In both the leaf-stalk is cylindrical and traversed,
as in the case of all cylindrical leaf-stalks, by a circle of fibro-vascular
cords, as in a branch, and which radiate in all directions in the blade of the
leaf. Now, if (as often happens to a slight extent) the central portion of the
leaf were much depressed, owing to the disproportionate growth of the
peripheral, as contrasted with the central portions, we should have a funnel-
like or tubular formation, precisely similar to many of the so-called calyx-
tubes. And, if we further suppose new growths to originate from the sides of
this funnel or tube, by chorisis or enation, we should have the homologue of
a tubular calyx, to the inner surface of which are attached petals, stamens,
&c. From the consideration of circumstances such as these just detailed,
together with that of the arrangement of the vascular cords, M. Casimir De
Candolle arrives at the conclusion that the calyx-tube is a ring-like
projection from an axis whose further direct development is arrested. The
secondary projections or growing-points correspond to the several fibro-
vascular cords of the primary ring, and are ultimately developed into sepals,
petals, stamens and ovaries (see pp. 394, 509).
Andrœcium.—The main points of morphological interest relating to the
andrœcium, referred to in this volume, are those concerning the structure of
the anther (see p. 292), the compound nature of the stamens in some orders
(see pp. 294, 345), and the nature of the andrœcium in orchids (see p. 380).
tube, and no one can say where the calyx ends and the receptacle begins.
Again, many leaves are known to originate in the form of a ring-like
protrusion from the axis, and from this primary ring originate secondary
developments. Thus the asserted difference between a leaf, with such a
history of development, and an axial structure becomes obliterated. From
this point of view, peltate leaves like those of Tropæolum or Nelumbium
become very significant. In both the leaf-stalk is cylindrical and traversed,
as in the case of all cylindrical leaf-stalks, by a circle of fibro-vascular
cords, as in a branch, and which radiate in all directions in the blade of the
leaf. Now, if (as often happens to a slight extent) the central portion of the
leaf were much depressed, owing to the disproportionate growth of the
peripheral, as contrasted with the central portions, we should have a funnel-
like or tubular formation, precisely similar to many of the so-called calyx-
tubes. And, if we further suppose new growths to originate from the sides of
this funnel or tube, by chorisis or enation, we should have the homologue of
a tubular calyx, to the inner surface of which are attached petals, stamens,
&c. From the consideration of circumstances such as these just detailed,
together with that of the arrangement of the vascular cords, M. Casimir De
Candolle arrives at the conclusion that the calyx-tube is a ring-like
projection from an axis whose further direct development is arrested. The
secondary projections or growing-points correspond to the several fibro-
vascular cords of the primary ring, and are ultimately developed into sepals,
petals, stamens and ovaries (see pp. 394, 509).
Andrœcium.—The main points of morphological interest relating to the
andrœcium, referred to in this volume, are those concerning the structure of
the anther (see p. 292), the compound nature of the stamens in some orders
(see pp. 294, 345), and the nature of the andrœcium in orchids (see p. 380).
Page 589
Inferior ovary.—Is the pistil always foliar in its morphological nature, or is
it, in some cases, as Schleiden taught, formed from the axis alone? To a
great extent the reply to this question is dependent on the conclusions that
may be arrived at as to the true nature of the calyx-tube. Considered from a
teratological point of view, there is no reason for considering the inferior
ovary to be purely axial. On the contrary, the evidence derived from this
source supports the ordinary opinion that the carpels are invaginated within
the expanded top of the flower-stalk and more or less adherent to it. Some
of the gourds afford good illustrations of this, the upper part of the carpels
in these fruits projecting beyond the axial portion. But this matter loses
much of its importance if the morphological identity of axis and leaf-organ
be conceded. The carpels in inferior ovaries seldom or never correspond to
the lamina of the leaf, and between the vaginal portion of the carpellary
leaf, and the axis who shall draw the distinction?
Placentation.—Some botanists have considered the placentas to be
portions of the carpel, and have compared the production of ovules on them
to the formation of buds on the leaf of Bryophyllum. Others have been led
to see in each placenta, even when it is, to all outward appearance, a portion
of the carpellary leaf, a direct prolongation from the axis, adherent to the
leaf. Teratology shows that ovules may be formed indifferently on leaf-
organs or on stem-organs. Sutural, parietal, axile, free-central placentation,
and, if there be more forms, all may be met with even in the same ovary
(see pp. 96, 508). Now, if there were such special tendencies in the axis, as
contrasted with the leaf, to produce ovules, it is hardly likely that such
anomalous arrangements as those just mentioned would be as frequent as
they are. But as leaves produce other leaves, from their edges or their
surfaces, and as they form buds in the same situations, just as axial organs
do,[561] there is surely little ground for considering the placentas, or
ovuliferous portions of the plant, to be of necessity axial. Here again, much
it, in some cases, as Schleiden taught, formed from the axis alone? To a
great extent the reply to this question is dependent on the conclusions that
may be arrived at as to the true nature of the calyx-tube. Considered from a
teratological point of view, there is no reason for considering the inferior
ovary to be purely axial. On the contrary, the evidence derived from this
source supports the ordinary opinion that the carpels are invaginated within
the expanded top of the flower-stalk and more or less adherent to it. Some
of the gourds afford good illustrations of this, the upper part of the carpels
in these fruits projecting beyond the axial portion. But this matter loses
much of its importance if the morphological identity of axis and leaf-organ
be conceded. The carpels in inferior ovaries seldom or never correspond to
the lamina of the leaf, and between the vaginal portion of the carpellary
leaf, and the axis who shall draw the distinction?
Placentation.—Some botanists have considered the placentas to be
portions of the carpel, and have compared the production of ovules on them
to the formation of buds on the leaf of Bryophyllum. Others have been led
to see in each placenta, even when it is, to all outward appearance, a portion
of the carpellary leaf, a direct prolongation from the axis, adherent to the
leaf. Teratology shows that ovules may be formed indifferently on leaf-
organs or on stem-organs. Sutural, parietal, axile, free-central placentation,
and, if there be more forms, all may be met with even in the same ovary
(see pp. 96, 508). Now, if there were such special tendencies in the axis, as
contrasted with the leaf, to produce ovules, it is hardly likely that such
anomalous arrangements as those just mentioned would be as frequent as
they are. But as leaves produce other leaves, from their edges or their
surfaces, and as they form buds in the same situations, just as axial organs
do,[561] there is surely little ground for considering the placentas, or
ovuliferous portions of the plant, to be of necessity axial. Here again, much
Page 590
of the difficulty vanishes if the morphological identity of the leaf-form and
of the stem-form be admitted.
Structure of the ovule.—The nature of the ovule and of its coverings has
been a fertile source of controversy. The teratological data bearing on this
subject have been given at pp. 262–272. These data strongly support the
notion of the foliar nature of the coatings, and of the axial nature of the
nucleus, taking leaf and axis either in the ordinary sense, or as
modifications one of the other. It has been shown that the ovular coats may
themselves become carpels, and that ovules may be developed upon ovules,
p. 268. Whether the intra-carpellary siliques of Cheiranthus, not
uncommonly met with (p. 182), are instances of ovular transmutation may
be open to doubt.
The axial nature of the nucleus has been inferred from its position, mode of
growth, and from its occasionally lengthening into a leafy or even a
floriferous shoot. Probably it may occasionally be invested by sheathing
coats, more analogous to tubular processes from the receptacle, than to
foliar organs, as is the case in Welwitschia. The discussion of this matter,
however, pertains rather to normal morphology than to teratology.
Morphology of conifers.—The nature of the pseudo-leaves of Sciadopitys,
and probably of other Conifers, is illustrated by teratology, as also is the
true constitution of the scale of the cone (see pp. 192, 245, 352), though it
must be admitted that little or no light is thrown on that much-contested
point—the true nature of the ovule of Gymnosperms.
Relative position of organs.—When organs are considered, not separately,
but in their relations to each other, the appearances presented are referable
to similar causes. Thus, the separation of parts usually united has been
shown to depend on an excess of development, the persistent union of parts,
usually separate in the adult state, has been traced to an arrest of the process
of the stem-form be admitted.
Structure of the ovule.—The nature of the ovule and of its coverings has
been a fertile source of controversy. The teratological data bearing on this
subject have been given at pp. 262–272. These data strongly support the
notion of the foliar nature of the coatings, and of the axial nature of the
nucleus, taking leaf and axis either in the ordinary sense, or as
modifications one of the other. It has been shown that the ovular coats may
themselves become carpels, and that ovules may be developed upon ovules,
p. 268. Whether the intra-carpellary siliques of Cheiranthus, not
uncommonly met with (p. 182), are instances of ovular transmutation may
be open to doubt.
The axial nature of the nucleus has been inferred from its position, mode of
growth, and from its occasionally lengthening into a leafy or even a
floriferous shoot. Probably it may occasionally be invested by sheathing
coats, more analogous to tubular processes from the receptacle, than to
foliar organs, as is the case in Welwitschia. The discussion of this matter,
however, pertains rather to normal morphology than to teratology.
Morphology of conifers.—The nature of the pseudo-leaves of Sciadopitys,
and probably of other Conifers, is illustrated by teratology, as also is the
true constitution of the scale of the cone (see pp. 192, 245, 352), though it
must be admitted that little or no light is thrown on that much-contested
point—the true nature of the ovule of Gymnosperms.
Relative position of organs.—When organs are considered, not separately,
but in their relations to each other, the appearances presented are referable
to similar causes. Thus, the separation of parts usually united has been
shown to depend on an excess of development, the persistent union of parts,
usually separate in the adult state, has been traced to an arrest of the process
Page 591
of development, by no means necessarily coexistent with diminished
growth. The diminished or increased number of parts is, in like manner,
attributable to analogous causes, as also are the variations in arrangement
and form, spoken of under the heads of Displacement, Peloria, Substitution,
&c.
In the instance of displacements, it has been shown how slight a change is
required to transform the so-called inferior ovary into a superior one. A
defective development of the top of the flower-stalk in some cases, in others
a lack of union between the tube of the receptacle or of the calyx
(comprising in those terms not only the apex of the receptacle, but the base
of the sepals) and the carpels, suffice to bring about this change in a
character which for systematic purposes is of great value.
Law of alternation.—The circumstances that interfere with the law of
alternation may be briefly alluded to. The deviations from the customary
arrangement have been very generally attributed to suppression, or to
chorisis. It is unquestionable that either of these affords an efficient
explanation of the arrangement in question, as also does that modification
of chorisis, as it may be considered, which has been treated of under the
head of Enation. Spiral torsion of the axis would likewise bring about
analogous results. Still, it is quite conceivable that opposition or
superposition of organs may occur without the intervention of any such
operations. This will be the more readily conceded when it is remembered
that the phyllotaxis of leaves not unfrequently varies on different branches
of the same individual tree, and that a similar variation in the flower would
at once disturb the customary alternate arrangement. Coalescence of the
vascular bundles in an unusual manner, and an irregular disposition of these
cords have also been considered to bring about deviations from the rule of
alternation, but in general the formation of the cords is subsequent to that of
the growing points or mamelons.
growth. The diminished or increased number of parts is, in like manner,
attributable to analogous causes, as also are the variations in arrangement
and form, spoken of under the heads of Displacement, Peloria, Substitution,
&c.
In the instance of displacements, it has been shown how slight a change is
required to transform the so-called inferior ovary into a superior one. A
defective development of the top of the flower-stalk in some cases, in others
a lack of union between the tube of the receptacle or of the calyx
(comprising in those terms not only the apex of the receptacle, but the base
of the sepals) and the carpels, suffice to bring about this change in a
character which for systematic purposes is of great value.
Law of alternation.—The circumstances that interfere with the law of
alternation may be briefly alluded to. The deviations from the customary
arrangement have been very generally attributed to suppression, or to
chorisis. It is unquestionable that either of these affords an efficient
explanation of the arrangement in question, as also does that modification
of chorisis, as it may be considered, which has been treated of under the
head of Enation. Spiral torsion of the axis would likewise bring about
analogous results. Still, it is quite conceivable that opposition or
superposition of organs may occur without the intervention of any such
operations. This will be the more readily conceded when it is remembered
that the phyllotaxis of leaves not unfrequently varies on different branches
of the same individual tree, and that a similar variation in the flower would
at once disturb the customary alternate arrangement. Coalescence of the
vascular bundles in an unusual manner, and an irregular disposition of these
cords have also been considered to bring about deviations from the rule of
alternation, but in general the formation of the cords is subsequent to that of
the growing points or mamelons.
Page 592
Adhesions, accompanied by displacements, occasionally produce similar
deviations, the nature of which is usually easily detected.
Co-relation.—The importance of this subject first prominently brought into
notice by Geoffroy St. Hilaire gains in force daily. Rarely is a malformation
an isolated phenomenon, almost always it is associated, from the operations
of cause or effect, with some others. Instances of this co-relation have been
cited in the preceding pages, and many more might have been mentioned,
had the consideration of the relationship between form and function formed
part of the plan of this volume. A change in itself slight, often acquires
importance from its association with other alterations. This is particularly
well seen in the case of the receptacle. Let an ordinarily concave thalamus
remain, from defective development, flat, and how great the change in the
appearance of the flower. Let the usually contracted receptacle be
lengthened, and the whole aspect of the flowers so affected is altered to
such an extent that, were their history not known, botanists would have no
hesitation in assigning them to widely separate groups in their schemes of
classification. Peloria, too, of either form, affords excellent illustrations of
the co-existence of one changed condition with another. Not only is the
form of one set of organs altered, but the number, the relative proportion,
and the direction of the other organs of the flower are altered likewise.[562]
Not only is the whole symmetry changed, but the physiological operations
carried on in the flower undergo corresponding alterations.
There are certain co-relations which do not appear to have hitherto attracted
the attention they merit; such, for instance, is that which exists between the
particular period at which an organ is developed and its position and form.
In normal morphology this has, to some extent, been worked out, as in the
case of definite and indefinite, centrifugal and centripetal inflorescences,
and in the definite or indefinite formation of shoots, &c.
deviations, the nature of which is usually easily detected.
Co-relation.—The importance of this subject first prominently brought into
notice by Geoffroy St. Hilaire gains in force daily. Rarely is a malformation
an isolated phenomenon, almost always it is associated, from the operations
of cause or effect, with some others. Instances of this co-relation have been
cited in the preceding pages, and many more might have been mentioned,
had the consideration of the relationship between form and function formed
part of the plan of this volume. A change in itself slight, often acquires
importance from its association with other alterations. This is particularly
well seen in the case of the receptacle. Let an ordinarily concave thalamus
remain, from defective development, flat, and how great the change in the
appearance of the flower. Let the usually contracted receptacle be
lengthened, and the whole aspect of the flowers so affected is altered to
such an extent that, were their history not known, botanists would have no
hesitation in assigning them to widely separate groups in their schemes of
classification. Peloria, too, of either form, affords excellent illustrations of
the co-existence of one changed condition with another. Not only is the
form of one set of organs altered, but the number, the relative proportion,
and the direction of the other organs of the flower are altered likewise.[562]
Not only is the whole symmetry changed, but the physiological operations
carried on in the flower undergo corresponding alterations.
There are certain co-relations which do not appear to have hitherto attracted
the attention they merit; such, for instance, is that which exists between the
particular period at which an organ is developed and its position and form.
In normal morphology this has, to some extent, been worked out, as in the
case of definite and indefinite, centrifugal and centripetal inflorescences,
and in the definite or indefinite formation of shoots, &c.
Page 593
Other instances may be cited in the frequent co-existence of regular flowers
and definite inflorescence, the terminal position of many peloriated flowers,
the relationship between indefinite inflorescence and prolongation of the
axis, &c.
Again, the simultaneous evolution of the parts of the flower and their
consequent verticillate arrangement, are often associated with the
production of different forms from those characteristic of organs developed
in succession, and, in consequence, arranged spirally. In the case of
simultaneous development we meet with a repetition of whorls, as in what
are termed hose-in-hose flowers (flores duplicati, triplicati, &c.), and also
with cases of peloria. In instances where the organs are formed successively
in spiral order, we meet with such changes as median prolification,
petalody, and phyllody. All these are alterations which we might anticipate
from the activity of the growing point being checked at a certain stage in
the one case, while it is continuous in the other. This relationship between
the definite and indefinite modes of growth and the form of the several
organs of the flower, is more constant in reality than it may appear to be
from a perusal of the lists of genera in the foregoing pages, in which it was
not possible to show sufficiently well the comparative frequency of any
given changes in individual plants. Had it been possible to give statistics
setting forth the frequency of certain deviations in plants or groups having a
particular organisation, as compared with the rarity of their occurrence in
other plants of a different conformation, these co-relationships would have
been rendered much more evident. A hundred different plants, for instance,
may be named in any particular list, of which fifty shall be of one type of
structure, and the remainder of another. And the co-relative changes in each
fifty may appear to be evenly balanced, but so far is this from being the
case, that the frequency of the occurrence of a particular change, in one
species in the list, may be so great as far to exceed the instances of its
manifestation in all the rest put together. This difficulty is only very
and definite inflorescence, the terminal position of many peloriated flowers,
the relationship between indefinite inflorescence and prolongation of the
axis, &c.
Again, the simultaneous evolution of the parts of the flower and their
consequent verticillate arrangement, are often associated with the
production of different forms from those characteristic of organs developed
in succession, and, in consequence, arranged spirally. In the case of
simultaneous development we meet with a repetition of whorls, as in what
are termed hose-in-hose flowers (flores duplicati, triplicati, &c.), and also
with cases of peloria. In instances where the organs are formed successively
in spiral order, we meet with such changes as median prolification,
petalody, and phyllody. All these are alterations which we might anticipate
from the activity of the growing point being checked at a certain stage in
the one case, while it is continuous in the other. This relationship between
the definite and indefinite modes of growth and the form of the several
organs of the flower, is more constant in reality than it may appear to be
from a perusal of the lists of genera in the foregoing pages, in which it was
not possible to show sufficiently well the comparative frequency of any
given changes in individual plants. Had it been possible to give statistics
setting forth the frequency of certain deviations in plants or groups having a
particular organisation, as compared with the rarity of their occurrence in
other plants of a different conformation, these co-relationships would have
been rendered much more evident. A hundred different plants, for instance,
may be named in any particular list, of which fifty shall be of one type of
structure, and the remainder of another. And the co-relative changes in each
fifty may appear to be evenly balanced, but so far is this from being the
case, that the frequency of the occurrence of a particular change, in one
species in the list, may be so great as far to exceed the instances of its
manifestation in all the rest put together. This difficulty is only very
Page 594
partially obviated by the addition of the * to signify especial frequency of
occurrence of any given malformation in the plants to whose names it is
affixed.
Compensation.—But little further need be said on this head. An atrophied
condition of one part is generally associated with an hypertrophied
condition of another, and scarcely a change takes place in one direction, but
it is associated with an inverse alteration in some other. This principle is not
universal, and its application must not be unduly strained. It requires
specially to be considered in reference to differences in the degree or kind
of functional activity exercised by the organs implicated—points beyond
the scope of the present volume.
Teratology and classification.—Lastly, there remain to be mentioned the
bearings of teratology on systematic botany. There are those who would
entirely exclude teratology from such matters. It may be expedient to do so
when the object sought is one of convenience and facility of determination
only, but when broader considerations are concerned, teratology must no
more be banished than variation. In most instances the one differs but in
degree from the other. If variation affords aid in our speculations as to the
affinities and genealogical descent of species and other groups, so does
teratology, and in a far higher degree.
Take the characters of exogens as distinct from endogens; even under
ordinary circumstances, no absolute distinction can be drawn between them.
There are plants normally of an intermediate character, while, to take
exceptional instances, there are exogens with the leaves and flowers of
endogens, and endogens whose outward organisation, at any rate,
assimilates them to exogens. Diclinous or monochlamydeous plants owe
their imperfect conformation to suppression, and may become structurally
complete by a species of peloria. Structurally hermaphrodite flowers
occurrence of any given malformation in the plants to whose names it is
affixed.
Compensation.—But little further need be said on this head. An atrophied
condition of one part is generally associated with an hypertrophied
condition of another, and scarcely a change takes place in one direction, but
it is associated with an inverse alteration in some other. This principle is not
universal, and its application must not be unduly strained. It requires
specially to be considered in reference to differences in the degree or kind
of functional activity exercised by the organs implicated—points beyond
the scope of the present volume.
Teratology and classification.—Lastly, there remain to be mentioned the
bearings of teratology on systematic botany. There are those who would
entirely exclude teratology from such matters. It may be expedient to do so
when the object sought is one of convenience and facility of determination
only, but when broader considerations are concerned, teratology must no
more be banished than variation. In most instances the one differs but in
degree from the other. If variation affords aid in our speculations as to the
affinities and genealogical descent of species and other groups, so does
teratology, and in a far higher degree.
Take the characters of exogens as distinct from endogens; even under
ordinary circumstances, no absolute distinction can be drawn between them.
There are plants normally of an intermediate character, while, to take
exceptional instances, there are exogens with the leaves and flowers of
endogens, and endogens whose outward organisation, at any rate,
assimilates them to exogens. Diclinous or monochlamydeous plants owe
their imperfect conformation to suppression, and may become structurally
complete by a species of peloria. Structurally hermaphrodite flowers
Page 595
become unisexual by suppression, or are rendered incomplete by the non-
development of one or more of their floral whorls. Hypogynous flowers
become perigynous by adhesion, or by lack of separation; perigynous ones
become hypogynous by an early detachment from the receptacle that bears
them, or by the arrested development of an ordinarily cup-like receptacle.
How the relative position of the carpels and the calyx may be altered has
already been alluded to, as has also the circumstance that while it is
common to find an habitually inferior or adherent ovary becoming superior
or free, it is much more rare to find the superior ovary adherent to the
receptacle or to the calyx.[563] Regular and irregular peloria, too, serve to
show how slight are the boundaries, not only between different genera, but
also between different families.
While, therefore, teratology may be an unsafe guide in strictly artificial
schemes, it is obvious that its teachings should have great weight in all
philosophical systems of classification.
The questions will constantly arise, does such and such a form represent the
ancestral condition of certain plants? Is it a reversion to that form? or is it,
on the other hand, the starting point of new forms?
Such questions cannot receive at present any satisfactory answer, but the
evidence we have seems to indicate that pre-existing forms were simpler,
and less specialised in structure than those now existing, and hence if we
meet with malformations of a simple kind, we may consider them as
possible reversions; while, if they present features of increased complexity,
and more sharply defined differentiation, we may assume them to be
evidences of a progressive rather than of a retrogressive tendency.
That monstrosities so called may become the starting points of new forms is
proved by circumstance that, in many cases, the peculiarities are inherited
development of one or more of their floral whorls. Hypogynous flowers
become perigynous by adhesion, or by lack of separation; perigynous ones
become hypogynous by an early detachment from the receptacle that bears
them, or by the arrested development of an ordinarily cup-like receptacle.
How the relative position of the carpels and the calyx may be altered has
already been alluded to, as has also the circumstance that while it is
common to find an habitually inferior or adherent ovary becoming superior
or free, it is much more rare to find the superior ovary adherent to the
receptacle or to the calyx.[563] Regular and irregular peloria, too, serve to
show how slight are the boundaries, not only between different genera, but
also between different families.
While, therefore, teratology may be an unsafe guide in strictly artificial
schemes, it is obvious that its teachings should have great weight in all
philosophical systems of classification.
The questions will constantly arise, does such and such a form represent the
ancestral condition of certain plants? Is it a reversion to that form? or is it,
on the other hand, the starting point of new forms?
Such questions cannot receive at present any satisfactory answer, but the
evidence we have seems to indicate that pre-existing forms were simpler,
and less specialised in structure than those now existing, and hence if we
meet with malformations of a simple kind, we may consider them as
possible reversions; while, if they present features of increased complexity,
and more sharply defined differentiation, we may assume them to be
evidences of a progressive rather than of a retrogressive tendency.
That monstrosities so called may become the starting points of new forms is
proved by circumstance that, in many cases, the peculiarities are inherited
Page 596
so that a new "race" is produced and perpetuated: and if a new race, why
not a new species? The difference is one of degree only.
FOOTNOTES:
[553] See Clos., 'Bull. Soc. Bot. Fr.,' 1856, vol. iii, p. 679.
[554] 'Théorie de la Feuille,' p. 26.
[555] An additional illustration of this may be cited, which has been brought under the notice
of the writer by Dr. Welwitsch recently, and in which some of the leaflets of the pinnate leaf of
a species of Macrolobium were absent, and their place supplied by flowers arranged in cymes.
[556] The presence of a bud at the extremity once considered to be an absolute distinction
between branch and leaf, which latter never forms a bud exactly at the apex—is invalidated by
the case of the Nepaul barley, p. 174.
[557] 'Journ. Linn. Soc.,' vol. x, p. 103 et seq.
[558] See also the receptacular tube (ovary?) of Bæckea bearing stamens, see p. 183. It
would be natural to see stamens springing from the receptacle but not from the ovary.
[559] In Passiflora the organogeny of the flower clearly shows the truth of this assertion, as
was indeed shown by Payer and Schleiden.
[560] See Payer, 'Organ. Veget.'
[561] It must, however, be borne in mind that no true leaf-organ has yet been seen with a bud
at its exact apex (unless it be the nepaul barley), while in the case of an axial organ such a
position of the bud is constant. The nearest approach is in the case of impari-pinnate leaves in
which the terminal leaflet is jointed to the common rachis, and in the leaves of some Meliaceæ
which continue to push forth new leaflets even after the leaf has attained maturity.
[562] A singular instance of co-relation was shown by Mr. Saunders at the Scientific
Committee of the Royal Horticultural Society, February 16th, 1868, in a hyacinth with
perfectly green, long, tubular, erect, not horizontally spreading flowers.
[563] An illustration of this latter nature in the case of a cherry, which was surmounted by
the calyx lobes, precisely as in the case of a pomaceous fruit, has been given at p. 424, adnot.
not a new species? The difference is one of degree only.
FOOTNOTES:
[553] See Clos., 'Bull. Soc. Bot. Fr.,' 1856, vol. iii, p. 679.
[554] 'Théorie de la Feuille,' p. 26.
[555] An additional illustration of this may be cited, which has been brought under the notice
of the writer by Dr. Welwitsch recently, and in which some of the leaflets of the pinnate leaf of
a species of Macrolobium were absent, and their place supplied by flowers arranged in cymes.
[556] The presence of a bud at the extremity once considered to be an absolute distinction
between branch and leaf, which latter never forms a bud exactly at the apex—is invalidated by
the case of the Nepaul barley, p. 174.
[557] 'Journ. Linn. Soc.,' vol. x, p. 103 et seq.
[558] See also the receptacular tube (ovary?) of Bæckea bearing stamens, see p. 183. It
would be natural to see stamens springing from the receptacle but not from the ovary.
[559] In Passiflora the organogeny of the flower clearly shows the truth of this assertion, as
was indeed shown by Payer and Schleiden.
[560] See Payer, 'Organ. Veget.'
[561] It must, however, be borne in mind that no true leaf-organ has yet been seen with a bud
at its exact apex (unless it be the nepaul barley), while in the case of an axial organ such a
position of the bud is constant. The nearest approach is in the case of impari-pinnate leaves in
which the terminal leaflet is jointed to the common rachis, and in the leaves of some Meliaceæ
which continue to push forth new leaflets even after the leaf has attained maturity.
[562] A singular instance of co-relation was shown by Mr. Saunders at the Scientific
Committee of the Royal Horticultural Society, February 16th, 1868, in a hyacinth with
perfectly green, long, tubular, erect, not horizontally spreading flowers.
[563] An illustration of this latter nature in the case of a cherry, which was surmounted by
the calyx lobes, precisely as in the case of a pomaceous fruit, has been given at p. 424, adnot.
Page 597
APPENDIX
DOUBLE FLOWERS.[564]
In ordinary language, the epithet double flowers is applied to flowers of
very varied structural conformation. The most common conditions
rendering a flower double, in the popular acceptation of the term, are
substitutions of petals or petal-like bodies for stamens and pistils, one or
both. (See Petalody, p. 283.) Another very common mode of doubling is
brought about by a real or apparent augmentation in the number of petals,
as by multiplication, fission, or chorisis. (See pp. 66, 343, 371, 376.)
Sometimes even the receptacle of the flower within the outer corolla,
divides, each subdivision becoming the centre of a new series of petals, as
in some very luxuriant camellias and anemones. The isolation of organs
which, under ordinary circumstances, are united together, is another
circumstance, giving rise, in popular parlance, to the use of the term double
flower. (See Adesmy, Solution, pp. 58, 76, 82.) Prolification is another very
frequent occurrence in the case of these flowers, while still other forms
arise from laciniation of the petals, or from the formation of excrescences
from the petals or stamens, in the form of supplementary petal-like lobes.
(See Enation, p. 443.)
As these matters are all treated of under their respective headings, it is not
necessary to allude to them again in detail. It may be well, however, to
DOUBLE FLOWERS.[564]
In ordinary language, the epithet double flowers is applied to flowers of
very varied structural conformation. The most common conditions
rendering a flower double, in the popular acceptation of the term, are
substitutions of petals or petal-like bodies for stamens and pistils, one or
both. (See Petalody, p. 283.) Another very common mode of doubling is
brought about by a real or apparent augmentation in the number of petals,
as by multiplication, fission, or chorisis. (See pp. 66, 343, 371, 376.)
Sometimes even the receptacle of the flower within the outer corolla,
divides, each subdivision becoming the centre of a new series of petals, as
in some very luxuriant camellias and anemones. The isolation of organs
which, under ordinary circumstances, are united together, is another
circumstance, giving rise, in popular parlance, to the use of the term double
flower. (See Adesmy, Solution, pp. 58, 76, 82.) Prolification is another very
frequent occurrence in the case of these flowers, while still other forms
arise from laciniation of the petals, or from the formation of excrescences
from the petals or stamens, in the form of supplementary petal-like lobes.
(See Enation, p. 443.)
As these matters are all treated of under their respective headings, it is not
necessary to allude to them again in detail. It may be well, however, to
Page 598
allude, in general terms, to the causes which have been assigned by various
writers for their formation, and to the means which have been adopted by
practical experimenters to secure the production of the flowers often so
much esteemed by the florist. It must be admitted that, in spite of all that
has been written on the subject, but very little is known about these matters.
In the case of the stock the following means have been adopted by
cultivators in order to obtain plants bearing double instead of single
flowers. There is first the crossing of single flowers with double ones,
effected by planting a double-flowered plant in proximity to a single-
flowered one; but this, it is obvious, could lead to no important results,
since the double flowers, having no pollen, could not possibly influence the
seed, which is borne only by the single-flowered plants. Another plan is the
degustation of the buds, that is to say, the chewing of the well-formed buds;
it is held that the single plants can be recognised by their sweeter taste and
greater consistence, and may thus be weeded out; but there is at least the
disadvantage attending this method, that the plants, single as well as double,
must all be grown up to the period when these buds are tolerably well
advanced. A third method which has been adopted is, that of sowing the
seeds at a particular lunar epoch, great confidence being placed in the plan
of planting them during the last quarter of the moon, but such confidence is
found to be misplaced. The plan of removing the stamens has had its
supporters, but as this must be done at an early stage of development, and
could only influence the result by diverting the vital force which would be
expended in the maturation of the pollen, to the perfecting of the seeds, it is
obvious that the plan is impracticable for all ordinary purposes, even if in
any degree efficient, which from the plasticity of vegetable development,
and the faculty of doubling which is inherent in the stock family, is not at
all improbable. Still another mark, the presence of a fifth petal in the single
or seed-bearing flower, has been held to indicate the assurance of obtaining
a crop of double-flowered plants from seeds saved from flowers possessing
writers for their formation, and to the means which have been adopted by
practical experimenters to secure the production of the flowers often so
much esteemed by the florist. It must be admitted that, in spite of all that
has been written on the subject, but very little is known about these matters.
In the case of the stock the following means have been adopted by
cultivators in order to obtain plants bearing double instead of single
flowers. There is first the crossing of single flowers with double ones,
effected by planting a double-flowered plant in proximity to a single-
flowered one; but this, it is obvious, could lead to no important results,
since the double flowers, having no pollen, could not possibly influence the
seed, which is borne only by the single-flowered plants. Another plan is the
degustation of the buds, that is to say, the chewing of the well-formed buds;
it is held that the single plants can be recognised by their sweeter taste and
greater consistence, and may thus be weeded out; but there is at least the
disadvantage attending this method, that the plants, single as well as double,
must all be grown up to the period when these buds are tolerably well
advanced. A third method which has been adopted is, that of sowing the
seeds at a particular lunar epoch, great confidence being placed in the plan
of planting them during the last quarter of the moon, but such confidence is
found to be misplaced. The plan of removing the stamens has had its
supporters, but as this must be done at an early stage of development, and
could only influence the result by diverting the vital force which would be
expended in the maturation of the pollen, to the perfecting of the seeds, it is
obvious that the plan is impracticable for all ordinary purposes, even if in
any degree efficient, which from the plasticity of vegetable development,
and the faculty of doubling which is inherent in the stock family, is not at
all improbable. Still another mark, the presence of a fifth petal in the single
or seed-bearing flower, has been held to indicate the assurance of obtaining
a crop of double-flowered plants from seeds saved from flowers possessing
Page 599
this peculiarity. To a certain extent, doubtless, this expectation would be
realised, owing to the plasticity and inherent quality just alluded to, but the
proportion would be too small for any useful practical purpose.
"The gardeners of Erfurt," observes M. Chaté, who has written a book[565]
on the subject, in which he makes known a means of obtaining double-
flowered stocks founded on more than fifty years' practice in his family,
"have, for a long time, to a certain extent monopolised the sale of seeds of
these plants. To obtain these seeds, the Erfurt gardeners cultivate the
flowers in pots, and place them on shelves in large greenhouses, giving
them only sufficient water to prevent them from dying. So cultivated the
plants become weakened, the pods shortened, and the seeds less numerous,
and better ripened; and these seeds give from 60 to 70 per cent. of double
flowers.
"The seeds from these plants are said to be mostly of an abnormal shape,
which is so striking that experienced cultivators are able to separate those
which would furnish double flowers from those which would produce
single ones."
M. Chaté's method, which he calls the French one, gives still greater results,
viz.: 80 per cent. of double flowers, and these produced by very simple
means. "When my seeds," he observes, "have been chosen with care, I plant
them, in the month of April, in good dry mould, in a position exposed to the
morning sun, this position being the most favourable. At the time of
flowering I nip off some of the flowering branches, and leave only ten or
twelve pods on the secondary branches, taking care to remove all the small
weak branches which shoot at this time. I leave none but the principal and
the secondary branches to bear the pods. All the sap is employed in
nourishing the seeds thus borne, which give a result of 80 per cent. of
double flowers. The pods under this management are thicker, and their
realised, owing to the plasticity and inherent quality just alluded to, but the
proportion would be too small for any useful practical purpose.
"The gardeners of Erfurt," observes M. Chaté, who has written a book[565]
on the subject, in which he makes known a means of obtaining double-
flowered stocks founded on more than fifty years' practice in his family,
"have, for a long time, to a certain extent monopolised the sale of seeds of
these plants. To obtain these seeds, the Erfurt gardeners cultivate the
flowers in pots, and place them on shelves in large greenhouses, giving
them only sufficient water to prevent them from dying. So cultivated the
plants become weakened, the pods shortened, and the seeds less numerous,
and better ripened; and these seeds give from 60 to 70 per cent. of double
flowers.
"The seeds from these plants are said to be mostly of an abnormal shape,
which is so striking that experienced cultivators are able to separate those
which would furnish double flowers from those which would produce
single ones."
M. Chaté's method, which he calls the French one, gives still greater results,
viz.: 80 per cent. of double flowers, and these produced by very simple
means. "When my seeds," he observes, "have been chosen with care, I plant
them, in the month of April, in good dry mould, in a position exposed to the
morning sun, this position being the most favourable. At the time of
flowering I nip off some of the flowering branches, and leave only ten or
twelve pods on the secondary branches, taking care to remove all the small
weak branches which shoot at this time. I leave none but the principal and
the secondary branches to bear the pods. All the sap is employed in
nourishing the seeds thus borne, which give a result of 80 per cent. of
double flowers. The pods under this management are thicker, and their
Page 600
maturation is more perfect. At the time of extracting the seeds the upper
portion of the pod is separated and placed aside, because it has been
ascertained that the plants coming from the seeds situated in this portion of
the pod, give 80 per cent. of single flowers. They yield, however, greater
variety than the others. This plan of suppressing that part of the pod which
yields single flowers in the largest proportion, greatly facilitates the
recognition of the single-flowered plants, because there remains to be
eliminated from among the seedlings only from 10 to 15 per cent.
This separation of the single from the double-flowered plants, M. Chaté
tells us is not so difficult as might be supposed. The single stocks, he
explains, have deep green leaves (glabrous in certain species), rounded at
the top, the heart being in the form of a shuttlecock, and the plant stout and
thickset in its general aspect, while the plants yielding double flowers have
very long leaves of a light green colour, hairy, and curled at the edges, the
heart consisting of whitish leaves, curved so that they enclose it completely.
Such is the substance of M. Chaté's method of securing so large a
proportion of double-flowered plants, and then of separating them from the
remaining single ones—a method which commends itself to the good sense
of the intelligent cultivator."[566]
Signor Rigamonti, a great cultivator of pinks, asserted that he was able to
distinguish double from single-flowered pinks, in the seedling state.
According to this gentleman, those seedlings which produce three
cotyledons in a whorl in place of two, form double flowers. In the case of
Primula sinensis the same results occurred. Some had three leaves in a ring,
others two; most had the leaves standing one over the other as usual. These
were divided into three sets, and when they flowered, the first lot were all
double, the second semi-double, the third single. But these statements have
not been confirmed by other observers; and the writer can safely assert that
seedling pinks occasionally produce three cotyledons, and subsequently
portion of the pod is separated and placed aside, because it has been
ascertained that the plants coming from the seeds situated in this portion of
the pod, give 80 per cent. of single flowers. They yield, however, greater
variety than the others. This plan of suppressing that part of the pod which
yields single flowers in the largest proportion, greatly facilitates the
recognition of the single-flowered plants, because there remains to be
eliminated from among the seedlings only from 10 to 15 per cent.
This separation of the single from the double-flowered plants, M. Chaté
tells us is not so difficult as might be supposed. The single stocks, he
explains, have deep green leaves (glabrous in certain species), rounded at
the top, the heart being in the form of a shuttlecock, and the plant stout and
thickset in its general aspect, while the plants yielding double flowers have
very long leaves of a light green colour, hairy, and curled at the edges, the
heart consisting of whitish leaves, curved so that they enclose it completely.
Such is the substance of M. Chaté's method of securing so large a
proportion of double-flowered plants, and then of separating them from the
remaining single ones—a method which commends itself to the good sense
of the intelligent cultivator."[566]
Signor Rigamonti, a great cultivator of pinks, asserted that he was able to
distinguish double from single-flowered pinks, in the seedling state.
According to this gentleman, those seedlings which produce three
cotyledons in a whorl in place of two, form double flowers. In the case of
Primula sinensis the same results occurred. Some had three leaves in a ring,
others two; most had the leaves standing one over the other as usual. These
were divided into three sets, and when they flowered, the first lot were all
double, the second semi-double, the third single. But these statements have
not been confirmed by other observers; and the writer can safely assert that
seedling pinks occasionally produce three cotyledons, and subsequently
Page 601
single flowers. He has never observed a double flower under these
circumstances, though it is true his experience in this matter has been but
small.
A writer in Otto's 'Gartenzeitung,' considers that double flowers are a
consequence of dryness of soil and atmosphere, and not of a luxurious soil,
rich in nutritious matter, having arrived at this conclusion from an
observation of the following circumstances:
"Fifty years ago we saw Kerria japonica in a hothouse with single flowers.
Twenty years later we met with it in several gardens, in the open air, but
always with double flowers. At this time we were assured that single-
flowered plants were no more to be found in the whole of Europe, and
botanists forming herbaria offered considerable sums for a branch of K.
japonica with single flowers. We were requested to take the plant in hand
for the purpose of inducing it to produce single flowers. We were advised to
plant it out in a rich soil, which was done, but, by chance, the situation was
sloping, consequently it did not retain moisture, and all the flowers
produced for several years in succession were double. Shortly after, the
captain of an English ship again brought plants bearing normal flowers
from Japan, which were soon spread over the continent, and of which we
received one plant. After three years all the young plants raised from
cuttings were double-flowered.
"In the year 1820 we several times visited a garden in the neighbourhood of
Vienna, well known on account of its plant culture. The gardener there
possessed an immense plant of Camellia japonica with single flowers, and
some small plants raised from this by cuttings, but no other variety of
camellia. He fertilised the flowers with their own pollen, harvested seeds,
which he sowed, and the plants raised from them were placed in an
extremely dry, lofty conservatory, where, after some years, instead of
circumstances, though it is true his experience in this matter has been but
small.
A writer in Otto's 'Gartenzeitung,' considers that double flowers are a
consequence of dryness of soil and atmosphere, and not of a luxurious soil,
rich in nutritious matter, having arrived at this conclusion from an
observation of the following circumstances:
"Fifty years ago we saw Kerria japonica in a hothouse with single flowers.
Twenty years later we met with it in several gardens, in the open air, but
always with double flowers. At this time we were assured that single-
flowered plants were no more to be found in the whole of Europe, and
botanists forming herbaria offered considerable sums for a branch of K.
japonica with single flowers. We were requested to take the plant in hand
for the purpose of inducing it to produce single flowers. We were advised to
plant it out in a rich soil, which was done, but, by chance, the situation was
sloping, consequently it did not retain moisture, and all the flowers
produced for several years in succession were double. Shortly after, the
captain of an English ship again brought plants bearing normal flowers
from Japan, which were soon spread over the continent, and of which we
received one plant. After three years all the young plants raised from
cuttings were double-flowered.
"In the year 1820 we several times visited a garden in the neighbourhood of
Vienna, well known on account of its plant culture. The gardener there
possessed an immense plant of Camellia japonica with single flowers, and
some small plants raised from this by cuttings, but no other variety of
camellia. He fertilised the flowers with their own pollen, harvested seeds,
which he sowed, and the plants raised from them were placed in an
extremely dry, lofty conservatory, where, after some years, instead of
Page 602
producing single flowers, they all produced double ones. The seedlings and
mother plant were planted in one and the same kind of earth, and some of
the flowers on the old plant also showed an inclination to become double.
"This, at that time, to us, enigmatical phenomenon, was kept in mind until
we had an opportunity of instituting comparisons between the climate of
Japan and China and our own, and we then concluded that in the case of a
plant imported from thence, and exposed to such different climatical
influences, the origin of the greater or less imperfection of its sexual organs
was probably owing to this change, as we had experienced in Kerria and
Camellia; and that the sterility of many other exotic plants might be
attributed to the same cause. The difference in the climatical relations of
Japan and Europe is very considerable. In Japan, previous to the new
growth of Kerria and Camellia, a rainy season of three months' duration
prevails; in Europe, on the contrary, dry winds prevail especially in the
eastern part, where our plains are often transformed into deserts. Is it,
therefore, remarkable that a plant introduced from Japan into Europe,
exposed to the influences of this great diversity of climate, should produce
imperfect sexual organs incapable of further propagating the plant from
seeds? A rich soil, with the necessary amount of moisture, will never
engender double flowers."[567]
Mr. Darwin[568] describes a peculiar form of Gentiana Amarella, in which
the parts of the flower were more or less replaced by compact aggregations
of purple scales in great numbers. A similar condition is, indeed, not
uncommon in this plant, and, as Mr. Darwin also remarked, on hard, dry,
bare, chalky banks, thus bearing out the views expressed by the writer in the
'Gartenzeitung' just cited. Some double flowers of Potentilla reptans found
growing wild near York, and transmitted to the writer by a correspondent,
were observed growing along a high wall, in a dry border, close to a beaten
mother plant were planted in one and the same kind of earth, and some of
the flowers on the old plant also showed an inclination to become double.
"This, at that time, to us, enigmatical phenomenon, was kept in mind until
we had an opportunity of instituting comparisons between the climate of
Japan and China and our own, and we then concluded that in the case of a
plant imported from thence, and exposed to such different climatical
influences, the origin of the greater or less imperfection of its sexual organs
was probably owing to this change, as we had experienced in Kerria and
Camellia; and that the sterility of many other exotic plants might be
attributed to the same cause. The difference in the climatical relations of
Japan and Europe is very considerable. In Japan, previous to the new
growth of Kerria and Camellia, a rainy season of three months' duration
prevails; in Europe, on the contrary, dry winds prevail especially in the
eastern part, where our plains are often transformed into deserts. Is it,
therefore, remarkable that a plant introduced from Japan into Europe,
exposed to the influences of this great diversity of climate, should produce
imperfect sexual organs incapable of further propagating the plant from
seeds? A rich soil, with the necessary amount of moisture, will never
engender double flowers."[567]
Mr. Darwin[568] describes a peculiar form of Gentiana Amarella, in which
the parts of the flower were more or less replaced by compact aggregations
of purple scales in great numbers. A similar condition is, indeed, not
uncommon in this plant, and, as Mr. Darwin also remarked, on hard, dry,
bare, chalky banks, thus bearing out the views expressed by the writer in the
'Gartenzeitung' just cited. Some double flowers of Potentilla reptans found
growing wild near York, and transmitted to the writer by a correspondent,
were observed growing along a high wall, in a dry border, close to a beaten
Page 603
path, bordering on a gravel pit, others were found on a raised bank, which,
from its elevation and exposure to the sun, was particularly dry.
On the other hand, the double-flowered Cardamine pratensis, which is
occasionally found in a wild state, always grows in very wet places.
Of late years a remarkable double-flowered race of Primula sinensis has
been obtained. In particular, Messrs. Windebank and Kingsbury, of
Southampton, have succeeded in raising a set of plants in which the flowers
are very double and very attractive in a florist's point of view. The corollas
in these flowers are not merely duplicated, but from their inner surface
spring, in some cases, funnel-shaped or tubular petals (p. 315), so regular in
form as quite to resemble a perfect corolla. These tubes are attached to the
inner side of the tube of the corolla, in the same way as are the stamens,
these latter organs being, it appears, absent. The carpels are present, but
open at the top, and bear numerous ovules, hence it was at first surmised
that these plants were obtained and perpetuated, by the application of pollen
from single flowers to these double-flowered varieties.
The raisers of this fine race however assert that "the double kinds are all
raised from the seed obtained from single flowers; the double blooms do
not produce seed, as a rule, and even if they did yield seed, and it were to
germinate, the plants so raised would simply produce single flowers." Semi-
double flowers will produce seed, but it is necessary that they should be
fertilised with the pollen from the single blooms. They rarely, however, if
ever, produce really double flowers when so fertilised, and the number of
semi-double flowers, even, is always small, the remainder, and,
consequently, the larger part, proving single. To obtain double varieties, the
raiser fertilises certain fine and striking single flowers, with the pollen of
other equally fine single blooms, and the desired result is obtained. This is
from its elevation and exposure to the sun, was particularly dry.
On the other hand, the double-flowered Cardamine pratensis, which is
occasionally found in a wild state, always grows in very wet places.
Of late years a remarkable double-flowered race of Primula sinensis has
been obtained. In particular, Messrs. Windebank and Kingsbury, of
Southampton, have succeeded in raising a set of plants in which the flowers
are very double and very attractive in a florist's point of view. The corollas
in these flowers are not merely duplicated, but from their inner surface
spring, in some cases, funnel-shaped or tubular petals (p. 315), so regular in
form as quite to resemble a perfect corolla. These tubes are attached to the
inner side of the tube of the corolla, in the same way as are the stamens,
these latter organs being, it appears, absent. The carpels are present, but
open at the top, and bear numerous ovules, hence it was at first surmised
that these plants were obtained and perpetuated, by the application of pollen
from single flowers to these double-flowered varieties.
The raisers of this fine race however assert that "the double kinds are all
raised from the seed obtained from single flowers; the double blooms do
not produce seed, as a rule, and even if they did yield seed, and it were to
germinate, the plants so raised would simply produce single flowers." Semi-
double flowers will produce seed, but it is necessary that they should be
fertilised with the pollen from the single blooms. They rarely, however, if
ever, produce really double flowers when so fertilised, and the number of
semi-double flowers, even, is always small, the remainder, and,
consequently, the larger part, proving single. To obtain double varieties, the
raiser fertilises certain fine and striking single flowers, with the pollen of
other equally fine single blooms, and the desired result is obtained. This is
Page 604
Messrs. Windebank and Kingsbury's modus operandi, the exact process or
mode of accomplishment being, however, a professional secret.[569]
From what has been said, as well as from other evidence which it is not
necessary to detail in this place, it may be seen that the causes assigned by
physiologists, and the plans proposed by cultivators for the production of
double flowers, are reducible to three heads, which may be classed under
Plethora, Starvation, and Sterility. These three seem inconsistent one with
the other, but are not so much so as they at first sight appear to be.
Tho advocates of the plethora theory have much in their favour: for
instance, the greater frequency of double flowers among cultivated plants
than among wild ones. The great preponderance of double flowers in plants
derived from the northern hemisphere, when contrasted with those procured
from the southern, as alluded to by Dr. Seemann, seems also to point to the
effect of cultivation in producing these flowers. Now, although this is, to a
large extent, due to the selection that has been for so long a period practised
by gardeners, still that process will not account for the appearance of double
flowers where no such selection has been exercised; as in the case of wild
plants. Some double peas, observed by Mr. Laxton, appeared suddenly; they
had not been selected or sought for, but they were produced, as it would
appear, as a result of high cultivation, and during the period when the plant
was in greatest vigour; and as the energies of the plant failed, so the
tendency to produce double flowers ceased. Indeed, in reference to this
subject, it is always important to bear in mind the time at which double
flowers are produced; thus, an annual plant subjected to cultivation, will, it
may be, produce single flowers for the firet year or two, then a few partially
double flowers are formed, and from these, by careful selection and
breeding, a double-flowered race may be secured. Sometimes, as in the peas
before alluded to, in the same season the earlier blossoms are single, while
later in the year double blossoms are produced. This happens, not only in
mode of accomplishment being, however, a professional secret.[569]
From what has been said, as well as from other evidence which it is not
necessary to detail in this place, it may be seen that the causes assigned by
physiologists, and the plans proposed by cultivators for the production of
double flowers, are reducible to three heads, which may be classed under
Plethora, Starvation, and Sterility. These three seem inconsistent one with
the other, but are not so much so as they at first sight appear to be.
Tho advocates of the plethora theory have much in their favour: for
instance, the greater frequency of double flowers among cultivated plants
than among wild ones. The great preponderance of double flowers in plants
derived from the northern hemisphere, when contrasted with those procured
from the southern, as alluded to by Dr. Seemann, seems also to point to the
effect of cultivation in producing these flowers. Now, although this is, to a
large extent, due to the selection that has been for so long a period practised
by gardeners, still that process will not account for the appearance of double
flowers where no such selection has been exercised; as in the case of wild
plants. Some double peas, observed by Mr. Laxton, appeared suddenly; they
had not been selected or sought for, but they were produced, as it would
appear, as a result of high cultivation, and during the period when the plant
was in greatest vigour; and as the energies of the plant failed, so the
tendency to produce double flowers ceased. Indeed, in reference to this
subject, it is always important to bear in mind the time at which double
flowers are produced; thus, an annual plant subjected to cultivation, will, it
may be, produce single flowers for the firet year or two, then a few partially
double flowers are formed, and from these, by careful selection and
breeding, a double-flowered race may be secured. Sometimes, as in the peas
before alluded to, in the same season the earlier blossoms are single, while
later in the year double blossoms are produced. This happens, not only in
Page 605
annuals, but also in perennials, and is not infrequent in the apple; an
illustration of this occurrence in this tree is given in the 'Gardeners'
Chronicle' for 1865, p. 554.[570] Sometimes the flowers on a particular
branch are double, while those on the rest of the plant are single.[571] On
these points, the evidence furnished by a double white hawthorn in the
Royal Botanic Gardens at Edinburgh is important. Professor Balfour kindly
wrote as follows in reply to an inquiry respecting this plant:—"A double
white hawthorn in the Royal Botanic Gardens produced double flowers in
spring. It retained its leaves during autumn and winter, until the following
spring. It then flowered in the second spring, but produced weak single
flowers only, and has continued to do so ever since. The flowering has been
always weak, since this change of flowers from double to single. Mr. M'Nab
attributes the change in the duration of the leaves to the filling up of the
ground round the tree, to the height of a foot and a half on the stem. He is
now trying the effect of extra manure in giving extra vigour to the plant."
Here, at least, the production of single flowers would seem to be the result
of debilitating causes, connected with the unusual persistence of the leaves,
&c., for while the tree was healthy, double flowers were produced.
A similar illustration came under the writer's own notice. Some seedling
balsams, of a strain which from long selection and hereditary tendency
produces, year after year, double flowers were, in the spring (of 1866),
allowed to remain in the seed-pans for many weeks after they were ready to
be potted off; they were hence partly starved, and when they bloomed, they
produced single flowers only. But these same plants, when more liberally
treated, produced an abundance of double flowers. Moreover, other
seedlings of the same batch, but sown later, and potted off at the usual time,
produced double flowers as usual. Of a like character is the fact that the
double Ranunculus asiaticus loses its doubleness if the roots are planted in
a poor soil.
illustration of this occurrence in this tree is given in the 'Gardeners'
Chronicle' for 1865, p. 554.[570] Sometimes the flowers on a particular
branch are double, while those on the rest of the plant are single.[571] On
these points, the evidence furnished by a double white hawthorn in the
Royal Botanic Gardens at Edinburgh is important. Professor Balfour kindly
wrote as follows in reply to an inquiry respecting this plant:—"A double
white hawthorn in the Royal Botanic Gardens produced double flowers in
spring. It retained its leaves during autumn and winter, until the following
spring. It then flowered in the second spring, but produced weak single
flowers only, and has continued to do so ever since. The flowering has been
always weak, since this change of flowers from double to single. Mr. M'Nab
attributes the change in the duration of the leaves to the filling up of the
ground round the tree, to the height of a foot and a half on the stem. He is
now trying the effect of extra manure in giving extra vigour to the plant."
Here, at least, the production of single flowers would seem to be the result
of debilitating causes, connected with the unusual persistence of the leaves,
&c., for while the tree was healthy, double flowers were produced.
A similar illustration came under the writer's own notice. Some seedling
balsams, of a strain which from long selection and hereditary tendency
produces, year after year, double flowers were, in the spring (of 1866),
allowed to remain in the seed-pans for many weeks after they were ready to
be potted off; they were hence partly starved, and when they bloomed, they
produced single flowers only. But these same plants, when more liberally
treated, produced an abundance of double flowers. Moreover, other
seedlings of the same batch, but sown later, and potted off at the usual time,
produced double flowers as usual. Of a like character is the fact that the
double Ranunculus asiaticus loses its doubleness if the roots are planted in
a poor soil.
Page 606
On the other hand, the way in which double stocks are stated to be produced
at Erfurt, viz.: by giving the plants a minimum supply of water, and the
other circumstances alluded to as showing the connection between the
production of double flowers, and a deficiency of water, as well as the
experiments of Mr. Monro, go to show that, so far from plethora, the
inducing cause must be more nearly allied to inanition, though the
impoverishing process is, to a certain extent, counteracted by only allowing
a few of the seed-pods to ripen, and thus concentrating in a small number of
flowers the nutriment intended for many.
Professor Edward Morren ('Bull. Acad. Roy. Belg.,' 2me ser., vol. xix, p.
224) considers the existence of true variegation in leaves, and the
production of double flowers, as antagonistic one to the other; the former is
a sign of weakness, the latter of strength. But it would seem that the
exceptions are so numerous—so many cases of the co-existence of
variegated leaves, and double flowers are known, at least in individual
plants if not in species—that no safe inferences can be drawn as to this
point. Since the above remarks were printed, Professor Morren has
published a second paper on the subject, upholding his former views as to
the incompatibility of variegated foliage (not mere colouration) and double
flowers. In this paper he criticises the objections raised by the present writer
and others, and examines some of the alleged exceptions. Some of these the
Belgian savant finds to prove his rule, inasmuch as although there is a co-
existence of variegated foliage and double flowers in these illustrations, yet
the plants are weakly, the flowers ill formed, or fall off before expansion.
Admitting all this, there still remain cases in which double flowers and
variegated foliage do exist in conjunction, and where the plants are
vigorous and the flowers well developed. Instances of this are known to
cultivators in species of Dianthus, Hemerocallis, Althæa, Pæonia, Rosa,
Ranunculus, Serissa, Saponaria, etc., and probably the art of the cultivator
would speedily be successful in raising other examples, were it a matter of
at Erfurt, viz.: by giving the plants a minimum supply of water, and the
other circumstances alluded to as showing the connection between the
production of double flowers, and a deficiency of water, as well as the
experiments of Mr. Monro, go to show that, so far from plethora, the
inducing cause must be more nearly allied to inanition, though the
impoverishing process is, to a certain extent, counteracted by only allowing
a few of the seed-pods to ripen, and thus concentrating in a small number of
flowers the nutriment intended for many.
Professor Edward Morren ('Bull. Acad. Roy. Belg.,' 2me ser., vol. xix, p.
224) considers the existence of true variegation in leaves, and the
production of double flowers, as antagonistic one to the other; the former is
a sign of weakness, the latter of strength. But it would seem that the
exceptions are so numerous—so many cases of the co-existence of
variegated leaves, and double flowers are known, at least in individual
plants if not in species—that no safe inferences can be drawn as to this
point. Since the above remarks were printed, Professor Morren has
published a second paper on the subject, upholding his former views as to
the incompatibility of variegated foliage (not mere colouration) and double
flowers. In this paper he criticises the objections raised by the present writer
and others, and examines some of the alleged exceptions. Some of these the
Belgian savant finds to prove his rule, inasmuch as although there is a co-
existence of variegated foliage and double flowers in these illustrations, yet
the plants are weakly, the flowers ill formed, or fall off before expansion.
Admitting all this, there still remain cases in which double flowers and
variegated foliage do exist in conjunction, and where the plants are
vigorous and the flowers well developed. Instances of this are known to
cultivators in species of Dianthus, Hemerocallis, Althæa, Pæonia, Rosa,
Ranunculus, Serissa, Saponaria, etc., and probably the art of the cultivator
would speedily be successful in raising other examples, were it a matter of
Page 607
importance or interest to them to do so. At any rate, the existence of a few
unimpeachable illustrations is sufficient to support the opinion of the
present writer, and objected to so strongly by M. Morren that, in the present
state of our knowledge, "no safe inferences can be drawn" from the facts
alluded to by the Belgian professor.[572]
Mr. Darwin[573] has thrown out the suggestion that the cause for the
appearance of double flowers may be sought for in some previous state of
things, bringing about sterility or imperfect formation, or functional activity
of the genitalia of the flower, and consequent compensatory increase of the
petaline element, either in the form of an increased number of bracts, petals,
&c., or in the substitution of petals for stamens and pistils, &c.
In considering these points the question arises whether they can be
reconciled one with another. And there is little doubt but that they may be.
The production of a flower is preceded by an arrest of vegetation; this is
obvious: the current of the plant's life becomes changed, the growth of the
leaves is checked, the lengthening of the branches is arrested as the flower-
bud forms; moreover, there is a close relationship in a large majority of
flowers between the outer envelopes of the flower and the scales of a leaf-
bud; this is especially so in regard to the venation, and is admitted by all
morphologists. So far, then, it may be said that the production of a flower,
like that of a bud, is due to a diminution of vegetative action; and as in
double flowers we have, for the most part, merely a repetition and
exuberant formation of floral envelopes, so we may attribute their formation
to a continuance of the same feeble vegetative action as that which
produced the first or normal series. How, then, can a copious supply of rich
food, such as is provided by cultivation, produce double flowers? To this
question, according to our theory, the reply would be that the quantity of
food is excessive, more than the plant can properly digest; and hence
vegetative action is stopped, at least partially—pretty much as it would be if
unimpeachable illustrations is sufficient to support the opinion of the
present writer, and objected to so strongly by M. Morren that, in the present
state of our knowledge, "no safe inferences can be drawn" from the facts
alluded to by the Belgian professor.[572]
Mr. Darwin[573] has thrown out the suggestion that the cause for the
appearance of double flowers may be sought for in some previous state of
things, bringing about sterility or imperfect formation, or functional activity
of the genitalia of the flower, and consequent compensatory increase of the
petaline element, either in the form of an increased number of bracts, petals,
&c., or in the substitution of petals for stamens and pistils, &c.
In considering these points the question arises whether they can be
reconciled one with another. And there is little doubt but that they may be.
The production of a flower is preceded by an arrest of vegetation; this is
obvious: the current of the plant's life becomes changed, the growth of the
leaves is checked, the lengthening of the branches is arrested as the flower-
bud forms; moreover, there is a close relationship in a large majority of
flowers between the outer envelopes of the flower and the scales of a leaf-
bud; this is especially so in regard to the venation, and is admitted by all
morphologists. So far, then, it may be said that the production of a flower,
like that of a bud, is due to a diminution of vegetative action; and as in
double flowers we have, for the most part, merely a repetition and
exuberant formation of floral envelopes, so we may attribute their formation
to a continuance of the same feeble vegetative action as that which
produced the first or normal series. How, then, can a copious supply of rich
food, such as is provided by cultivation, produce double flowers? To this
question, according to our theory, the reply would be that the quantity of
food is excessive, more than the plant can properly digest; and hence
vegetative action is stopped, at least partially—pretty much as it would be if
Page 608
the plant were placed in the opposite condition of starvation. The effect of
supplying a plant (or an animal) with an excessive supply of food, which it
cannot assimilate, is in many respects similar to that which results from
partially cutting off the supplies. And the same reasoning applies to sterility.
If by high culture, or the supply of an undue quantity of nourishment, the
constitution of the plant be impaired, or if the plant be pampered, it is no
wonderful thing that sterility should ensue. Hence, then, may it not be
asserted as a general principle that in the production of double flowers a
partial arrest of development, if not of growth, however produced, is an
essential preliminary? All the attendant phenomena, such as the obliteration
of the stamens, the augmentation in the number of floral whorls, the
occurrence of prolification, are consistent with the supposition of a primary
arrest of development, more or less complete, as the case may be: at one
time permanent, at another time relaxed and intermittent, or in a third set of
cases the vegetative activity or power of growth may be restored, and from
the centre of the flower may spring a perfect branch with perfect leaves, the
production of sheaths only being superseded by the development of leaves,
in which all the parts—sheath, stalk, and blade—are present.
When once the disposition to form double flowers is established, that
tendency becomes hereditary: there are races of single Stocks in which, out
of hundreds of plants, scarcely one double-flowered form is met with; but
when the tendency to produce double blooms is set up, single flowers
become the exception: thus, in the Balsams, before mentioned, not one in
fifty now produces single flowers, and the seeds of these double Balsams
produce double-flowered seedlings, with scarcely a "rogue" among them.
The following list of plants producing double flowers of any kind is taken
from that given in 'Seemann's Journal of Botany,' vol. ii, p. 177, and to
which some additions have been made. Miscalled double flowers, such as
those of the Compositæ, Viburnum Hydrangea, &c., are excluded.
supplying a plant (or an animal) with an excessive supply of food, which it
cannot assimilate, is in many respects similar to that which results from
partially cutting off the supplies. And the same reasoning applies to sterility.
If by high culture, or the supply of an undue quantity of nourishment, the
constitution of the plant be impaired, or if the plant be pampered, it is no
wonderful thing that sterility should ensue. Hence, then, may it not be
asserted as a general principle that in the production of double flowers a
partial arrest of development, if not of growth, however produced, is an
essential preliminary? All the attendant phenomena, such as the obliteration
of the stamens, the augmentation in the number of floral whorls, the
occurrence of prolification, are consistent with the supposition of a primary
arrest of development, more or less complete, as the case may be: at one
time permanent, at another time relaxed and intermittent, or in a third set of
cases the vegetative activity or power of growth may be restored, and from
the centre of the flower may spring a perfect branch with perfect leaves, the
production of sheaths only being superseded by the development of leaves,
in which all the parts—sheath, stalk, and blade—are present.
When once the disposition to form double flowers is established, that
tendency becomes hereditary: there are races of single Stocks in which, out
of hundreds of plants, scarcely one double-flowered form is met with; but
when the tendency to produce double blooms is set up, single flowers
become the exception: thus, in the Balsams, before mentioned, not one in
fifty now produces single flowers, and the seeds of these double Balsams
produce double-flowered seedlings, with scarcely a "rogue" among them.
The following list of plants producing double flowers of any kind is taken
from that given in 'Seemann's Journal of Botany,' vol. ii, p. 177, and to
which some additions have been made. Miscalled double flowers, such as
those of the Compositæ, Viburnum Hydrangea, &c., are excluded.
Page 609
Ranunculaceæ.
Clematis Viticella, Linn., S. Europe.
florida, Thunb., Japan.
Fortunei, Moore, Japan.
patens, Desne, Japan.
Anemone japonica, Sieb. et Zucc., Japan.
coronaria, Linn., S. Europe, Asia Minor.
hortensis, var. Linn., S. Europe.
palmata, Linn., N. Africa, Spain, Portugal.
nemorosa, Linn., Europe, N. America, Siberia.
sylvestris, Linn., S. Europe, Siberia.
Hepatica triloba, Chaix., Europe.
Ranunculus bulbosus, Linn., Europe, N. Amer.
repens, Linn., Europe, Siberia, N. Amer.
acris, Linn., Europe, Siberia.
aconitifolius, Linn., Europe.
gramineus, Linn., Italy, France, Portugal, Switzerland.
bullatus, Linn., S. Europe.
asiaticus, Linn., The East.
Ficaria ranunculoides, Mœnch., Europe.
Thalictrum anemoides, Michæ., N. America.
Caltha palustris, Linn., Europe, Asia, N. America.
Trollius europæus, Linn., Europe.
nepalensis, Himalaya.
Nigella damascena, Linn., Mediterranean.
Aquilegia vulgaris, Linn., Europe.
canadensis, Linn., N. America.
Delphinium Ajacis, Linn., S. Europe.
grandiflorum, Linn., Siberia, N. America.
Consolida, Linn., Europe, N. America.
cheilanthum, Fisch., Siberia.
elegans, D. C., North America.
Adonis autumnalis, Linn., Europe.
vernalis, Linn., Europe, Asia.
Pæonia Moutan, Sims, China, Japan.
officinalis, Retz., Europe.
tenuifolia, Linn., Tauria.
albiflora, Pall., Siberia.
Clematis Viticella, Linn., S. Europe.
florida, Thunb., Japan.
Fortunei, Moore, Japan.
patens, Desne, Japan.
Anemone japonica, Sieb. et Zucc., Japan.
coronaria, Linn., S. Europe, Asia Minor.
hortensis, var. Linn., S. Europe.
palmata, Linn., N. Africa, Spain, Portugal.
nemorosa, Linn., Europe, N. America, Siberia.
sylvestris, Linn., S. Europe, Siberia.
Hepatica triloba, Chaix., Europe.
Ranunculus bulbosus, Linn., Europe, N. Amer.
repens, Linn., Europe, Siberia, N. Amer.
acris, Linn., Europe, Siberia.
aconitifolius, Linn., Europe.
gramineus, Linn., Italy, France, Portugal, Switzerland.
bullatus, Linn., S. Europe.
asiaticus, Linn., The East.
Ficaria ranunculoides, Mœnch., Europe.
Thalictrum anemoides, Michæ., N. America.
Caltha palustris, Linn., Europe, Asia, N. America.
Trollius europæus, Linn., Europe.
nepalensis, Himalaya.
Nigella damascena, Linn., Mediterranean.
Aquilegia vulgaris, Linn., Europe.
canadensis, Linn., N. America.
Delphinium Ajacis, Linn., S. Europe.
grandiflorum, Linn., Siberia, N. America.
Consolida, Linn., Europe, N. America.
cheilanthum, Fisch., Siberia.
elegans, D. C., North America.
Adonis autumnalis, Linn., Europe.
vernalis, Linn., Europe, Asia.
Pæonia Moutan, Sims, China, Japan.
officinalis, Retz., Europe.
tenuifolia, Linn., Tauria.
albiflora, Pall., Siberia.
Page 610
paradoxa, Andr., S. Europe.
Nymphæaceæ.
Nelumbium speciosum, Willd., Africa, Asia.
Berberidaceæ.
Berberis, sp. cult.
Papaveraceæ.
Papaver Rhœas, Linn., Europe.
bracteatum, Lindl., Russia.
somniferum, Linn., S. Europe, Asia Minor, Egypt.
Chelidonium majus, Linn., Europe, Asia.
Sanguinaria canadensis, Linn., N. America.
Podophyllum peltatum, Linn., N. America.
Cruciferæ.
Mathiola incana, R. Br., Mediterranean.
glabrata, D. C.
annua, Sweet., South Europe, Syria.
Cheiranthus Cheiri, Linn., Europe.
Iberis umbellata, Linn., Europe.
amara, Linn., Europe.
Cardamine pratensis, Linn., Europe, Asia, Africa, America.
Hesperis matronalis, Linn., Europe, Siberia.
Barbarea vulgaris, R. Br., Europe.
Sinapis arvensis, Linn., Europe.
Brassica oleracea. Linn., Europe.
Cistaceæ.
Helianthemum vulgare, Spach., Europe, N. Africa.
Violaceæ.
Viola odorata, Linn., Europe, Siberia.
Nymphæaceæ.
Nelumbium speciosum, Willd., Africa, Asia.
Berberidaceæ.
Berberis, sp. cult.
Papaveraceæ.
Papaver Rhœas, Linn., Europe.
bracteatum, Lindl., Russia.
somniferum, Linn., S. Europe, Asia Minor, Egypt.
Chelidonium majus, Linn., Europe, Asia.
Sanguinaria canadensis, Linn., N. America.
Podophyllum peltatum, Linn., N. America.
Cruciferæ.
Mathiola incana, R. Br., Mediterranean.
glabrata, D. C.
annua, Sweet., South Europe, Syria.
Cheiranthus Cheiri, Linn., Europe.
Iberis umbellata, Linn., Europe.
amara, Linn., Europe.
Cardamine pratensis, Linn., Europe, Asia, Africa, America.
Hesperis matronalis, Linn., Europe, Siberia.
Barbarea vulgaris, R. Br., Europe.
Sinapis arvensis, Linn., Europe.
Brassica oleracea. Linn., Europe.
Cistaceæ.
Helianthemum vulgare, Spach., Europe, N. Africa.
Violaceæ.
Viola odorata, Linn., Europe, Siberia.
Page 611
grandiflora, Linn., Europe,
tricolor, Linn., Europe.
Caryophylleæ.
Dianthus barbatus, Linn., France, Germany.
chinensis, D. C., China.
Poiretianus, Seringe, ?
Caryophyllus, Linn., France, Italy.
arboreus, Linn., Crete.
hybridus (gardens).
corymbosus, Sibth., Asia Minor.
plumarius, Linn., Europe, Siberia, N. America.
deltoides, Linn., Europe.
Saponaria officinalis, Linn., Europe.
Lychnis sylvestris, Schkr., Europe.
vespertina, Linn., Europe.
flos cuculi, Linn., Europe.
Viscaria, Linn., Europe.
chalcedonica, Linn., Japan, Asia Minor.
Silene inflata, Sm.; var. maritima, D. C., Europe.
Alsineæ.
Sagina procumbens, Linn., Europe.
Malvaceæ.
Hibiscus Rosa sinensis, Linn., E. Indies.
flavescens, Cav., China.
alba, Hook., China.
syriacus, Linn., Syria, Carniola.
Althæa rosea, Cav., Caucasus, &c.
Malva rotundifolia, Linn., Europe.
moschata, D. C., Europe.
Hippocastaneæ.
Æsculus Hippocastanum, Linn., Europe, N. America.
tricolor, Linn., Europe.
Caryophylleæ.
Dianthus barbatus, Linn., France, Germany.
chinensis, D. C., China.
Poiretianus, Seringe, ?
Caryophyllus, Linn., France, Italy.
arboreus, Linn., Crete.
hybridus (gardens).
corymbosus, Sibth., Asia Minor.
plumarius, Linn., Europe, Siberia, N. America.
deltoides, Linn., Europe.
Saponaria officinalis, Linn., Europe.
Lychnis sylvestris, Schkr., Europe.
vespertina, Linn., Europe.
flos cuculi, Linn., Europe.
Viscaria, Linn., Europe.
chalcedonica, Linn., Japan, Asia Minor.
Silene inflata, Sm.; var. maritima, D. C., Europe.
Alsineæ.
Sagina procumbens, Linn., Europe.
Malvaceæ.
Hibiscus Rosa sinensis, Linn., E. Indies.
flavescens, Cav., China.
alba, Hook., China.
syriacus, Linn., Syria, Carniola.
Althæa rosea, Cav., Caucasus, &c.
Malva rotundifolia, Linn., Europe.
moschata, D. C., Europe.
Hippocastaneæ.
Æsculus Hippocastanum, Linn., Europe, N. America.
Page 612
Geraniaceæ.
Geranium pratense, Linn., Europe, Siberia.
sylvaticum. Linn., Europe.
Pelargonium zonale, Willd., S. Africa.
Tropæolum majus, Linn., Peru.
minus, Linn., Peru.
Oxalis cernua, Thunb., S. Africa.
Impatiens Balsamina, Linn., E. Ind.
Ternströmiaceæ.
Camellia reticulata, Lindl., China.
Sasanqua, Thunb., China.
japonica, Linn., Japan.
Thea maliflora, Seem., Japan.
Aurantiaceæ.
Citrus Aurantium, Linn., Asia, South Europe.
Papilionaceæ.
Trifolium repens, Linn., Europe, S. America.
Medicago sp., ?., Europe.
Ulex europæus, Link., Europe.
Spartianthus junceus, Linn., S. Europe.
Clitoria Ternatea, Linn., E. India.
Orobus viscoides, D. C., Croatia, &c.
vernus, Linn., Europe.
Genista tinctoria, Linn., Europe.
sibirica, Linn., Siberia.
scoparia, Lam., Europe.
Cytisus albus, Link., Portugal.
Anthyllis Vulneraria, Linn., Europe.
Coronilla Emerus, D. C., Europe.
Lotus corniculatus, Linn., Europe.
Rosaceæ.
Geranium pratense, Linn., Europe, Siberia.
sylvaticum. Linn., Europe.
Pelargonium zonale, Willd., S. Africa.
Tropæolum majus, Linn., Peru.
minus, Linn., Peru.
Oxalis cernua, Thunb., S. Africa.
Impatiens Balsamina, Linn., E. Ind.
Ternströmiaceæ.
Camellia reticulata, Lindl., China.
Sasanqua, Thunb., China.
japonica, Linn., Japan.
Thea maliflora, Seem., Japan.
Aurantiaceæ.
Citrus Aurantium, Linn., Asia, South Europe.
Papilionaceæ.
Trifolium repens, Linn., Europe, S. America.
Medicago sp., ?., Europe.
Ulex europæus, Link., Europe.
Spartianthus junceus, Linn., S. Europe.
Clitoria Ternatea, Linn., E. India.
Orobus viscoides, D. C., Croatia, &c.
vernus, Linn., Europe.
Genista tinctoria, Linn., Europe.
sibirica, Linn., Siberia.
scoparia, Lam., Europe.
Cytisus albus, Link., Portugal.
Anthyllis Vulneraria, Linn., Europe.
Coronilla Emerus, D. C., Europe.
Lotus corniculatus, Linn., Europe.
Rosaceæ.
Page 613
Rosa lutea, Mill., Europe.
cinnamomea, Linn., Europe, N. America.
spinosissima, Linn., Central Asia.
Carolina, Linn., N. America.
villosa, Linn., Europe, Central Asia.
centifolia, Linn.
damascena, Linn., Syria.
rubiginosa, Linn., Europe, Asia, N. America.
moschata, Ait., Madeira, N. Africa.
canina, Linn., Europe.
alba, Linn., Europe, Caucasus.
indica, Linn., China.
nivea, D. C., China.
Eglanteria, Linn., Europe.
gallica, Linn., Europe, Caucasus.
pimpinellifolia, Linn., Europe, Central Asia.
Banksiæ, R. Br., China.
sulphurea, Ait., East.
Rubus fruticosus, Linn., Europe.
rosifolius, Linn., Mauritius, E. India.
corylifolius, Smith, Europe.
cæsius, Linn., Europe.
Kerria japonica, D. C., Japan.
Spiræa Filipendula, Linn., Europe.
Ulmaria, Linn., Europe.
prunifolia, Sieb. et Zucc., Japan.
Reevesii, Lindl., China.
strobilacea, Sieb. et Zucc., Japan.
Fragaria vesca, Linn., Europe, N. America.
Potentilla alpestris, Hall. f., Europe.
reptans, Linn., Europe, Asia.
Tormentilla, Schrank, Europe, Asia.
anserina, Linn., Europe.
Geum rivale, Linn., Europe.
Pomaceæ.
Cratægus Oxyacantha, Linn., Europe.
Crus galli, Linn., N. America.
cinnamomea, Linn., Europe, N. America.
spinosissima, Linn., Central Asia.
Carolina, Linn., N. America.
villosa, Linn., Europe, Central Asia.
centifolia, Linn.
damascena, Linn., Syria.
rubiginosa, Linn., Europe, Asia, N. America.
moschata, Ait., Madeira, N. Africa.
canina, Linn., Europe.
alba, Linn., Europe, Caucasus.
indica, Linn., China.
nivea, D. C., China.
Eglanteria, Linn., Europe.
gallica, Linn., Europe, Caucasus.
pimpinellifolia, Linn., Europe, Central Asia.
Banksiæ, R. Br., China.
sulphurea, Ait., East.
Rubus fruticosus, Linn., Europe.
rosifolius, Linn., Mauritius, E. India.
corylifolius, Smith, Europe.
cæsius, Linn., Europe.
Kerria japonica, D. C., Japan.
Spiræa Filipendula, Linn., Europe.
Ulmaria, Linn., Europe.
prunifolia, Sieb. et Zucc., Japan.
Reevesii, Lindl., China.
strobilacea, Sieb. et Zucc., Japan.
Fragaria vesca, Linn., Europe, N. America.
Potentilla alpestris, Hall. f., Europe.
reptans, Linn., Europe, Asia.
Tormentilla, Schrank, Europe, Asia.
anserina, Linn., Europe.
Geum rivale, Linn., Europe.
Pomaceæ.
Cratægus Oxyacantha, Linn., Europe.
Crus galli, Linn., N. America.
Page 614
Cydonia japonica, Pers., Japan.
Pyrus communis, Linn., Europe.
Malus, Linn., Europe.
Eriobotrya japonica, Lindl., Japan.
Amygdaleæ.
Amygdalus Persica, Linn., Persia.
communis, Linn., Mauritania.
Prunus domestica, Linn., Europe.
spinosa, Linn., Europe, N. America.
avium, Linn., Europe.
Cerasus, Linn., Europe.
Kerii, Steud., Japan.
japonica, Thunb., China, Japan.
insititia, Linn., Europe.
triloba, Lindl., China.
Myrtaceæ.
Myrtus communis, Linn., S. Europe.
Punica Granatum, Linn., S. Europe, Marocco.
Philadelphaceæ.
Philadelphus Coronarius, linn., S. Europe.
Deutzia Crenata, sieb. Et Zucc., Japan.
Onagraceæ.
Fuchsia globosa, Lindl. (and var. hort. pl.), Mexico.
Epilobium tetragonum, D.C., Europe.
Clarkia pulchella, Pursh., California.
elegans, Douglas, N. America.
Portulacaceæ.
Portulaca grandiflora, Hook, Chili.
Pyrus communis, Linn., Europe.
Malus, Linn., Europe.
Eriobotrya japonica, Lindl., Japan.
Amygdaleæ.
Amygdalus Persica, Linn., Persia.
communis, Linn., Mauritania.
Prunus domestica, Linn., Europe.
spinosa, Linn., Europe, N. America.
avium, Linn., Europe.
Cerasus, Linn., Europe.
Kerii, Steud., Japan.
japonica, Thunb., China, Japan.
insititia, Linn., Europe.
triloba, Lindl., China.
Myrtaceæ.
Myrtus communis, Linn., S. Europe.
Punica Granatum, Linn., S. Europe, Marocco.
Philadelphaceæ.
Philadelphus Coronarius, linn., S. Europe.
Deutzia Crenata, sieb. Et Zucc., Japan.
Onagraceæ.
Fuchsia globosa, Lindl. (and var. hort. pl.), Mexico.
Epilobium tetragonum, D.C., Europe.
Clarkia pulchella, Pursh., California.
elegans, Douglas, N. America.
Portulacaceæ.
Portulaca grandiflora, Hook, Chili.
Page 615
Grossulariaceæ.
Ribes sanguineum, Pursh., N. America.
Saxifragaceæ.
Saxifraga granulata, Linn., Europe.
Umbelliferæ.
Daucus Carota, Linn., Europe.
Rubiaceæ.
Ixora grandiflora, De Cand., E. India.
Serissa fœtida, Comm., China, Japan.
Gardenia Fortuniana, Hook., China.
florida, Linn., China, E. India.
radicans, Thunb., Japan.
Caprifoliaceæ.
Lonicera Periclymenum, Linn., Europe.
Sambucus nigra, Linn., Europe.
Campanulaceæ.
Campanula latifolia, Linn., Europe, Asia.
Tenorei, Morett, Naples.
Trachelium, Linn., Europe.
Vidallii, H. C. Wats., Europe.
pyramidalis, Linn., S. Europe.
rotundifolia, Linn., Europe, N. America.
persicifolia, Linn., Europe.
glomerata, Linn., Europe, Asia.
Medium, Linn., Europe.
rhomboidea, Linn., Europe.
Platycodon grandiflorum, D. C., Siberia.
Ribes sanguineum, Pursh., N. America.
Saxifragaceæ.
Saxifraga granulata, Linn., Europe.
Umbelliferæ.
Daucus Carota, Linn., Europe.
Rubiaceæ.
Ixora grandiflora, De Cand., E. India.
Serissa fœtida, Comm., China, Japan.
Gardenia Fortuniana, Hook., China.
florida, Linn., China, E. India.
radicans, Thunb., Japan.
Caprifoliaceæ.
Lonicera Periclymenum, Linn., Europe.
Sambucus nigra, Linn., Europe.
Campanulaceæ.
Campanula latifolia, Linn., Europe, Asia.
Tenorei, Morett, Naples.
Trachelium, Linn., Europe.
Vidallii, H. C. Wats., Europe.
pyramidalis, Linn., S. Europe.
rotundifolia, Linn., Europe, N. America.
persicifolia, Linn., Europe.
glomerata, Linn., Europe, Asia.
Medium, Linn., Europe.
rhomboidea, Linn., Europe.
Platycodon grandiflorum, D. C., Siberia.
Page 616
Ericaceæ.
Calluna vulgaris, Linn., Europe, N. America.
Rhododendron indicum, Sweet., E. India.
ponticum, Linn., Asia Minor.
Azalea nudiflora, Linn., N. America.
glauca, Lam., N. America.
Arbutus Unedo, Linn., S. Europe.
Erica Tetralix, Linn., Europe.
cinerea, Linn., Europe.
hyemalis, gardens.
Epacridaceæ.
Epacris impressa, R. Br., Australia.
Primulaceæ.
Primula villosa, Jacq., Europe.
Auricula, Linn., Europe.
denticulata, Smith, E. India.
acaulis, Jacq., Europe.
clatior, Jacq., Europe.
prænitens, Ker. = sinensis, Lindl., China.
Lysimachia Nummularia, Roem et Schult., Europe.
Anagallis tenella, Linn., Europe.
Jasminaceæ.
Jasminum officinale, Linn., S. Europe.
Sambac., Ait., E. India.
hirsutum, Hook., China.
grandiflorum, Lindl., S. Europe.
Oleaceæ.
Syringa persica, Linn., Persia.
vulgaris, Linn., Europe, Persia.
Calluna vulgaris, Linn., Europe, N. America.
Rhododendron indicum, Sweet., E. India.
ponticum, Linn., Asia Minor.
Azalea nudiflora, Linn., N. America.
glauca, Lam., N. America.
Arbutus Unedo, Linn., S. Europe.
Erica Tetralix, Linn., Europe.
cinerea, Linn., Europe.
hyemalis, gardens.
Epacridaceæ.
Epacris impressa, R. Br., Australia.
Primulaceæ.
Primula villosa, Jacq., Europe.
Auricula, Linn., Europe.
denticulata, Smith, E. India.
acaulis, Jacq., Europe.
clatior, Jacq., Europe.
prænitens, Ker. = sinensis, Lindl., China.
Lysimachia Nummularia, Roem et Schult., Europe.
Anagallis tenella, Linn., Europe.
Jasminaceæ.
Jasminum officinale, Linn., S. Europe.
Sambac., Ait., E. India.
hirsutum, Hook., China.
grandiflorum, Lindl., S. Europe.
Oleaceæ.
Syringa persica, Linn., Persia.
vulgaris, Linn., Europe, Persia.
Page 617
Apocyneæ.
Vinca minor, Linn., Europe.
major, Linn., Europe.
Nerium odorum, Ait., E. India.
Oleander, Linn., S. Europe.
Tabernæmontana coronaria, Willd., E. India.
Allamanda cathartica, Aubl., S. America.
Convolvulaceæ.
Calystegia sepium, R. Br., Europe, America, Asia.
pubescens, Lindl., China.
Convolvulus tricolor, Linn., S. Europe.
Ipomœa pandurata, Meyer, S. America.
Solanaceæ.
Datura cornigera, Hook., Peru.
fastuosa, Linn., S. America, Egypt.
arborea, Linn., S. America.
chlorantha, Hook.
humilis, Desf.
Petunia nyctaginiflora, Juss., S. America.
violacea, Hook, S. America.
Solanum Dulcamara, Linn., Europe.
Gentianaceæ.
Gentiana Amarella, Linn., Europe.
Orobanchaceæ.
Orobanche sp.
Scrophulariaceæ.
Mimulus luteus, Linn., Chili.
Antirrhinum majus, Linn., S. Europe.
Vinca minor, Linn., Europe.
major, Linn., Europe.
Nerium odorum, Ait., E. India.
Oleander, Linn., S. Europe.
Tabernæmontana coronaria, Willd., E. India.
Allamanda cathartica, Aubl., S. America.
Convolvulaceæ.
Calystegia sepium, R. Br., Europe, America, Asia.
pubescens, Lindl., China.
Convolvulus tricolor, Linn., S. Europe.
Ipomœa pandurata, Meyer, S. America.
Solanaceæ.
Datura cornigera, Hook., Peru.
fastuosa, Linn., S. America, Egypt.
arborea, Linn., S. America.
chlorantha, Hook.
humilis, Desf.
Petunia nyctaginiflora, Juss., S. America.
violacea, Hook, S. America.
Solanum Dulcamara, Linn., Europe.
Gentianaceæ.
Gentiana Amarella, Linn., Europe.
Orobanchaceæ.
Orobanche sp.
Scrophulariaceæ.
Mimulus luteus, Linn., Chili.
Antirrhinum majus, Linn., S. Europe.
Page 618
Digitalis purpurea, Linn., Europe.
Linaria vulgaris, Mill., Europe, N. America.
Veronica, sp.
Calceolaria, var. cult.
Gesneraceæ.
Achimenes longiflora, D. C., Mexico.
Gloxinia var. hort.
Verbenaceæ.
Clerodendron fragrans, Willd., Japan.
Verbena var. hort.
Nyctagineæ.
Mirabilis Jalapa, Linn., Trop. America.
Laurineæ.
Laurus nobilis, Linn., S. Europe.
Sassafras, Linn., N. America.
Iridaceæ.
Gladiolus tristis, Linn., Cape of Good Hope.
Crocus aureus, Sibth, Europe, Asia Minor.
Susianus, Curt., Asia Minor.
pusillus, Tenore, Italy.
vernus, Smith, S. Europe.
Iris sibirica, Linn., Europe.
Iris Kæmpferi, Siebold, Japan.
Amaryllidaceæ.
Galanthus nivalis, Linn., Europe.
Leucoium vernum, Linn., Europe.
Sternbergia lutea, Gawl., Europe, Asia Minor.
Hippeastrum equestre, Herb., S. America.
Linaria vulgaris, Mill., Europe, N. America.
Veronica, sp.
Calceolaria, var. cult.
Gesneraceæ.
Achimenes longiflora, D. C., Mexico.
Gloxinia var. hort.
Verbenaceæ.
Clerodendron fragrans, Willd., Japan.
Verbena var. hort.
Nyctagineæ.
Mirabilis Jalapa, Linn., Trop. America.
Laurineæ.
Laurus nobilis, Linn., S. Europe.
Sassafras, Linn., N. America.
Iridaceæ.
Gladiolus tristis, Linn., Cape of Good Hope.
Crocus aureus, Sibth, Europe, Asia Minor.
Susianus, Curt., Asia Minor.
pusillus, Tenore, Italy.
vernus, Smith, S. Europe.
Iris sibirica, Linn., Europe.
Iris Kæmpferi, Siebold, Japan.
Amaryllidaceæ.
Galanthus nivalis, Linn., Europe.
Leucoium vernum, Linn., Europe.
Sternbergia lutea, Gawl., Europe, Asia Minor.
Hippeastrum equestre, Herb., S. America.
Page 619
Narcissus cernuus, Salisb., S. Europe.
Telamonius, Schult., Europe.
lobularis, Schult.
concolor, Schult., Portugal.
biflorus, Curt., Europe.
italicus, Ker., Italy.
incomparabilis, Curt., Italy.
Cypri, Haw., Cyprus.
Pseudo-Narcissus, Linn., Europe.
poeticus, Linn., Europe.
Jonquilla, Linn., S. Europe, East.
Tazetta, Linn., S. Europe.
poculiformis, Salisb., S. Europe.
Orchidaceæ.
Orchis Morio, Linn., Europe.
mascula, Linn., Europe.
pyramidalis, Linn., Europe.
Ophrys fucifera, Linn., Europe.
See also pp. 380, 509.
Hydrocharidaceæ.
Hydrocharis Morsus ranæ, Linn., Europe.
Asphodeleæ.
Asphodelus luteus, Linn., S. Europe.
Liliaceæ.
Tulipa Gesneriana, Linn., Asia Minor.
sylvestris, Linn., S. Europe.
Scilla autumnalis, Linn., Europe.
nutans, Smith, S. Europe.
Convallaria majalis, Linn., Europe, America.
Polygonatum, Linn., Europe.
Trillium grandiflorum, Spreng., America.
Fritillaria Meleagris, Linn., Europe.
Telamonius, Schult., Europe.
lobularis, Schult.
concolor, Schult., Portugal.
biflorus, Curt., Europe.
italicus, Ker., Italy.
incomparabilis, Curt., Italy.
Cypri, Haw., Cyprus.
Pseudo-Narcissus, Linn., Europe.
poeticus, Linn., Europe.
Jonquilla, Linn., S. Europe, East.
Tazetta, Linn., S. Europe.
poculiformis, Salisb., S. Europe.
Orchidaceæ.
Orchis Morio, Linn., Europe.
mascula, Linn., Europe.
pyramidalis, Linn., Europe.
Ophrys fucifera, Linn., Europe.
See also pp. 380, 509.
Hydrocharidaceæ.
Hydrocharis Morsus ranæ, Linn., Europe.
Asphodeleæ.
Asphodelus luteus, Linn., S. Europe.
Liliaceæ.
Tulipa Gesneriana, Linn., Asia Minor.
sylvestris, Linn., S. Europe.
Scilla autumnalis, Linn., Europe.
nutans, Smith, S. Europe.
Convallaria majalis, Linn., Europe, America.
Polygonatum, Linn., Europe.
Trillium grandiflorum, Spreng., America.
Fritillaria Meleagris, Linn., Europe.
Page 620
imperialis, Linn., Persia.
Lilium Martagon, Linn., Europe.
candidum, Linn., Syria, Persia.
Hyacinthus orientalis, Linn., East.
Polianthes tuberosa, Linn., E. India.
Hemerocallis disticha, Don., Nepal.
Kwanso, gardens.
fulva, Linn., S. Europe.
Colchicaceæ.
Colchicum autumnale, Linn., Europe.
Tofieldia calyculata, Wahl., Europe.
Butomaceæ.
Sagittaria latifolia, Willd., N. America.
sagittifolia, Linn., Europe, Asia, America.
Commelynaceæ.
Tradescantia virginica, Linn., N. America.
alba, gardens.
FOOTNOTES:
[564] This appendix forms a portion of a paper published in the 'Proceedings of the
International Botanical Congress,' London, 1886, p. 127, and which it has been deemed
advisable to reproduce with sundry additions and modifications.
[565] 'Traité des Giroflées,' per E. Chaté.
[566] Leading Article in the 'Gardeners' Chronicle,' p. 74, 1866.
[567] Otto's 'Gartenzeitung,' 1866.
[568] 'Gard. Chron.,' 1843, p. 628.
[569] 'Gard. Chron.,' 1867, p. 381.—Art. "Chinese primroses."
[570] See also p. 79, fig. 36. A similar flower is figured in 'Hort. Eystett. Ic. Arb. Vern.,' fol.
5. "Fructus nondum observatus est fortassis alimento uberius in flores refuso, nullus sperari
Lilium Martagon, Linn., Europe.
candidum, Linn., Syria, Persia.
Hyacinthus orientalis, Linn., East.
Polianthes tuberosa, Linn., E. India.
Hemerocallis disticha, Don., Nepal.
Kwanso, gardens.
fulva, Linn., S. Europe.
Colchicaceæ.
Colchicum autumnale, Linn., Europe.
Tofieldia calyculata, Wahl., Europe.
Butomaceæ.
Sagittaria latifolia, Willd., N. America.
sagittifolia, Linn., Europe, Asia, America.
Commelynaceæ.
Tradescantia virginica, Linn., N. America.
alba, gardens.
FOOTNOTES:
[564] This appendix forms a portion of a paper published in the 'Proceedings of the
International Botanical Congress,' London, 1886, p. 127, and which it has been deemed
advisable to reproduce with sundry additions and modifications.
[565] 'Traité des Giroflées,' per E. Chaté.
[566] Leading Article in the 'Gardeners' Chronicle,' p. 74, 1866.
[567] Otto's 'Gartenzeitung,' 1866.
[568] 'Gard. Chron.,' 1843, p. 628.
[569] 'Gard. Chron.,' 1867, p. 381.—Art. "Chinese primroses."
[570] See also p. 79, fig. 36. A similar flower is figured in 'Hort. Eystett. Ic. Arb. Vern.,' fol.
5. "Fructus nondum observatus est fortassis alimento uberius in flores refuso, nullus sperari
Page 621
possit."
[571] See De Candolle, 'Plant. Rar. Genev.,' 1829, p. 91; and Alph. de Candolle.' Géog. Bot.,'
p. 1080.
[572] See 'Gardeners' Chronicle,' 1868, p. 1113.
[573] Ibid., 1843, p. 628.
[571] See De Candolle, 'Plant. Rar. Genev.,' 1829, p. 91; and Alph. de Candolle.' Géog. Bot.,'
p. 1080.
[572] See 'Gardeners' Chronicle,' 1868, p. 1113.
[573] Ibid., 1843, p. 628.
Page 622
NOTE.
During the progress of the foregoing pages through the press, several
additional illustrations of particular malformations have come under notice.
Some of the more important of these may here be recorded.
Fasciation (see p. 11).—The following plants may be added to the list:—
Acer eriocarpum, Arabis albida, Brassica oleracea, var., Guarea, sp.,
Artabotrys sp. In all, with the exception of the first-named, the fasciation
occurred in the inflorescence. In some species of Artabotrys, indeed,
fasciation and curvation of the inflorescence are common.
Synanthy (p. 39).—Several additional instances of adhesion of two or more
flowers in Calanthe vestita, C. Veitchii, and other forms of this genus may
be cited. These furnish further illustrations of the much greater liability of
some plants to particular changes as compared with others. Scilla bifolia,
Gagea arvensis, and Viola odorata may be added to the list of synanthic
plants.
Alterations of placentation, &c. (see pp. 98, 483).—M. Casimir De
Candolle, in a letter to the author, dated March 8th, 1869, thus writes of the
existence of a double row of carpels in Pyrus spectabilis and Cratægus
Oxyacantha, "a longitudinal section of a double flower of Pyrus spectabilis
shows two rows of carpels, placed one above another. The arrangement of
During the progress of the foregoing pages through the press, several
additional illustrations of particular malformations have come under notice.
Some of the more important of these may here be recorded.
Fasciation (see p. 11).—The following plants may be added to the list:—
Acer eriocarpum, Arabis albida, Brassica oleracea, var., Guarea, sp.,
Artabotrys sp. In all, with the exception of the first-named, the fasciation
occurred in the inflorescence. In some species of Artabotrys, indeed,
fasciation and curvation of the inflorescence are common.
Synanthy (p. 39).—Several additional instances of adhesion of two or more
flowers in Calanthe vestita, C. Veitchii, and other forms of this genus may
be cited. These furnish further illustrations of the much greater liability of
some plants to particular changes as compared with others. Scilla bifolia,
Gagea arvensis, and Viola odorata may be added to the list of synanthic
plants.
Alterations of placentation, &c. (see pp. 98, 483).—M. Casimir De
Candolle, in a letter to the author, dated March 8th, 1869, thus writes of the
existence of a double row of carpels in Pyrus spectabilis and Cratægus
Oxyacantha, "a longitudinal section of a double flower of Pyrus spectabilis
shows two rows of carpels, placed one above another. The arrangement of
Page 623
the vascular bundles shows that the upper row is external in relation to the
lower series. The carpels of the latter are wholly coalescent as in a pear,
while those of the upper verticil are only partially coherent or sometimes
quite distinct. The placentation is constantly axile in the inferior row and
parietal in the upper one. The number of ovules in each carpel of the
superior row varies greatly, and they are often, but not always, inserted in
two longitudinal ranks, as is constantly the case in the lower carpels.
Double flowers of Cratægus Oxyacantha present the same anomalies." For
analogous instances in Digitalis, see p. 98. See also p. 380, Saxifraga.
Prolification, p. 120.—A. P. De Candolle, "Organographie Végétale," tab.
40, figures an instance of suppression of one lobe of the ovary in Iris
chinensis, and of the presence at the base of the flower of an adventitious
and imperfect flower-bud, as in the Phlomis, mentioned at p. 119.
Monœcious Misleto, p. 193.—In this specimen, exhibited at one of the
meetings of the Scientific Committee of the Royal Horticultural Society in
1869, there were both male and female flowers on the same bush. The plant
was of the male sex, with numerous long slender whip-like, somewhat
pendulous, branches bearing comparatively large broad yellowish leaves,
and fully developed male flowers at the end. From the side of one of these
male branches, near the base, protruded a tuft of short, stiff branches,
bearing small, narrow, dark green leaves, ripe berries and immature female
flowers. There was no evidence of grafting or parasitism, of the female
branch on the male, the bark and the wood being perfectly continuous so
that the only tenable supposition is that this was a case of dimorphism.
Adventitious leaflet and pitcher, see pp. 30 and 355. In a species of
Picrasma, in which the leaves are impari-pinnate and spread horizontally,
an adventitious leaflet was observed to project at right angles to the plane of
the primary leaf. It emerged at a point nearly corresponding to that at which
lower series. The carpels of the latter are wholly coalescent as in a pear,
while those of the upper verticil are only partially coherent or sometimes
quite distinct. The placentation is constantly axile in the inferior row and
parietal in the upper one. The number of ovules in each carpel of the
superior row varies greatly, and they are often, but not always, inserted in
two longitudinal ranks, as is constantly the case in the lower carpels.
Double flowers of Cratægus Oxyacantha present the same anomalies." For
analogous instances in Digitalis, see p. 98. See also p. 380, Saxifraga.
Prolification, p. 120.—A. P. De Candolle, "Organographie Végétale," tab.
40, figures an instance of suppression of one lobe of the ovary in Iris
chinensis, and of the presence at the base of the flower of an adventitious
and imperfect flower-bud, as in the Phlomis, mentioned at p. 119.
Monœcious Misleto, p. 193.—In this specimen, exhibited at one of the
meetings of the Scientific Committee of the Royal Horticultural Society in
1869, there were both male and female flowers on the same bush. The plant
was of the male sex, with numerous long slender whip-like, somewhat
pendulous, branches bearing comparatively large broad yellowish leaves,
and fully developed male flowers at the end. From the side of one of these
male branches, near the base, protruded a tuft of short, stiff branches,
bearing small, narrow, dark green leaves, ripe berries and immature female
flowers. There was no evidence of grafting or parasitism, of the female
branch on the male, the bark and the wood being perfectly continuous so
that the only tenable supposition is that this was a case of dimorphism.
Adventitious leaflet and pitcher, see pp. 30 and 355. In a species of
Picrasma, in which the leaves are impari-pinnate and spread horizontally,
an adventitious leaflet was observed to project at right angles to the plane of
the primary leaf. It emerged at a point nearly corresponding to that at which
Page 624
the normal pinnæ were given off. The appearance presented was thus like
that of a whorl of three leaves, except that the shining surface of the
adventitious leaflet, corresponding to the upper face of the normal leaflets,
was directed towards the axis, i.e., away from the corresponding portion of
the neighbouring pinnæ, while the dull surface, corresponding to the lower
part of an ordinary leaflet, looked towards the apex of the main leaf, or
away from the axis. In one instance, a stalked pitcher was given off from
the same point as that from which the supernumerary leaflet emerged, the
pitcher being apparently formed from the cohesion (congenital) of the
margins of a leaflet.
In the normal leaf of this plant there is between the bases of the pinnæ, a
small reddish gland or stipel? attached to, or projecting from, the upper
surface of the rachis. It appeared from some transitional forms that the
adventitious leaflet, just mentioned, was due to the exaggerated
development of this gland, but no clue was afforded as to the origin of the
ascidium. It was not practicable to examine the arrangement of the vascular
bundles in the rachis.
Additional labella in Phaius.—A flower of Phaius grandiflorus was found
in the same condition as the Catasetum, mentioned at pp. 291 and 382.
Tubular stem.—A species of Sempervivum, exhibited by Mr. Salter, of
Hammersmith, at one of the summer exhibitions of flowers at the Royal
Horticultural Society in 1868, under the name of S. Bollei, deserves notice
from its bearing on the question of such structures as the calyx-tubes, the
hip of the rose and such like, see pp. 394, 482. In this plant the leaves
appeared to be arranged some on the outside, others on the inside, of an
erect hollow cylinder, some six inches in height. The oldest leaves were
outside, the youngest within, so that the appearance presented was as if the
that of a whorl of three leaves, except that the shining surface of the
adventitious leaflet, corresponding to the upper face of the normal leaflets,
was directed towards the axis, i.e., away from the corresponding portion of
the neighbouring pinnæ, while the dull surface, corresponding to the lower
part of an ordinary leaflet, looked towards the apex of the main leaf, or
away from the axis. In one instance, a stalked pitcher was given off from
the same point as that from which the supernumerary leaflet emerged, the
pitcher being apparently formed from the cohesion (congenital) of the
margins of a leaflet.
In the normal leaf of this plant there is between the bases of the pinnæ, a
small reddish gland or stipel? attached to, or projecting from, the upper
surface of the rachis. It appeared from some transitional forms that the
adventitious leaflet, just mentioned, was due to the exaggerated
development of this gland, but no clue was afforded as to the origin of the
ascidium. It was not practicable to examine the arrangement of the vascular
bundles in the rachis.
Additional labella in Phaius.—A flower of Phaius grandiflorus was found
in the same condition as the Catasetum, mentioned at pp. 291 and 382.
Tubular stem.—A species of Sempervivum, exhibited by Mr. Salter, of
Hammersmith, at one of the summer exhibitions of flowers at the Royal
Horticultural Society in 1868, under the name of S. Bollei, deserves notice
from its bearing on the question of such structures as the calyx-tubes, the
hip of the rose and such like, see pp. 394, 482. In this plant the leaves
appeared to be arranged some on the outside, others on the inside, of an
erect hollow cylinder, some six inches in height. The oldest leaves were
outside, the youngest within, so that the appearance presented was as if the
Page 625
summit of the axis had been pushed or drawn in, much as the finger of a
tight glove might be invaginated in withdrawing it from the hand.
The plant in question thus furnishes an actual illustration of the
supposititious case mentioned at p. 482.
Double flowers, see pp. 499, et seq.—The following species may be added
to those already recorded: Lychnis coronaria, Hibiscus mutabilis, Lotus
major, Pisum sativum, Godetia sp., Ipomœa purpurea, Convolvulus minor,
Heliotropium peruvianum, Trillium grandiflorum, and Phaius grandiflorus.
tight glove might be invaginated in withdrawing it from the hand.
The plant in question thus furnishes an actual illustration of the
supposititious case mentioned at p. 482.
Double flowers, see pp. 499, et seq.—The following species may be added
to those already recorded: Lychnis coronaria, Hibiscus mutabilis, Lotus
major, Pisum sativum, Godetia sp., Ipomœa purpurea, Convolvulus minor,
Heliotropium peruvianum, Trillium grandiflorum, and Phaius grandiflorus.
Page 626
INDEX TO SUBJECTS.
A.
Abortion, 455, 467
of axile organs, 455
calyx, 460
corolla, 460
indusium, 467
leaves, 458
ovules, 466
perianth, 460
pistil, 464
receptacle, 457
stamens, 463
Acaulescence, 393
Acaulosia, 393, 456
Acheilary, 398
Adesiny, 58, 76
Adhesion, 32
of axes, 50, 55
embryos, 56
leaves, 33
parts of flower, 34
roots, 53
Adventitious buds, 156, 176
flowers, 174, 176, 180
gemmæ, 173
leaves, 162
roots, 156
shoots, 161
Albinism, 337
Alternation, 3, 485
Andrœcium, enlargement of, 430
A.
Abortion, 455, 467
of axile organs, 455
calyx, 460
corolla, 460
indusium, 467
leaves, 458
ovules, 466
perianth, 460
pistil, 464
receptacle, 457
stamens, 463
Acaulescence, 393
Acaulosia, 393, 456
Acheilary, 398
Adesiny, 58, 76
Adhesion, 32
of axes, 50, 55
embryos, 56
leaves, 33
parts of flower, 34
roots, 53
Adventitious buds, 156, 176
flowers, 174, 176, 180
gemmæ, 173
leaves, 162
roots, 156
shoots, 161
Albinism, 337
Alternation, 3, 485
Andrœcium, enlargement of, 430
Page 627
meiophylly of, 398
meiotaxy of, 405
of orchids, 380
pleiotaxy of, 379
polyphylly of, 361
Androgynism, 193
Anther (see connective), contabescence of, 463
morphology of, 291
ovuliferous, 200
petalody of, 291
sutures of, 291
Apilary, 397
Apostasis, 440
Aphylly, 395
Arrangement, 1
Ascidia, 30, 313
bibliography, 30
plants with, 30
Atrophy, 454 (see abortion)
Antophyllogeny, 355
Avalidouires (vines), 211
Axes, abortion of, 455
adhesion of, 50, 55
cohesion of, 9
enation from, 444
enlargement of, 418
fission of, 60
suppression of, 398
Axile organs, see Axes
B.
Barley, Nepaul, 174
Bigarades cornues, 303
Bladder-plums, 465
Bracts, multiplication of, 358
staminody of, 298
phyllody of, 242
Buds, adventitious on fruits, 178
on leaves, 170, 174
in ovary, 180
on petals, 177
in pith, 171
on roots, 160 (see flower-buds, prolification)
variations of, 336
Bulbs, displacement of, 84
multiplication of, 172, 350
meiotaxy of, 405
of orchids, 380
pleiotaxy of, 379
polyphylly of, 361
Androgynism, 193
Anther (see connective), contabescence of, 463
morphology of, 291
ovuliferous, 200
petalody of, 291
sutures of, 291
Apilary, 397
Apostasis, 440
Aphylly, 395
Arrangement, 1
Ascidia, 30, 313
bibliography, 30
plants with, 30
Atrophy, 454 (see abortion)
Antophyllogeny, 355
Avalidouires (vines), 211
Axes, abortion of, 455
adhesion of, 50, 55
cohesion of, 9
enation from, 444
enlargement of, 418
fission of, 60
suppression of, 398
Axile organs, see Axes
B.
Barley, Nepaul, 174
Bigarades cornues, 303
Bladder-plums, 465
Bracts, multiplication of, 358
staminody of, 298
phyllody of, 242
Buds, adventitious on fruits, 178
on leaves, 170, 174
in ovary, 180
on petals, 177
in pith, 171
on roots, 160 (see flower-buds, prolification)
variations of, 336
Bulbs, displacement of, 84
multiplication of, 172, 350
Page 628
Burrs, 347, 420
C.
Catacorolla, 450
Calycanthemy, 283
Calyphyomy, 34
Calyx, abortion of, 461
dialysis of, 70
meiophylly of, 396
meiotaxy of, 403
obsolete, 460
petalody of, 283
pleiotaxy of, 374
polyphylly of, 359
solution of from ovary, 77
Calyx-tube, 394, 480, 509
Carnation wheat-ear, 371
Carpels (see Pistil, Fruit, Ovary), adventitious, 182
dialysis of, 73
enation from, 453
fission of, 68
in ovary, 182
Cauliflower, 421
Cenanthy, 408
Chloranthy, 273, 279
bibliography, 280
remarks on, 279
plants subject to, 280
Chorisis, 59, 343
parallel, 344
collateral, 344
Chromatism, 339
Cladodes, 328
Classification, teratology in relation to, 488
Cohesion of leaves, 21, 25
of petals, 28
pistils, 29
sepals, 27
stamens, 29
stems, 9
Colour, alterations of, 337
Columella, 395
Coniferæ, leaves of, 217, 352, 484
inflorescence of, 245
scales of, 192, 245, 484
Compensation, 488
C.
Catacorolla, 450
Calycanthemy, 283
Calyphyomy, 34
Calyx, abortion of, 461
dialysis of, 70
meiophylly of, 396
meiotaxy of, 403
obsolete, 460
petalody of, 283
pleiotaxy of, 374
polyphylly of, 359
solution of from ovary, 77
Calyx-tube, 394, 480, 509
Carnation wheat-ear, 371
Carpels (see Pistil, Fruit, Ovary), adventitious, 182
dialysis of, 73
enation from, 453
fission of, 68
in ovary, 182
Cauliflower, 421
Cenanthy, 408
Chloranthy, 273, 279
bibliography, 280
remarks on, 279
plants subject to, 280
Chorisis, 59, 343
parallel, 344
collateral, 344
Chromatism, 339
Cladodes, 328
Classification, teratology in relation to, 488
Cohesion of leaves, 21, 25
of petals, 28
pistils, 29
sepals, 27
stamens, 29
stems, 9
Colour, alterations of, 337
Columella, 395
Coniferæ, leaves of, 217, 352, 484
inflorescence of, 245
scales of, 192, 245, 484
Compensation, 488
Page 629
Connective, petalody of, 293
Consistence, alterations of, 432
Contabescence, 463
Contortion, 317
Co-relation, 486
Cornute leaves, 328
Corolla, abortion of, 461
dialysis of, 71
duplicate, 376
hose in hose, 377
meiophylly of, 397
meiotaxy of, 403
pleiotaxy of, 374
polyphylly of, 359
virescence of, 338
Cotyledons, increased number of, 370
shoots, below, 161
Cuttings, formation of, 159
D.
Dédoublement, 59 (see Chorisis)
Deflexion, 209
Deformities, 311
Degeneration, 470
Depauperate ferns, 466
Diadelphia, 29
Dialysis, 58, 69
of calyx, 71
carpels, 73
corolla, 72
plants subject to, 72
of leaves—margins of, 70
stamens, 73
Diaphysis, 103
Diœcious plants, 192, 193
Dimorphism, 333
Dimorphic flowers, 403
Direction, changes of, 201
Diremption, 87
Disjunction, see Dialysis, Fission, Solution
Displacement, 84
of bulbs, 84
carpels, 96
flower-parts of, 91
inflorescence, 85
leaves, 87
Consistence, alterations of, 432
Contabescence, 463
Contortion, 317
Co-relation, 486
Cornute leaves, 328
Corolla, abortion of, 461
dialysis of, 71
duplicate, 376
hose in hose, 377
meiophylly of, 397
meiotaxy of, 403
pleiotaxy of, 374
polyphylly of, 359
virescence of, 338
Cotyledons, increased number of, 370
shoots, below, 161
Cuttings, formation of, 159
D.
Dédoublement, 59 (see Chorisis)
Deflexion, 209
Deformities, 311
Degeneration, 470
Depauperate ferns, 466
Diadelphia, 29
Dialysis, 58, 69
of calyx, 71
carpels, 73
corolla, 72
plants subject to, 72
of leaves—margins of, 70
stamens, 73
Diaphysis, 103
Diœcious plants, 192, 193
Dimorphism, 333
Dimorphic flowers, 403
Direction, changes of, 201
Diremption, 87
Disjunction, see Dialysis, Fission, Solution
Displacement, 84
of bulbs, 84
carpels, 96
flower-parts of, 91
inflorescence, 85
leaves, 87
Page 630
ovules, 96
placentas, 96
Distension, 419
E.
Ecblastesis, 107, 138
Elongation, 433
of parts of flower, 438
flower-stalks, 435
inflorescence, 434
leaves, 437
nucleus of ovule, 269
placenta, 440
receptacle, 440
root, 434
thalamus, 440
Embryos, adhesion of, 56
increased number of, 369
Enation, 443
from axile organs, 444
carpels, 453
corolla, 449
foliar organs, 445
sepals, 448
stamens, 453
Enlargement, 417
of andrœcium, 430
axile organs, 418
buds, 420
flower stalk, 421
fruit, 431
gynœcium, 430
perianth, 428
placenta, 425
Epanody, 226
Epistrophy, 226
Etiolation, 337
Eversion, 204
Excrescences, 444 (see Enation)
F.
Fasciation, 11
plants affected with, 20, 508
Fastigiation, 202
placentas, 96
Distension, 419
E.
Ecblastesis, 107, 138
Elongation, 433
of parts of flower, 438
flower-stalks, 435
inflorescence, 434
leaves, 437
nucleus of ovule, 269
placenta, 440
receptacle, 440
root, 434
thalamus, 440
Embryos, adhesion of, 56
increased number of, 369
Enation, 443
from axile organs, 444
carpels, 453
corolla, 449
foliar organs, 445
sepals, 448
stamens, 453
Enlargement, 417
of andrœcium, 430
axile organs, 418
buds, 420
flower stalk, 421
fruit, 431
gynœcium, 430
perianth, 428
placenta, 425
Epanody, 226
Epistrophy, 226
Etiolation, 337
Eversion, 204
Excrescences, 444 (see Enation)
F.
Fasciation, 11
plants affected with, 20, 508
Fastigiation, 202
Page 631
Ferns, crested, 63, 447
depauperated, 466
exindusiate, 467
supra-soriferous, 189
Filaments, see Stamens
petaloid, 290
4-winged, 290
Finger and toe, 69
Fission, 59
of carpels, 68
leaf-organs, 61, 66
plants, subject to, 66
of petals, 66
stem-organs, 60
stamens, 68
Flattening, 328
Floral organs, displacement of, 91
elongation of, 439
metamorphy of, 281
Florets, increased number of, 351, 390
Flowers, adventitious, 174
on fruits, 177
on leaves, 174
in ovary, 180
on petals, 177
on spines, 177
apetalous, 404
double, 490, 510
hermaphrodite, 196
homomorphic, 188
increased number of, 390
mutilated, 403
unisexual, 193
Flower-stalk, enlargement of, 421
Flower-bud (see Prolification), replaced by leaves or scales, 164
in place of leaf-buds, 176
Foliar organs (see leaves), adhesion of, 32
Foliar organs, cohesion of, 21, 25
enation from, 445
fission of, 61
suppression of, 396
Form, alterations of, 213
juvenile, persistence of, 217
Frondescence, 241, 279, see Phyllody, Virescence
Fruit, adhesion of, 44
enlargement of, 431
depauperated, 466
exindusiate, 467
supra-soriferous, 189
Filaments, see Stamens
petaloid, 290
4-winged, 290
Finger and toe, 69
Fission, 59
of carpels, 68
leaf-organs, 61, 66
plants, subject to, 66
of petals, 66
stem-organs, 60
stamens, 68
Flattening, 328
Floral organs, displacement of, 91
elongation of, 439
metamorphy of, 281
Florets, increased number of, 351, 390
Flowers, adventitious, 174
on fruits, 177
on leaves, 174
in ovary, 180
on petals, 177
on spines, 177
apetalous, 404
double, 490, 510
hermaphrodite, 196
homomorphic, 188
increased number of, 390
mutilated, 403
unisexual, 193
Flower-stalk, enlargement of, 421
Flower-bud (see Prolification), replaced by leaves or scales, 164
in place of leaf-buds, 176
Foliar organs (see leaves), adhesion of, 32
Foliar organs, cohesion of, 21, 25
enation from, 445
fission of, 61
suppression of, 396
Form, alterations of, 213
juvenile, persistence of, 217
Frondescence, 241, 279, see Phyllody, Virescence
Fruit, adhesion of, 44
enlargement of, 431
Page 632
G.
Gemmæ, formation of, 173
Glands, formation of, 473
Gnaurs, 158, 347, 417, 419
Grafting, 53, 56
Greffe des Charlatans, 56
Growth interrupted, 327
irregular, 228
Gymnaxony, 211
Gynantherus, 305
Gynœcium (see pistils)
enlargement of, 430
meiophylly of, 399
meiotaxy of, 406
pleiotaxy of, 388
polyphylly of, 363
suppression of, 406
H.
Hairs, formation of, 472
Hermaphroditism, 197
Heterogamy, 190
Heteromorphy, 311
Heterophylly, 330
Heterotaxy, 156
Homology, 476
Homomorphy, 188
Hose in hose corollas, 291, 377
Hypertrophy, 415, see Enlargement
I.
Independence, 58
Indusium, abortion of, 467
Inflorescence, displacement of, 84
elongation of, 434
prolification of, 102, 115
Interrupted growth, 327
Inversion of organs, 206
Irregularity, 213
Irregular growth, 228
K.
Gemmæ, formation of, 173
Glands, formation of, 473
Gnaurs, 158, 347, 417, 419
Grafting, 53, 56
Greffe des Charlatans, 56
Growth interrupted, 327
irregular, 228
Gymnaxony, 211
Gynantherus, 305
Gynœcium (see pistils)
enlargement of, 430
meiophylly of, 399
meiotaxy of, 406
pleiotaxy of, 388
polyphylly of, 363
suppression of, 406
H.
Hairs, formation of, 472
Hermaphroditism, 197
Heterogamy, 190
Heteromorphy, 311
Heterophylly, 330
Heterotaxy, 156
Homology, 476
Homomorphy, 188
Hose in hose corollas, 291, 377
Hypertrophy, 415, see Enlargement
I.
Independence, 58
Indusium, abortion of, 467
Inflorescence, displacement of, 84
elongation of, 434
prolification of, 102, 115
Interrupted growth, 327
Inversion of organs, 206
Irregularity, 213
Irregular growth, 228
K.
Page 633
Knaurs, (see gnaurs)
Kail, 426
L.
Laciniation, see fission
Layering, 156
Leaders, formation of, 203
Leaf-sheath, 477
Leaves, see foliar organs
abortion of, 458
adhesion of,
by surfaces, 33
to stem, 34
adventitious, 162-165, 509
cornute, 328
displacement of, 86
elongation of, 437
enlargement of, 421
frondiferous, 355
geminate, 352
multiplication of, 358
nature of, 477
palmate-passage of to pinnate, 439
spiral torsion of, 326
supernumerary, 353
Lily, double white, 375
M.
Meiotaxy of andrœcium, 405
of calyx, 403
corolla, 403
gynœcium, 405
Meiophylly of andrœcium, 398
of calyx, 397
corolla, 397
gynœcium, 399
perianth, 397
Mellarose, 134
Metaphery, 91
Metamorphy, 239, 281
calycanthemy, 283
chloranthy, 273
petalody, 283
phyllody, 241
Kail, 426
L.
Laciniation, see fission
Layering, 156
Leaders, formation of, 203
Leaf-sheath, 477
Leaves, see foliar organs
abortion of, 458
adhesion of,
by surfaces, 33
to stem, 34
adventitious, 162-165, 509
cornute, 328
displacement of, 86
elongation of, 437
enlargement of, 421
frondiferous, 355
geminate, 352
multiplication of, 358
nature of, 477
palmate-passage of to pinnate, 439
spiral torsion of, 326
supernumerary, 353
Lily, double white, 375
M.
Meiotaxy of andrœcium, 405
of calyx, 403
corolla, 403
gynœcium, 405
Meiophylly of andrœcium, 398
of calyx, 397
corolla, 397
gynœcium, 399
perianth, 397
Mellarose, 134
Metaphery, 91
Metamorphy, 239, 281
calycanthemy, 283
chloranthy, 273
petalody, 283
phyllody, 241
Page 634
pistillody, 302
sepalody, 282
staminody, 298
Mischomany, 348
Monadelphia, 29
Monœcious, 192, 193, 509
Monosy, 58
Morphology, 479
Multiplication, see pleiotaxy, pleiophylly
of bracts, 358, 371
bulbs, 350
cotyledons, 370
embryos, 369
florets, 351
foliar organs, 352
whorls, 371
N.
Nepaul Barley, 174
Number, alterations of, 341
increased, 343, 353
diminished, 392
O.
Orchids, andrœcium of, 380
prolification in, 153
Organs, rudimentary, see Atrophy
Ovary, inferior, nature of, 394, 482
solution from calyx, 77
stamens in, 184
Ovules, abortion of, 466
increase of, 367
in place of pollen, 200
polliniferous, 183
petalody of, 297
phyllody of, 262
pistillody of, 310
suppression of, 407
malformations of, 262
bibliography of, 272
P.
sepalody, 282
staminody, 298
Mischomany, 348
Monadelphia, 29
Monœcious, 192, 193, 509
Monosy, 58
Morphology, 479
Multiplication, see pleiotaxy, pleiophylly
of bracts, 358, 371
bulbs, 350
cotyledons, 370
embryos, 369
florets, 351
foliar organs, 352
whorls, 371
N.
Nepaul Barley, 174
Number, alterations of, 341
increased, 343, 353
diminished, 392
O.
Orchids, andrœcium of, 380
prolification in, 153
Organs, rudimentary, see Atrophy
Ovary, inferior, nature of, 394, 482
solution from calyx, 77
stamens in, 184
Ovules, abortion of, 466
increase of, 367
in place of pollen, 200
polliniferous, 183
petalody of, 297
phyllody of, 262
pistillody of, 310
suppression of, 407
malformations of, 262
bibliography of, 272
P.
Page 635
Parasitical plants, 55
Peduncles, elongation of, 435
Peloria, 207, 228
bibliography of, 227, 239
regular, 219
plants subject to, 226
irregular, 229
plants, subject to, 239
Perianth, abortion of, 460
enlargement of, 428
meiophylly of, 396
pistillody of, 303
pleiotaxy of, 375
Persistence, xxxvi adnot., 217
Petalody, 283
of accessory organs, 297
anther, 291
calyx, 283
connective, 292
ovules, 297
pistils, 297
stamens, 284
plants, subject to, 295
Petals, cohesion of, 28
enation from, 448
fission of, 66
phyllody of, 251
staminody of, 298
tubular, 23, 314
Phyllode, 328
Phyllody, 240 (see Virescence, Chloranthy)
of accessory organs, 272
bracts, 242
calyx, 244
plants subject to, 250
in Conifers, 245
of corolla, 251
plants subject to, 252
of ovules, 262
plants subject to, 271
pistils, 256
plants subject to, 261
stamens, 253
plants subject to, 256
Phyllomania, 352
Phyllomorphy, see Phyllody
Phyllotaxy, 1, 320
Pistil, abortion of, 464
Peduncles, elongation of, 435
Peloria, 207, 228
bibliography of, 227, 239
regular, 219
plants subject to, 226
irregular, 229
plants, subject to, 239
Perianth, abortion of, 460
enlargement of, 428
meiophylly of, 396
pistillody of, 303
pleiotaxy of, 375
Persistence, xxxvi adnot., 217
Petalody, 283
of accessory organs, 297
anther, 291
calyx, 283
connective, 292
ovules, 297
pistils, 297
stamens, 284
plants, subject to, 295
Petals, cohesion of, 28
enation from, 448
fission of, 66
phyllody of, 251
staminody of, 298
tubular, 23, 314
Phyllode, 328
Phyllody, 240 (see Virescence, Chloranthy)
of accessory organs, 272
bracts, 242
calyx, 244
plants subject to, 250
in Conifers, 245
of corolla, 251
plants subject to, 252
of ovules, 262
plants subject to, 271
pistils, 256
plants subject to, 261
stamens, 253
plants subject to, 256
Phyllomania, 352
Phyllomorphy, see Phyllody
Phyllotaxy, 1, 320
Pistil, abortion of, 464
Page 636
cohesion of, 29
petalody of, 296
staminody of, 298
Pistillody, 302
of ovules, 268, 310
perianth, 302
sepals, 302
stamens, 303
plants subject to, 310
Pitchers, 30, 313, 509, see Ascidia
Placentation, changes in, 96, 508
bibliography, 100
nature of, 483
Placenta, elongation of, 439
enlargement of, 421
Pleiomorphy, 228
Pleiophylly, 353
Pleiotaxy, 371
of andrœcium, 375
bracts, 371
calyx, 374
corolla, 376
gynœcium, 388
perianth, 375
Plien, 346
Plymouth Strawberry, 275
Polyadelphia, 29
Pollen in ovules, 183
abortion of, 463
replaced by ovules, 200
Polyclady, 346
Polycotyledony, 370
Polyembryony, 369
Polymorphy, 328
Polyphylly of andrœcium, 361
of calyx, 350
corolla, 359
plants subject to, 360
of flower, 363
gynœcium, 363
Polyphylly, bibliography, 364
plants subject to, 364
Position, changes of, 83
relative, 485
Prolification, 100
axillary, 138
foliar, 141
floral, 142
petalody of, 296
staminody of, 298
Pistillody, 302
of ovules, 268, 310
perianth, 302
sepals, 302
stamens, 303
plants subject to, 310
Pitchers, 30, 313, 509, see Ascidia
Placentation, changes in, 96, 508
bibliography, 100
nature of, 483
Placenta, elongation of, 439
enlargement of, 421
Pleiomorphy, 228
Pleiophylly, 353
Pleiotaxy, 371
of andrœcium, 375
bracts, 371
calyx, 374
corolla, 376
gynœcium, 388
perianth, 375
Plien, 346
Plymouth Strawberry, 275
Polyadelphia, 29
Pollen in ovules, 183
abortion of, 463
replaced by ovules, 200
Polyclady, 346
Polycotyledony, 370
Polyembryony, 369
Polymorphy, 328
Polyphylly of andrœcium, 361
of calyx, 350
corolla, 359
plants subject to, 360
of flower, 363
gynœcium, 363
Polyphylly, bibliography, 364
plants subject to, 364
Position, changes of, 83
relative, 485
Prolification, 100
axillary, 138
foliar, 141
floral, 142
Page 637
plants affected with, 148
bibliography of, 154
complicated, 151
of embryo, 155
of flower, 115
coincident changes, 128
median foliar, 116
median floral, 119, 508
plants affected with, 137
of fruit, 134
inflorescence, 102
bibliography, 115
median foliar, 103
median floral, 105
lateral floral, 107
lateral foliar, 106
R.
Receptacle, abortion of, 457
elongation of, 116, 440
spiral torsion of, 325
Reflexion, 209
Regularity, 213
Rhizotaxy, 1
Rose Willow, 166, 168
Roots, adventitious, 156
elongation of, 434
Rudimentary organs, 469
S.
Saint Valery Apple, 135, 282, 304, 375, 388
Savoys, 426
Scales, formation of, 164, 448, 470
Scape, leaves on, 163
Seeds, abortion of, 407
union of, 50
Sepals, adhesion of, to petals, 34
cohesion of, 27
enation from, 448
phyllody of, 243
pistillody of, 303
staminody of, 298
Sepalody, 282
Separation, 58
bibliography of, 154
complicated, 151
of embryo, 155
of flower, 115
coincident changes, 128
median foliar, 116
median floral, 119, 508
plants affected with, 137
of fruit, 134
inflorescence, 102
bibliography, 115
median foliar, 103
median floral, 105
lateral floral, 107
lateral foliar, 106
R.
Receptacle, abortion of, 457
elongation of, 116, 440
spiral torsion of, 325
Reflexion, 209
Regularity, 213
Rhizotaxy, 1
Rose Willow, 166, 168
Roots, adventitious, 156
elongation of, 434
Rudimentary organs, 469
S.
Saint Valery Apple, 135, 282, 304, 375, 388
Savoys, 426
Scales, formation of, 164, 448, 470
Scape, leaves on, 163
Seeds, abortion of, 407
union of, 50
Sepals, adhesion of, to petals, 34
cohesion of, 27
enation from, 448
phyllody of, 243
pistillody of, 303
staminody of, 298
Sepalody, 282
Separation, 58
Page 638
Sex, changes of, 190, 509
Shamrock, four-leaved, 356
Shoots below cotyledons, 167; see Leaders
Size, alterations in, 411
Solenaidie, 21, 316
Solution, 59, 76
bibliography, 82
of calyx, 77
plants subject to, 82
of stamens, 82
Spathes, increased number of, 357
Speiranthy, 91, 325
Spiral torsion, 319
plants subject to, 325
of leaf, 326
of receptacle, 324
Spines, 456
Sports, 336
Spurs, formation of, 228, 315
Stamens, see Andrœcium
abortion of, 463
adhesion of, 34, 35
cohesion of, 29
compound, 294, 345
dialysis, 73
enation from, 453
fission of, 68
in ovary, 183
petalody of, 283
phyllody of, 253
pistillody of, 303
tubular, 316
Staminody, of accessory organs, 301
of bracts, 298
petals, 298
pistils, 299
sepals, 298
Stasimorphy, 216
Stem, see axes
Stipules, increased number of, 357
Strawberry, Plymouth, 275
blind, 195
Suppression, 393
of andrœcium, 405
flower, 408
foliar organs, 395
Suppression of ovules, 407
remarks on, 409
Shamrock, four-leaved, 356
Shoots below cotyledons, 167; see Leaders
Size, alterations in, 411
Solenaidie, 21, 316
Solution, 59, 76
bibliography, 82
of calyx, 77
plants subject to, 82
of stamens, 82
Spathes, increased number of, 357
Speiranthy, 91, 325
Spiral torsion, 319
plants subject to, 325
of leaf, 326
of receptacle, 324
Spines, 456
Sports, 336
Spurs, formation of, 228, 315
Stamens, see Andrœcium
abortion of, 463
adhesion of, 34, 35
cohesion of, 29
compound, 294, 345
dialysis, 73
enation from, 453
fission of, 68
in ovary, 183
petalody of, 283
phyllody of, 253
pistillody of, 303
tubular, 316
Staminody, of accessory organs, 301
of bracts, 298
petals, 298
pistils, 299
sepals, 298
Stasimorphy, 216
Stem, see axes
Stipules, increased number of, 357
Strawberry, Plymouth, 275
blind, 195
Suppression, 393
of andrœcium, 405
flower, 408
foliar organs, 395
Suppression of ovules, 407
remarks on, 409
Page 639
of seeds, 407
Symmetry, 213
Synanthy, 37
bibliography, 45
plants subject to, 44, 508
Syncarpy, 45
Syngenesia, 29
Synophty, 57
Synspermy, 50
T.
Tendrils, adventitious, 326
formation of, 473
Thalamus, see Receptacle
Thorns, 456
Torsion spiral, 319
Tubers, 421
in axils of leaves, 142
Tubes, formation of, 312, 509 (see Ascidia, Solenaidy, Spurs)
Tubular petals, 314
stamens, 316
U.
Union, 8
Unisexuality, 195
Uovoli, 420
V.
Varieties, dwarf, 411
Venation, 338
Virescence, 338
Viviparous plants, 106, 168
W.
Warts, 444
Wheat-ear carnation, 371
Symmetry, 213
Synanthy, 37
bibliography, 45
plants subject to, 44, 508
Syncarpy, 45
Syngenesia, 29
Synophty, 57
Synspermy, 50
T.
Tendrils, adventitious, 326
formation of, 473
Thalamus, see Receptacle
Thorns, 456
Torsion spiral, 319
Tubers, 421
in axils of leaves, 142
Tubes, formation of, 312, 509 (see Ascidia, Solenaidy, Spurs)
Tubular petals, 314
stamens, 316
U.
Union, 8
Unisexuality, 195
Uovoli, 420
V.
Varieties, dwarf, 411
Venation, 338
Virescence, 338
Viviparous plants, 106, 168
W.
Warts, 444
Wheat-ear carnation, 371
Page 640
INDEX OF NAMES OF PLANTS.
[In the following Index the names of the orders that are incidentally
mentioned are printed in small capitals, those of the genera and species in
ordinary type. The names are inserted as found in the several records, &c.,
without in general any attempt having been made to determine their
accuracy. For this reason the authority for the specific name is rarely given,
such citations being here unnecessary if not impracticable. It may, however,
be assumed that the names made use of are those generally adopted by
naturalists.
This index will be found useful for statistical purposes. It will show at a
glance, at least approximately, how often certain genera and species are
affected with malformation, as contrasted with others. The nature of the
malformation may of course be ascertained by referring to the particular
page indicated by the number. The proportion of wild to cultivated plants
may also be approximately ascertained, and the effects of cultivation
estimated. The disproportionate frequency with which some species are
affected, e.g., Trifolium repens, &c., as contrasted with other closely allied,
and perhaps equally common species, under apparently identical conditions,
is also made manifest.]
A.
[In the following Index the names of the orders that are incidentally
mentioned are printed in small capitals, those of the genera and species in
ordinary type. The names are inserted as found in the several records, &c.,
without in general any attempt having been made to determine their
accuracy. For this reason the authority for the specific name is rarely given,
such citations being here unnecessary if not impracticable. It may, however,
be assumed that the names made use of are those generally adopted by
naturalists.
This index will be found useful for statistical purposes. It will show at a
glance, at least approximately, how often certain genera and species are
affected with malformation, as contrasted with others. The nature of the
malformation may of course be ascertained by referring to the particular
page indicated by the number. The proportion of wild to cultivated plants
may also be approximately ascertained, and the effects of cultivation
estimated. The disproportionate frequency with which some species are
affected, e.g., Trifolium repens, &c., as contrasted with other closely allied,
and perhaps equally common species, under apparently identical conditions,
is also made manifest.]
A.
Page 641
Abies Brunoniana, 245
excelsa, 21, 61, 192, 298, 325, 456,
Larix, 90
pectinata, 52
Acacia, 329
Aceras anthropophora, 238
Acer, 359, 364, 367, 508
platanoides, 66, 459
pseudo-platanus, 20, 371
Aceranthus, 225, 226
Achimenes, 106, 114, 296
longiflora, 506
Aconitum, 148, 231, 359, 360, 399, 404, 407, 464
Napellus, 39, 44, 238
Acorus, 225
Actæa spicata, 251, 252
Adenium obesum, 417
Adenorophium luxurians, 254
Adonis, 262, 295
autumnalis, 500
vernalis, 500
Adoxa, 367
Ærides odoratum, 398
Æschynanthus, 44, 297
Æsculus Hippocastanum, 50, 66, 295, 369, 438, 459, 501
Affonsea, 364
Afzelia, 397
Agaricus, xxiii, 54
Agave, 177, 432
Americana, 33, 45
Angelica, 365
Agrimonia, 137, 406
Agrostemma, 148
Githago, 271
Agrostis alba, 169
Ailanthus glandulosa, 21
Aira alpina, 169
cæspitosa, 169
Ajuga Iva, 404
pyramidalis, 20
reptans, 243
Alcea, 149, 297
Alchemilla minima, 171
Aldrovanda vesiculosa, 86
Alisma, 115, 329
natans, 107
parnassifolia, 167
Alismaceæ, 115
excelsa, 21, 61, 192, 298, 325, 456,
Larix, 90
pectinata, 52
Acacia, 329
Aceras anthropophora, 238
Acer, 359, 364, 367, 508
platanoides, 66, 459
pseudo-platanus, 20, 371
Aceranthus, 225, 226
Achimenes, 106, 114, 296
longiflora, 506
Aconitum, 148, 231, 359, 360, 399, 404, 407, 464
Napellus, 39, 44, 238
Acorus, 225
Actæa spicata, 251, 252
Adenium obesum, 417
Adenorophium luxurians, 254
Adonis, 262, 295
autumnalis, 500
vernalis, 500
Adoxa, 367
Ærides odoratum, 398
Æschynanthus, 44, 297
Æsculus Hippocastanum, 50, 66, 295, 369, 438, 459, 501
Affonsea, 364
Afzelia, 397
Agaricus, xxiii, 54
Agave, 177, 432
Americana, 33, 45
Angelica, 365
Agrimonia, 137, 406
Agrostemma, 148
Githago, 271
Agrostis alba, 169
Ailanthus glandulosa, 21
Aira alpina, 169
cæspitosa, 169
Ajuga Iva, 404
pyramidalis, 20
reptans, 243
Alcea, 149, 297
Alchemilla minima, 171
Aldrovanda vesiculosa, 86
Alisma, 115, 329
natans, 107
parnassifolia, 167
Alismaceæ, 115
Page 642
Allamanda, 296
cathartica, 288, 390, 505
Allium, 106, 114, 170, 299, 360, 365, 367
fragrans, 369
vineale, 150
Alliaria officinalis, 269, 271
Almond, see Amygdalus.
Alnus, 349
fruticosa, 192
imperialis, 459
incana, 21
glutinosa, 66, 244, 349
laciniata, 65
Alopecurus pratensis, 169
Alsine media, 67, 252, 404
Alstrœmeria, 319, 326
Althæa, 295
rosea, 20, 501
Alyssum, 137
incanum, 252
Amaryllidaceæ, 115, 138, 150
Amaryllis, 307, 310, 432
Ambrina ambrosioides, 397
Amentaceæ, 114, 435
Amorpha, 21, 397
fruticosa, 23, 30, 243
Amygdaleæ, 500; see Rosaceæ
Amygdalus, 122, 137, 155, 295, 297, 364
communis, 250, 252, 369, 503
Persica, 74, 176, 503
Anagallis, 73, 138, 141, 150, 288, 296, 360, 365, 367
arvensis, 117, 161, 256, 263, 271, 278, 281, 284
collina, 44,
phœnicea, 141, 253, 271, 441, 461
tenella, 505
Webbiana, 281
Anomodon alternatus, 49
Ananassa, 350
Anchusa, 132, 138, 339, 365
ochroleuca, 125, 259, 262, 281
paniculata, 262
Androsace maxima, 16, 20
Anemiopsis californica, 63, 66
Anemone, 113, 121, 136, 140, 142, 148, 258, 289, 295, 296, 297, 360, 374, 430, 463, 491, (see
Hepatica)
coronaria, 107, 250, 256, 339, 499
hortensis, 107, 250, 339, 499
japonica, 161, 499
cathartica, 288, 390, 505
Allium, 106, 114, 170, 299, 360, 365, 367
fragrans, 369
vineale, 150
Alliaria officinalis, 269, 271
Almond, see Amygdalus.
Alnus, 349
fruticosa, 192
imperialis, 459
incana, 21
glutinosa, 66, 244, 349
laciniata, 65
Alopecurus pratensis, 169
Alsine media, 67, 252, 404
Alstrœmeria, 319, 326
Althæa, 295
rosea, 20, 501
Alyssum, 137
incanum, 252
Amaryllidaceæ, 115, 138, 150
Amaryllis, 307, 310, 432
Ambrina ambrosioides, 397
Amentaceæ, 114, 435
Amorpha, 21, 397
fruticosa, 23, 30, 243
Amygdaleæ, 500; see Rosaceæ
Amygdalus, 122, 137, 155, 295, 297, 364
communis, 250, 252, 369, 503
Persica, 74, 176, 503
Anagallis, 73, 138, 141, 150, 288, 296, 360, 365, 367
arvensis, 117, 161, 256, 263, 271, 278, 281, 284
collina, 44,
phœnicea, 141, 253, 271, 441, 461
tenella, 505
Webbiana, 281
Anomodon alternatus, 49
Ananassa, 350
Anchusa, 132, 138, 339, 365
ochroleuca, 125, 259, 262, 281
paniculata, 262
Androsace maxima, 16, 20
Anemiopsis californica, 63, 66
Anemone, 113, 121, 136, 140, 142, 148, 258, 289, 295, 296, 297, 360, 374, 430, 463, 491, (see
Hepatica)
coronaria, 107, 250, 256, 339, 499
hortensis, 107, 250, 339, 499
japonica, 161, 499
Page 643
nemorosa, 250, 256, 339, 499
palmata, 499
Anemone pavonina, 499
Pulsatilla, 250
rivularis, 165
sylvestris, 250, 499
Angelica, 114, 137, 143, 150
Razoulzii, 244, 437
Anthemis arvensis, 20
nobilis, 20
retusa, 44
Anthoxanthum, 61
odoratum, 61
Anthriscus, 442
Anthurium Scherzerianum, 358
Anthyllis, 295
Antirrhinum majus, 20, 22, 24, 31, 44, 57, 73, 82, 104, 114, 121, 131, 225, 226, 227, 230, 233,
235, 238, 253, 272, 296, 301, 315, 316, 318, 363, 365, 378, 398, 505
Apargia, 114
autumnalis, 20
Apium, 82, 113, 150
graveolens, 66, 158
Petroselinum, 370, 437
Apocynaceæ, 137
Apple, St. Valery, 135, 282, 304, 375, 388
Aquilegia canadensis, 500
Skinneri, 266, 271
vulgaris, 24, 70, 127, 136, 220, 226, 252, 257, 260, 261, 271, 280, 286, 287, 288, 293,
295, 74, 390, 500
Arabis, 148, 508
alpina, 397, 461, 463
pumila, 170
sagittata, 44
Araucaria, 245
Arbutus Unedo, 291, 292, 296, 377, 504
Archidendron, 365
Ardisia serrulata, 369
Arenaria serpyllifolia, 461
tetraquetra, 398, 399, 464, 469
Armeria, 114
Aristolochia Clematitis, 38, 45, 314
caudata, 231, 238
sipho, 23
Armoracia rusticana, 64, 299
Arnoseris, 114
Artabotrys, 456, 508
Artemisia, 405
Artocarpus, 407
palmata, 499
Anemone pavonina, 499
Pulsatilla, 250
rivularis, 165
sylvestris, 250, 499
Angelica, 114, 137, 143, 150
Razoulzii, 244, 437
Anthemis arvensis, 20
nobilis, 20
retusa, 44
Anthoxanthum, 61
odoratum, 61
Anthriscus, 442
Anthurium Scherzerianum, 358
Anthyllis, 295
Antirrhinum majus, 20, 22, 24, 31, 44, 57, 73, 82, 104, 114, 121, 131, 225, 226, 227, 230, 233,
235, 238, 253, 272, 296, 301, 315, 316, 318, 363, 365, 378, 398, 505
Apargia, 114
autumnalis, 20
Apium, 82, 113, 150
graveolens, 66, 158
Petroselinum, 370, 437
Apocynaceæ, 137
Apple, St. Valery, 135, 282, 304, 375, 388
Aquilegia canadensis, 500
Skinneri, 266, 271
vulgaris, 24, 70, 127, 136, 220, 226, 252, 257, 260, 261, 271, 280, 286, 287, 288, 293,
295, 74, 390, 500
Arabis, 148, 508
alpina, 397, 461, 463
pumila, 170
sagittata, 44
Araucaria, 245
Arbutus Unedo, 291, 292, 296, 377, 504
Archidendron, 365
Ardisia serrulata, 369
Arenaria serpyllifolia, 461
tetraquetra, 398, 399, 464, 469
Armeria, 114
Aristolochia Clematitis, 38, 45, 314
caudata, 231, 238
sipho, 23
Armoracia rusticana, 64, 299
Arnoseris, 114
Artabotrys, 456, 508
Artemisia, 405
Artocarpus, 407
Page 644
Arum, 329
maculatum, 66, 225, 227, 245, 358
Asparagus officinalis, 12, 19, 21, 320, 325, 471
Asphodelus, 138, 296
Asphodelus luteus, 506
ramosus, 298, 310
Asplenium Trichomanes, 190
Astrantia, 114
major, 368
Athamanta, 82, 137, 149
Cervaria, 250, 442
Atriplex, 227
hortensis, 224
Atropa Belladonna, 44
Aucuba, 21
japonica, 21, 197
Aurantiaceæ, 137, 149, 502
Avena, 319, 351, 391
chinensis, 298
Azalea, 35, 114
glauca, 504
indica, 44, 73, 166, 209, 289, 296
nudiflora, 504
B.
maculatum, 66, 225, 227, 245, 358
Asparagus officinalis, 12, 19, 21, 320, 325, 471
Asphodelus, 138, 296
Asphodelus luteus, 506
ramosus, 298, 310
Asplenium Trichomanes, 190
Astrantia, 114
major, 368
Athamanta, 82, 137, 149
Cervaria, 250, 442
Atriplex, 227
hortensis, 224
Atropa Belladonna, 44
Aucuba, 21
japonica, 21, 197
Aurantiaceæ, 137, 149, 502
Avena, 319, 351, 391
chinensis, 298
Azalea, 35, 114
glauca, 504
indica, 44, 73, 166, 209, 289, 296
nudiflora, 504
B.
Page 645
Babingtonia, 185
Bæckea diosmifolia, 183
Balsamineæ, 359, 404, 501
Balsam, see Impatiens
Bamboo, see Bambusa
Bambusa, 307, 310, 324, 365
Barbarea, 295
vulgaris, 310, 500
Barkhausia, taraxacifolia, 20
Barley, Nepaul, 174
Bauhinia, 328
Begoniaceæ, 114
Begonia, 31, 81, 106, 114, 162, 170, 352
frigida, 199, 303, 310
fuchsioides, 281
phyllomaniaca, 170
Bellevalia, 408, 461, 467
comosa, 348
Bellis perennis, 17, 20, 31, 106, 114, 164, 244
Berberis, 272, 295, 458, 460, 500
vulgaris, 20
Beta, 19, 325, 365
vulgaris, 20
Betonica Alopecuros, 42, 44, 226
Betula alba, 66, 346, 456, 472
populifolia, 66
Bidens, 114, 165, 223
Bignoniaceæ, 222
Bignonia, 272, 296, 327
Bikkhia, 80
Blitum, 45
polymorphum, 397, 458
Bocconia, 224
cordata, 310
Bowiea volubilis, 409
Boraginaceæ, 132, 138
Bouchea hyderabadensis, 166
Bougainvillea, 339
Brachythecium plumosum, 49
Brassica, 20, 136, 139, 295, 364, 367
Napus, 27, 205
oleracea, 30, 33, 66, 67, 106, 113, 136, 141, 142, 148, 250, 252, 264, 271, 280, 351,
408, 421, 426, 442, 445, 501, 508
Rapa, 181
Breynia, 198
Bromelia, 103
Bromus velutinus, 358
Bæckea diosmifolia, 183
Balsamineæ, 359, 404, 501
Balsam, see Impatiens
Bamboo, see Bambusa
Bambusa, 307, 310, 324, 365
Barbarea, 295
vulgaris, 310, 500
Barkhausia, taraxacifolia, 20
Barley, Nepaul, 174
Bauhinia, 328
Begoniaceæ, 114
Begonia, 31, 81, 106, 114, 162, 170, 352
frigida, 199, 303, 310
fuchsioides, 281
phyllomaniaca, 170
Bellevalia, 408, 461, 467
comosa, 348
Bellis perennis, 17, 20, 31, 106, 114, 164, 244
Berberis, 272, 295, 458, 460, 500
vulgaris, 20
Beta, 19, 325, 365
vulgaris, 20
Betonica Alopecuros, 42, 44, 226
Betula alba, 66, 346, 456, 472
populifolia, 66
Bidens, 114, 165, 223
Bignoniaceæ, 222
Bignonia, 272, 296, 327
Bikkhia, 80
Blitum, 45
polymorphum, 397, 458
Bocconia, 224
cordata, 310
Bowiea volubilis, 409
Boraginaceæ, 132, 138
Bouchea hyderabadensis, 166
Bougainvillea, 339
Brachythecium plumosum, 49
Brassica, 20, 136, 139, 295, 364, 367
Napus, 27, 205
oleracea, 30, 33, 66, 67, 106, 113, 136, 141, 142, 148, 250, 252, 264, 271, 280, 351,
408, 421, 426, 442, 445, 501, 508
Rapa, 181
Breynia, 198
Bromelia, 103
Bromus velutinus, 358
Page 646
Broussonettia papyrifera, 331, 459
Bruniaceæ, 80, 81
Brunia microphylla, 81
Bryonia, 360, 367
Bryophyllum calycinum, 158, 171, 483
proliferum, 166
Bryum cæspititium, 49
Bunias, 97, 136, 280, 360
Bunium creticum, 159
flexuosum, 20
Bupleurum, 149, 459
falcatum, 15, 20
Butomaceæ, 507
Buxbaumia indusiata, 49
Byrsonima, 137
Byttneriaceæ, 362
C.
Cabomba aquatica, 458
Cactaceæ, 81, 113, 149, 395
Cactus, 160
Cachrys taurica, 197
Cæsalpinia, 365
digyna, 48
Cakile maritima, 246, 250
Calanthe, 227, 398, 402, 508
vestita, 39, 45, 227, 402, 508
Calceolaria, 41, 44, 230, 233, 284, 296, 397, 405, 406, 505
crenatifolia, 238
floribunda, 316
rugosa, 238
Calendula, 114, 138, 339, 370
officinalis, 252, 280, 339
Calla palustris, 357
Callitriche, 196
autumnalis, 406
vernalis, 406
Caltha, 136, 148, 295
palustris, 141, 250, 442, 500
Calluna, 296, 504
Calycophyllum, 249, 283, 429
Calystegia, 114, 296
Calystegia Sepium, 505
pubescens, 505
Camellia japonica, 288, 295, 297, 491, 494, 502
reticulata, 502
Bruniaceæ, 80, 81
Brunia microphylla, 81
Bryonia, 360, 367
Bryophyllum calycinum, 158, 171, 483
proliferum, 166
Bryum cæspititium, 49
Bunias, 97, 136, 280, 360
Bunium creticum, 159
flexuosum, 20
Bupleurum, 149, 459
falcatum, 15, 20
Butomaceæ, 507
Buxbaumia indusiata, 49
Byrsonima, 137
Byttneriaceæ, 362
C.
Cabomba aquatica, 458
Cactaceæ, 81, 113, 149, 395
Cactus, 160
Cachrys taurica, 197
Cæsalpinia, 365
digyna, 48
Cakile maritima, 246, 250
Calanthe, 227, 398, 402, 508
vestita, 39, 45, 227, 402, 508
Calceolaria, 41, 44, 230, 233, 284, 296, 397, 405, 406, 505
crenatifolia, 238
floribunda, 316
rugosa, 238
Calendula, 114, 138, 339, 370
officinalis, 252, 280, 339
Calla palustris, 357
Callitriche, 196
autumnalis, 406
vernalis, 406
Caltha, 136, 148, 295
palustris, 141, 250, 442, 500
Calluna, 296, 504
Calycophyllum, 249, 283, 429
Calystegia, 114, 296
Calystegia Sepium, 505
pubescens, 505
Camellia japonica, 288, 295, 297, 491, 494, 502
reticulata, 502
Page 647
Sasanqua, 502
Campanulaceæ, 80, 114, 127, 131, 138, 139, 150, 334
Campanula, 71, 72, 73, 82, 138, 150, 250, 285, 296, 365, 367, 404, 442, 472
canescens, 403
colorata, 403
glomerata, 242, 300, 504
latifolia, 504
Medium, 20, 37, 44, 61, 251, 448, 504
persicifolia, 44, 284, 300, 429, 504
pyramidalis, 281, 504
Rapunculus, 429
rapunculoides, 20, 252, 300, 310, 375
rhomboidea, 504
rotundifolia, 377, 378, 504
Tenorei, 504
thyrsoidea, 20
Trachelium, 504
Vidallii, 504
Campanumæa, 80, 81
Camphorosma monspeliaca, 456
Canna, 285
Cannabis, 82, 194, 197
sativa, 81
Cannamois virgata, 197
Capparidaceæ, 148, 390
Caprifoliaceæ, 45
Capsella bursa pastoris, 298, 361
Cardamine, 295, 357
hirsuta, 398
Impatiens, 404
pratensis, 65, 170, 181, 495, 500
sylvatica, 398
Carduus arvensis, 20
crispus, 166, 339
heterophyllus, 260, 262, 250
tataricus, 250, 260, 262
Carex, 115, 138, 150, 191, 194, 350
acuta, 143, 198
cæspitosa, 199
glauca, 143, 199
maritima, 369
paludosa, 199
vulpina, 428
Carica Papaya, 199
Carlemannia, 398
Carlina, 114
vulgaris, 20
Carpinus, 346
Campanulaceæ, 80, 114, 127, 131, 138, 139, 150, 334
Campanula, 71, 72, 73, 82, 138, 150, 250, 285, 296, 365, 367, 404, 442, 472
canescens, 403
colorata, 403
glomerata, 242, 300, 504
latifolia, 504
Medium, 20, 37, 44, 61, 251, 448, 504
persicifolia, 44, 284, 300, 429, 504
pyramidalis, 281, 504
Rapunculus, 429
rapunculoides, 20, 252, 300, 310, 375
rhomboidea, 504
rotundifolia, 377, 378, 504
Tenorei, 504
thyrsoidea, 20
Trachelium, 504
Vidallii, 504
Campanumæa, 80, 81
Camphorosma monspeliaca, 456
Canna, 285
Cannabis, 82, 194, 197
sativa, 81
Cannamois virgata, 197
Capparidaceæ, 148, 390
Caprifoliaceæ, 45
Capsella bursa pastoris, 298, 361
Cardamine, 295, 357
hirsuta, 398
Impatiens, 404
pratensis, 65, 170, 181, 495, 500
sylvatica, 398
Carduus arvensis, 20
crispus, 166, 339
heterophyllus, 260, 262, 250
tataricus, 250, 260, 262
Carex, 115, 138, 150, 191, 194, 350
acuta, 143, 198
cæspitosa, 199
glauca, 143, 199
maritima, 369
paludosa, 199
vulpina, 428
Carica Papaya, 199
Carlemannia, 398
Carlina, 114
vulgaris, 20
Carpinus, 346
Page 648
Betulus, 66
Carthamus, 138
Carum, 82, 114, 365
Bulbocastanum, 159
Carui, 244, 271, 285, 339, 437
Caryophyllaceæ, 99, 113, 120, 137, 139, 140, 148, 250, 379, 397, 398, 404, 406, 407, 410,
418, 442, 443, 448
Casuarina rigida, 325
Cassia, 364, 369
marylandica, 30
Castanea vesca, 11, 66, 104, 114, 319, 435
Catabrosa aquatica, 351
Catalpa, 399
Catasetum, 291, 296, 334
eburneum, 384
Cattleya amethystina, 401
Forbesii, 34, 384, 398
marginata, 223, 227
Mossiæ, 224, 227, 238
violacea, 383, 397
Caucalis leptophylla, 33
Caulophyllum, 75, 125
Caylussa, 137
Cedrus Libani, 61
Celastraceæ, 149
Celastrus, 149
Celosia, 19, 20
Centaurea, 37, 114
collina, 34
Jacea, 43, 243
Scabiosa, 20
moschata, 44
Centranthus, 247
macrosiphon, 250
ruber, 42, 44
Cephalotus follicularis, 314
Cerastium, 62, 97, 262, 397, 398
glomeratum, 280, 358, 463
tetandrum, 463
triviale, 252, 280
vulgatum, 252, 404
Cerasus, 74, 117, 149, 250, 260, 424, 489
avium, 262
caproniana, 364
Lauro-cerasus, 64, 66, 370
vulgaris, 252, 262
Ceratonia Siliqua, 30
Cercis, 364
Carthamus, 138
Carum, 82, 114, 365
Bulbocastanum, 159
Carui, 244, 271, 285, 339, 437
Caryophyllaceæ, 99, 113, 120, 137, 139, 140, 148, 250, 379, 397, 398, 404, 406, 407, 410,
418, 442, 443, 448
Casuarina rigida, 325
Cassia, 364, 369
marylandica, 30
Castanea vesca, 11, 66, 104, 114, 319, 435
Catabrosa aquatica, 351
Catalpa, 399
Catasetum, 291, 296, 334
eburneum, 384
Cattleya amethystina, 401
Forbesii, 34, 384, 398
marginata, 223, 227
Mossiæ, 224, 227, 238
violacea, 383, 397
Caucalis leptophylla, 33
Caulophyllum, 75, 125
Caylussa, 137
Cedrus Libani, 61
Celastraceæ, 149
Celastrus, 149
Celosia, 19, 20
Centaurea, 37, 114
collina, 34
Jacea, 43, 243
Scabiosa, 20
moschata, 44
Centranthus, 247
macrosiphon, 250
ruber, 42, 44
Cephalotus follicularis, 314
Cerastium, 62, 97, 262, 397, 398
glomeratum, 280, 358, 463
tetandrum, 463
triviale, 252, 280
vulgatum, 252, 404
Cerasus, 74, 117, 149, 250, 260, 424, 489
avium, 262
caproniana, 364
Lauro-cerasus, 64, 66, 370
vulgaris, 252, 262
Ceratonia Siliqua, 30
Cercis, 364
Page 649
siliquastrum, 325
Chamærops humilis, 300
Chærophyllum, 113
Cheiranthus, 121, 131, 136, 148, 295, 364
Cheiri, 20, 33, 35, 36, 250, 252, 271, 404, 427, 500
var. gynantherus, 305, 310
Cheiranthus incanus, 250
Chelone, 361
barbata, 238
Chelidonium majus, 66, 170, 280, 295, 500
Chenopodiaceæ, 397, 405, 406
Chenopodium, 45, 365, 367
glaucum, 397
murale, 428
Quinoa, 62, 66
Vulvaria, 458
Chirita sinensis, 170
Chlorophytum Sternbergianum, 107
Chorozema ilicifolium, 21
Chrysanthemum, 16, 72, 365
indicum, 20, 188, 472
Leucanthemum, 20
Chrysosplenium, 367
Cichorium Intybus, 20, 44
Cicuta virosa, 406
Cinchonaceæ, 429
Cionidium Moorei, 190
Cirsium, 114, 138
arvense, 250, 457
lanceolatum, 20
tricephalodes, 252, 339
Cissus, 211
Cistaceæ, 137
Cistus vaginatus, 473
Citrus, 137, 149, 364, 453
Aurantium, 33, 35, 44, 56, 75, 134, 303, 310, 335, 369, 388, 389, 391, 502
Clarkia, 24, 295
elegans, 177, 503
pulchella, 503
Cleistanthus polystachyus, 198
Clematis, 136, 148, 288, 295, 367
florida, 499
Fortunei, 499
patens, 499
Viticella, 28, 499
Cleome, 137, 148
Cleonia lusitanica, 238
Clerodendron fragrans, 506
Chamærops humilis, 300
Chærophyllum, 113
Cheiranthus, 121, 131, 136, 148, 295, 364
Cheiri, 20, 33, 35, 36, 250, 252, 271, 404, 427, 500
var. gynantherus, 305, 310
Cheiranthus incanus, 250
Chelone, 361
barbata, 238
Chelidonium majus, 66, 170, 280, 295, 500
Chenopodiaceæ, 397, 405, 406
Chenopodium, 45, 365, 367
glaucum, 397
murale, 428
Quinoa, 62, 66
Vulvaria, 458
Chirita sinensis, 170
Chlorophytum Sternbergianum, 107
Chorozema ilicifolium, 21
Chrysanthemum, 16, 72, 365
indicum, 20, 188, 472
Leucanthemum, 20
Chrysosplenium, 367
Cichorium Intybus, 20, 44
Cicuta virosa, 406
Cinchonaceæ, 429
Cionidium Moorei, 190
Cirsium, 114, 138
arvense, 250, 457
lanceolatum, 20
tricephalodes, 252, 339
Cissus, 211
Cistaceæ, 137
Cistus vaginatus, 473
Citrus, 137, 149, 364, 453
Aurantium, 33, 35, 44, 56, 75, 134, 303, 310, 335, 369, 388, 389, 391, 502
Clarkia, 24, 295
elegans, 177, 503
pulchella, 503
Cleistanthus polystachyus, 198
Clematis, 136, 148, 288, 295, 367
florida, 499
Fortunei, 499
patens, 499
Viticella, 28, 499
Cleome, 137, 148
Cleonia lusitanica, 238
Clerodendron fragrans, 506
Page 650
Cliffortia, 396
Clinacium dendroides, 49
Clitoria Ternatea, 231, 238, 295, 502
Clusia rosea, 11
Cluytia semperflorens, 198
Cneorum, 365
Cnicus palustris, 20
Cnidium, 113
Cobæa scandens, 73, 82, 272, 326, 365
Coccoloba platycladon, 328
Cochlearia Armoracia, 64, 299, 310, 331
Cocos, 115, 365
nucifera, 429
Codiæum variegatum, 31, 314, 326, 328, 459
Cœlebogyne ilicifolia, 369
Cola acuminata, 370
Colchicum autumnale, 45, 67, 73, 250, 287, 296, 406, 407, 507
Coleus, 365, 459
Columnea Schiedeana, 226
Columbine, see Aquilegia
Colutea, 465
Commelyna, 73, 296, 507
Commelynaceæ, 245, 507
Compositæ, 72, 73, 86, 107, 114, 127, 131, 138, 165, 223, 226, 235, 339, 406, 407, 430, 434,
437, 439, 442
Conceveiba macrophylla, 198
Coniferæ, 56, 65, 103, 114, 191, 245, 369, 435
Conium maculatum, 114
Conostephium, 120
Convallaria maialis, 73, 150, 250, 296, 360, 367, 375, 400, 442, 507
Polygonatum, 507
Convolvulaceæ, 114, 137, 150
Convolvulus, 73, 114, 137, 142, 150, 296, 510
arvensis, 20
Sepium, 20, 108, 250
tricolor, 505
Conyza squarrosa, 20
Corallorhiza innata, 238
Circeia, 410
Corchorus acutangulus, 397
Coreopsis, 20, 73, 114, 138
Drummondi, 339
Cornus, 37, 44, 358
mas, 358, 374
sanguinea, 44
suecica, 374
Coronilla, 106, 113, 149, 295
Emerus, 502
Clinacium dendroides, 49
Clitoria Ternatea, 231, 238, 295, 502
Clusia rosea, 11
Cluytia semperflorens, 198
Cneorum, 365
Cnicus palustris, 20
Cnidium, 113
Cobæa scandens, 73, 82, 272, 326, 365
Coccoloba platycladon, 328
Cochlearia Armoracia, 64, 299, 310, 331
Cocos, 115, 365
nucifera, 429
Codiæum variegatum, 31, 314, 326, 328, 459
Cœlebogyne ilicifolia, 369
Cola acuminata, 370
Colchicum autumnale, 45, 67, 73, 250, 287, 296, 406, 407, 507
Coleus, 365, 459
Columnea Schiedeana, 226
Columbine, see Aquilegia
Colutea, 465
Commelyna, 73, 296, 507
Commelynaceæ, 245, 507
Compositæ, 72, 73, 86, 107, 114, 127, 131, 138, 165, 223, 226, 235, 339, 406, 407, 430, 434,
437, 439, 442
Conceveiba macrophylla, 198
Coniferæ, 56, 65, 103, 114, 191, 245, 369, 435
Conium maculatum, 114
Conostephium, 120
Convallaria maialis, 73, 150, 250, 296, 360, 367, 375, 400, 442, 507
Polygonatum, 507
Convolvulaceæ, 114, 137, 150
Convolvulus, 73, 114, 137, 142, 150, 296, 510
arvensis, 20
Sepium, 20, 108, 250
tricolor, 505
Conyza squarrosa, 20
Corallorhiza innata, 238
Circeia, 410
Corchorus acutangulus, 397
Coreopsis, 20, 73, 114, 138
Drummondi, 339
Cornus, 37, 44, 358
mas, 358, 374
sanguinea, 44
suecica, 374
Coronilla, 106, 113, 149, 295
Emerus, 502
Page 651
Correa, 72, 73, 370
Cortusa Mathioli, 133, 138, 263
Corydalis aurea, 280
solida, 243
tuberosa, 235, 336, 238
Corylas Avellana, 21, 31, 48, 66, 114, 349, 354, 364, 365, 368
Cotoneaster microphylla, 21
Cotula fœtida, 19, 20
Crassulaceæ, 113
Crassula, 113
arborescens, 26, 31
Cratægus, 82, 113, 149, 295, 364, 404, 419
Crus galli, 503
monogyna, 44, 400
Cratægus Oxyacantha, 57, 66, 78, 317, 370, 503, 508
tanacetifolia, 107, 162
Crepis, 271
virens, 20
Crinum, 432
Crocus, 29, 35, 45, 67, 287, 289, 296, 319, 361, 399, 400, 434, 462
aureus, 506
nudiflorus, 302
pusillus, 506
vernus, 506
Crozophora tinctoria, 198
Cruciferæ, 73, 76, 98, 113, 136, 139, 141, 148, 257, 297, 364, 379, 406, 410, 428, 442, 500
Cryptomeria japonica, 103, 114, 245, 435
Cucubalus, 149
Cucumis, 36, 82, 138, 248, 259, 326, 367
Cucurbitaceæ, 71, 80, 81, 137, 247
Cucurbita, 197, 201, 250, 307, 310, 365, 474
Cuphea miniata, 211, 424
Cupressus funebris, 218
Cycadeæ, 56, 369
Cyclamen, 67, 104, 114, 296, 319, 359, 360
linearifolium, 329
Cyclodon, 80
Cydonia vulgaris, 71, 79, 295, 423
japonica, 503
Cynanchum fuscatum, 369
nigrum, 369
Cynosurus cristatus, 169
Cyperaceæ, 115, 138, 150, 169, 350
Cypripedium, 27, 92, 381, 386
candidum, 401
Hookeræ, 386
insigne, 91
Cyrtanthus, 177
Cortusa Mathioli, 133, 138, 263
Corydalis aurea, 280
solida, 243
tuberosa, 235, 336, 238
Corylas Avellana, 21, 31, 48, 66, 114, 349, 354, 364, 365, 368
Cotoneaster microphylla, 21
Cotula fœtida, 19, 20
Crassulaceæ, 113
Crassula, 113
arborescens, 26, 31
Cratægus, 82, 113, 149, 295, 364, 404, 419
Crus galli, 503
monogyna, 44, 400
Cratægus Oxyacantha, 57, 66, 78, 317, 370, 503, 508
tanacetifolia, 107, 162
Crepis, 271
virens, 20
Crinum, 432
Crocus, 29, 35, 45, 67, 287, 289, 296, 319, 361, 399, 400, 434, 462
aureus, 506
nudiflorus, 302
pusillus, 506
vernus, 506
Crozophora tinctoria, 198
Cruciferæ, 73, 76, 98, 113, 136, 139, 141, 148, 257, 297, 364, 379, 406, 410, 428, 442, 500
Cryptomeria japonica, 103, 114, 245, 435
Cucubalus, 149
Cucumis, 36, 82, 138, 248, 259, 326, 367
Cucurbitaceæ, 71, 80, 81, 137, 247
Cucurbita, 197, 201, 250, 307, 310, 365, 474
Cuphea miniata, 211, 424
Cupressus funebris, 218
Cycadeæ, 56, 369
Cyclamen, 67, 104, 114, 296, 319, 359, 360
linearifolium, 329
Cyclodon, 80
Cydonia vulgaris, 71, 79, 295, 423
japonica, 503
Cynanchum fuscatum, 369
nigrum, 369
Cynosurus cristatus, 169
Cyperaceæ, 115, 138, 150, 169, 350
Cypripedium, 27, 92, 381, 386
candidum, 401
Hookeræ, 386
insigne, 91
Cyrtanthus, 177
Page 652
Cytisus, 113, 295, 336
albus, 502
Laburnum, 21, 66, 157, 189, 222, 226, 231, 238, 356, 459
nigricans, 15, 104
D.
Dactylis, 115
glomerata, 169
Dahlia, 44, 72, 188, 244, 433
variabilis, 20
Danaë, 60
Daphne indica, 21
odora, 21
Datura, 285, 291, 296, 365, 378, 407
Datura arborea, 505
cornigera, 505
chlorantha, 505
fastuosa, 296, 379, 450, 505
humilis, 505
Daucus Carota, 53, 57, 82, 113, 121, 124, 125, 137, 149, 244, 250, 252, 256, 260, 262, 296,
298, 339, 365, 368, 369, 370, 457, 504
Delphinium, 44, 136, 148, 225, 283, 286, 295, 344, 364, 367, 374, 388, 399, 407
Ajacis, 250, 252, 261, 271, 339, 500
amœnum, 261
cheilanthum, 500
Consolida, 373, 500
crassicaule, 252, 256, 261, 271, 339
dictyocarpum, 271, 432
elatum, 20, 126, 237, 238, 261, 267, 271, 309, 310, 339
elegans, 500
grandiflorum, 500
peregrinum, 219, 226
Dendrobium, 227
nobile, 94, 398
normale, 224, 383
Desmodium canadense, 271
marylandicum, 467
Deutzia, 295
crenata, 503
Dianthus, 67, 113, 121, 129, 137, 139, 145, 146, 149, 166, 261, 268, 289, 295, 297, 310, 360,
364, 371, 379, 397, 471
arboreus, 501
barbatus, 325, 404, 501
Caryophyllus, 501
corymbosus, 501
deltoides, 501
albus, 502
Laburnum, 21, 66, 157, 189, 222, 226, 231, 238, 356, 459
nigricans, 15, 104
D.
Dactylis, 115
glomerata, 169
Dahlia, 44, 72, 188, 244, 433
variabilis, 20
Danaë, 60
Daphne indica, 21
odora, 21
Datura, 285, 291, 296, 365, 378, 407
Datura arborea, 505
cornigera, 505
chlorantha, 505
fastuosa, 296, 379, 450, 505
humilis, 505
Daucus Carota, 53, 57, 82, 113, 121, 124, 125, 137, 149, 244, 250, 252, 256, 260, 262, 296,
298, 339, 365, 368, 369, 370, 457, 504
Delphinium, 44, 136, 148, 225, 283, 286, 295, 344, 364, 367, 374, 388, 399, 407
Ajacis, 250, 252, 261, 271, 339, 500
amœnum, 261
cheilanthum, 500
Consolida, 373, 500
crassicaule, 252, 256, 261, 271, 339
dictyocarpum, 271, 432
elatum, 20, 126, 237, 238, 261, 267, 271, 309, 310, 339
elegans, 500
grandiflorum, 500
peregrinum, 219, 226
Dendrobium, 227
nobile, 94, 398
normale, 224, 383
Desmodium canadense, 271
marylandicum, 467
Deutzia, 295
crenata, 503
Dianthus, 67, 113, 121, 129, 137, 139, 145, 146, 149, 166, 261, 268, 289, 295, 297, 310, 360,
364, 371, 379, 397, 471
arboreus, 501
barbatus, 325, 404, 501
Caryophyllus, 501
corymbosus, 501
deltoides, 501
Page 653
hybridus, 501
plumarius, 501
Poiretianus, 501
sinensis, 370, 501
Dictamnus, 121, 122, 137, 140
albus, 256, 271
Fraxinella, 252, 262, 271, 278, 280
Dielytra, 236, 237
Digitalis lutea, 60
orientalis, 238
purpurea, 20, 40, 44, 73, 98, 121, 129, 137, 150, 226, 233, 238, 296, 298, 315, 365, 373,
398, 459, 472, 505
Dilleniaceæ, 398
Dionæa, 308
Muscipula, 310
Diosma, 369
Diphaca, 365
Diplotaxis, 136, 148, 364
muralis, 252, 458
tenuifolia, 73, 250, 261, 274, 280, 397, 398, 430
Dipsacaceæ, 86, 107, 114, 138
Dipsacus, 419, 429
fullonum, 20, 62, 66, 281, 320, 321, 325
Gmelini, 325
pilosus, 20, 325
sylvestris, 10, 20
Dipterocarpus, 249
Ditaxis lancifolia, 380
Dodecatheon, 138
Dodonæa viscosa, 20
Draba, 364
Dracocephalum austriacum, 238
moldavicum, 20
speciosum, 320, 325
Dracontium pertusum, 25
Drosera intermedia, 170, 265, 271, 277, 473
E.
Ebenus cretica, 26
Eccremocarpus scaber, 222, 226, 326
Echeveria, 113
Echinophora maritima, 252
Echium pyrenaicum, 20
simplex, 20
vulgare, 374
Echinocactus, 149, 178, 417
plumarius, 501
Poiretianus, 501
sinensis, 370, 501
Dictamnus, 121, 122, 137, 140
albus, 256, 271
Fraxinella, 252, 262, 271, 278, 280
Dielytra, 236, 237
Digitalis lutea, 60
orientalis, 238
purpurea, 20, 40, 44, 73, 98, 121, 129, 137, 150, 226, 233, 238, 296, 298, 315, 365, 373,
398, 459, 472, 505
Dilleniaceæ, 398
Dionæa, 308
Muscipula, 310
Diosma, 369
Diphaca, 365
Diplotaxis, 136, 148, 364
muralis, 252, 458
tenuifolia, 73, 250, 261, 274, 280, 397, 398, 430
Dipsacaceæ, 86, 107, 114, 138
Dipsacus, 419, 429
fullonum, 20, 62, 66, 281, 320, 321, 325
Gmelini, 325
pilosus, 20, 325
sylvestris, 10, 20
Dipterocarpus, 249
Ditaxis lancifolia, 380
Dodecatheon, 138
Dodonæa viscosa, 20
Draba, 364
Dracocephalum austriacum, 238
moldavicum, 20
speciosum, 320, 325
Dracontium pertusum, 25
Drosera intermedia, 170, 265, 271, 277, 473
E.
Ebenus cretica, 26
Eccremocarpus scaber, 222, 226, 326
Echeveria, 113
Echinophora maritima, 252
Echium pyrenaicum, 20
simplex, 20
vulgare, 374
Echinocactus, 149, 178, 417
Page 654
Elegia, 115
Empetrum nigrum, 197
Encamptodon perichætialis, 174
Epacridaceæ, 120, 137
Epacris, 103, 137
impressa, 61, 379, 504
Epidendrum, 114
elongatum, 107
Stamfordianum, 401
Epimedium, 23, 226
Musschianum, 390
Epiphyllum, 328
Epipactis palustris, 325
Epilobium, 81, 137, 273
angustifolium, 20
hirsutum, 246, 250, 252, 256, 262, 281, 442
palustre, 271
tetragonum, 503
Episcia bicolor, 170
Equisetaceæ, 189, 350
Equisetum, 61, 325
fluviatile, 325
limosum, 325
Telmateia, 323, 325
Eranthis hyemalis, 23, 70
Ericaceæ, 114, 119, 137
Erica, 21, 73, 137, 296
cinerea, 504
hyemalis, 378, 504
multiflora, 372
Tetralix, 286, 310, 405, 406, 504
Eriobotrya japonica, 295
Erodium, 463
Ervum Lens, 20, 25, 66
Erucago, 136
Erucastrum Pollichii, 271
canariense, 280
Eryngium, 113, 368, 442
viviparum, 104
Erysimum, 136, 148
Barbarea, 252
cheiranthoides, 252
officinale, 252
Erythrochiton hypophyllanthus, 32, 174
Escholtzia crocea, 250
Eucomis, 103
Eugenia Jambos, 369
Euonymus japonicus, 20
Empetrum nigrum, 197
Encamptodon perichætialis, 174
Epacridaceæ, 120, 137
Epacris, 103, 137
impressa, 61, 379, 504
Epidendrum, 114
elongatum, 107
Stamfordianum, 401
Epimedium, 23, 226
Musschianum, 390
Epiphyllum, 328
Epipactis palustris, 325
Epilobium, 81, 137, 273
angustifolium, 20
hirsutum, 246, 250, 252, 256, 262, 281, 442
palustre, 271
tetragonum, 503
Episcia bicolor, 170
Equisetaceæ, 189, 350
Equisetum, 61, 325
fluviatile, 325
limosum, 325
Telmateia, 323, 325
Eranthis hyemalis, 23, 70
Ericaceæ, 114, 119, 137
Erica, 21, 73, 137, 296
cinerea, 504
hyemalis, 378, 504
multiflora, 372
Tetralix, 286, 310, 405, 406, 504
Eriobotrya japonica, 295
Erodium, 463
Ervum Lens, 20, 25, 66
Erucago, 136
Erucastrum Pollichii, 271
canariense, 280
Eryngium, 113, 368, 442
viviparum, 104
Erysimum, 136, 148
Barbarea, 252
cheiranthoides, 252
officinale, 252
Erythrochiton hypophyllanthus, 32, 174
Escholtzia crocea, 250
Eucomis, 103
Eugenia Jambos, 369
Euonymus japonicus, 20
Page 655
latifolius, 369
Euphorbiaceæ, 114, 150, 369, 395
Euphorbia, 114, 150, 365, 371, 395
Characias, 20
Cyparissias, 20, 244
Esula, 198, 307, 310
exigua, 20
geniculata, 253, 256
helioscopia, 56
Lathyris, 244
rosea, 369
palustris, 106, 299
Peplus, 162
pusilla, 244
segetalis, 281
F.
Faba, see Vicia
vulgaris, 397
Fabiana, 237
Fagus silvatica, 65, 66, 197, 318, 370, 459
Festuca, 115
nemoralis, 169
ovina, 169
Ficaria, (see Ranunculus Ficaria)
ranunculoides, 70, 295, 500
Ficoideæ, 43
Ficus Carica, 114, 204, 435
stipulata, 332
Filago, 114
germanica, 108
Filices, 21, 190, 447
Fourcroya, 115
Fragaria, xxxvi adnot., 250, 295
alpina, 271
botryformis, 47
monophylla, 396
vesca, 20, 70, 116, 195, 275, 281, 406, 503
Fraxinus excelsior, 13, 21, 66, 325, 396, 421
Ornus, 21
Fritillaria imperialis, 21, 45, 296, 462, 506
Meleagris, 506
Fuchsia, 35, 38, 44, 57, 81, 91, 127, 199, 247, 250, 288, 290, 291, 292, 294, 295, 316, 359,
360, 364, 367, 400, 443
globosa, 503
Funckia, 369
Euphorbiaceæ, 114, 150, 369, 395
Euphorbia, 114, 150, 365, 371, 395
Characias, 20
Cyparissias, 20, 244
Esula, 198, 307, 310
exigua, 20
geniculata, 253, 256
helioscopia, 56
Lathyris, 244
rosea, 369
palustris, 106, 299
Peplus, 162
pusilla, 244
segetalis, 281
F.
Faba, see Vicia
vulgaris, 397
Fabiana, 237
Fagus silvatica, 65, 66, 197, 318, 370, 459
Festuca, 115
nemoralis, 169
ovina, 169
Ficaria, (see Ranunculus Ficaria)
ranunculoides, 70, 295, 500
Ficoideæ, 43
Ficus Carica, 114, 204, 435
stipulata, 332
Filago, 114
germanica, 108
Filices, 21, 190, 447
Fourcroya, 115
Fragaria, xxxvi adnot., 250, 295
alpina, 271
botryformis, 47
monophylla, 396
vesca, 20, 70, 116, 195, 275, 281, 406, 503
Fraxinus excelsior, 13, 21, 66, 325, 396, 421
Ornus, 21
Fritillaria imperialis, 21, 45, 296, 462, 506
Meleagris, 506
Fuchsia, 35, 38, 44, 57, 81, 91, 127, 199, 247, 250, 288, 290, 291, 292, 294, 295, 316, 359,
360, 364, 367, 400, 443
globosa, 503
Funckia, 369
Page 656
G.
Page 657
Gagea, 365, 367
arvensis, 375, 508
Gaillardia, 269, 271
Galanthus, 296
nivalis, 300, 506
Galeobdolon luteum, 226, 238
Galeopsis, 429
Ladanum, 238
ochroleuca, 44
Tetrahit, 429
Galium Aparine, 205, 325
Mollugo, 321, 325
verum, 325
Gaudichaudieæ, 334
Gardenia, 296, 377
florida, 504
Fortuniana, 504
radicans, 504
Gaura biennis, 20
Genista, 295
tinctoria, 502
sibirica, 502
Scoparia, 502
Gentianaceæ, 137, 150, 505
Gentiana, 71, 73, 137, 150, 252, 296
Amarella, 166, 305, 310, 371, 505
campestris, 250, 299, 442
purpurea, 389
Geraniaceæ, 113, 137, 501
Geranium, 20, 137, 221, 246, 250, 252, 292, 295, 418
columbinum, 461
nodosum, 34, 65
pratense, 501
sylvaticum, 501
Gesneraceæ, 38, 114, 222, 505
Gesnera, 33, 44, 95, 171, 339, 357, 427
Geroltiana, 88, 89
zebrina, 355
Geum, 121, 137, 465
coccineum, 275
rivale, 122, 130, 131, 250, 252, 281, 503
Gilia capitata, 281
glomeruliflora, 253, 271
Gladiolus, 21, 296
tristis, 506
Glancium luteum, 66
Gleditschia, 30, 177, 364
arvensis, 375, 508
Gaillardia, 269, 271
Galanthus, 296
nivalis, 300, 506
Galeobdolon luteum, 226, 238
Galeopsis, 429
Ladanum, 238
ochroleuca, 44
Tetrahit, 429
Galium Aparine, 205, 325
Mollugo, 321, 325
verum, 325
Gaudichaudieæ, 334
Gardenia, 296, 377
florida, 504
Fortuniana, 504
radicans, 504
Gaura biennis, 20
Genista, 295
tinctoria, 502
sibirica, 502
Scoparia, 502
Gentianaceæ, 137, 150, 505
Gentiana, 71, 73, 137, 150, 252, 296
Amarella, 166, 305, 310, 371, 505
campestris, 250, 299, 442
purpurea, 389
Geraniaceæ, 113, 137, 501
Geranium, 20, 137, 221, 246, 250, 252, 292, 295, 418
columbinum, 461
nodosum, 34, 65
pratense, 501
sylvaticum, 501
Gesneraceæ, 38, 114, 222, 505
Gesnera, 33, 44, 95, 171, 339, 357, 427
Geroltiana, 88, 89
zebrina, 355
Geum, 121, 137, 465
coccineum, 275
rivale, 122, 130, 131, 250, 252, 281, 503
Gilia capitata, 281
glomeruliflora, 253, 271
Gladiolus, 21, 296
tristis, 506
Glancium luteum, 66
Gleditschia, 30, 177, 364
Page 658
triacanthos, 44, 48
Glochidion, 310
Gloxinia, 171, 206, 207, 222, 226, 238, 284, 291, 296, 365, 451, 506
Glyceria aquatica, 169
fluitans, 169, 278
Godetia, 295, 510
Godoya, 374
Gomphia, 281
Gongora, 35
Goodenia ovata, 21, 31
Graminaceæ, 115, 138, 278, 350, 391
Gratiola, 296
Guarea, 508
Gypsophila, 149
H.
Habenaria, 138, 238
chlorantha, 382
Halenia, 222
heterantha, 222, 226
Hedera Helix, 65
Hedypnois, 114
Helianthemum, 132, 137, 295, 404
vulgare, 501
Helianthus, 38, 44, 66
annuus, 20
tuberosus, 20
Helleborus, 23, 288
fœtidus, 442
olympicus, 284
Heliotropium peruvianum, 510
Helwingia, 174
Hemerocallis, 138, 296, 310, 507
disticha, 507
fulva, 507
Hepatica, 295, 463
triloba, 500
Heracleum, 82, 113, 137, 150, 262, 365, 368
Sphondylium, 252, 256, 339
Hermesia castaneifolia, 194
Herreria parviflora, 141, 150
Hesperis, 19, 136, 295
matronalis, 20, 252, 280, 325, 500
Heterocentron, 354
Hibiscus, 137, 293, 295, 297, 360, 510
albus, 501
Glochidion, 310
Gloxinia, 171, 206, 207, 222, 226, 238, 284, 291, 296, 365, 451, 506
Glyceria aquatica, 169
fluitans, 169, 278
Godetia, 295, 510
Godoya, 374
Gomphia, 281
Gongora, 35
Goodenia ovata, 21, 31
Graminaceæ, 115, 138, 278, 350, 391
Gratiola, 296
Guarea, 508
Gypsophila, 149
H.
Habenaria, 138, 238
chlorantha, 382
Halenia, 222
heterantha, 222, 226
Hedera Helix, 65
Hedypnois, 114
Helianthemum, 132, 137, 295, 404
vulgare, 501
Helianthus, 38, 44, 66
annuus, 20
tuberosus, 20
Helleborus, 23, 288
fœtidus, 442
olympicus, 284
Heliotropium peruvianum, 510
Helwingia, 174
Hemerocallis, 138, 296, 310, 507
disticha, 507
fulva, 507
Hepatica, 295, 463
triloba, 500
Heracleum, 82, 113, 137, 150, 262, 365, 368
Sphondylium, 252, 256, 339
Hermesia castaneifolia, 194
Herreria parviflora, 141, 150
Hesperis, 19, 136, 295
matronalis, 20, 252, 280, 325, 500
Heterocentron, 354
Hibiscus, 137, 293, 295, 297, 360, 510
albus, 501
Page 659
flavescens, 501
Rosa sinensis, 501
Syriacus, 20, 501
Hieracium, 138
aureum, 17
præaltum, 339
Pilosella, 20
umbellatum, 20
Hïppocastaneæ, 501
Hippeastrum, 296
equestre, 506
Hippuris, 196
vulgaris, 325, 406
Hodgsonia, 326, 474
Holeus mollis, 169
Honckenya peploides, 196, 406, 461
Hordeum, 115, 351
nepalense, 174, 175
trifurcatum, 174, 175
Humulus Lupulus, 193, 244, 435, 472
Hyacinthus, 138, 150, 262, 296, 360, 361, 486 adnot
comosus, 409
orientalis, 21, 45, 48, 54, 172, 189, 286, 299, 348, 507
Pouzolzii, 170
Hydrangea, 417
Hydrocera triflora, 359
Hydrocharidaceæ, 506
Hydrocharis, 296
morsus ranæ, 506
Hydrocotyle, 113
Hydrophyllaceæ, 138
Hydrophyllum, 138
Hymenocallis, 404
americana, 462, 463
Hyoscyamus, 430
Hypericum, 442
perforatum, 369, 458
Hypnum triquetrum, 49
Hypochæris, 73, 138
radicata, 250, 339, 437, 457
Hyssopus officinalis, 20, 325
I.
Iberis, 295, 364
amara, 500
umbellata, 500
Rosa sinensis, 501
Syriacus, 20, 501
Hieracium, 138
aureum, 17
præaltum, 339
Pilosella, 20
umbellatum, 20
Hïppocastaneæ, 501
Hippeastrum, 296
equestre, 506
Hippuris, 196
vulgaris, 325, 406
Hodgsonia, 326, 474
Holeus mollis, 169
Honckenya peploides, 196, 406, 461
Hordeum, 115, 351
nepalense, 174, 175
trifurcatum, 174, 175
Humulus Lupulus, 193, 244, 435, 472
Hyacinthus, 138, 150, 262, 296, 360, 361, 486 adnot
comosus, 409
orientalis, 21, 45, 48, 54, 172, 189, 286, 299, 348, 507
Pouzolzii, 170
Hydrangea, 417
Hydrocera triflora, 359
Hydrocharidaceæ, 506
Hydrocharis, 296
morsus ranæ, 506
Hydrocotyle, 113
Hydrophyllaceæ, 138
Hydrophyllum, 138
Hymenocallis, 404
americana, 462, 463
Hyoscyamus, 430
Hypericum, 442
perforatum, 369, 458
Hypnum triquetrum, 49
Hypochæris, 73, 138
radicata, 250, 339, 437, 457
Hyssopus officinalis, 20, 325
I.
Iberis, 295, 364
amara, 500
umbellata, 500
Page 660
Ilex Aquifolium, 21, 66, 447
Impatiens, 161, 231, 295, 299
Balsamina, 238, 502
Imperatoria, 82
Indigofera, 459
Inula, 20
Ionopsidium acaule, 362
Ipomœa, 296, 510
pandurata, 505
Iridaceæ, 138, 506
Irina, 64
glabra, 65
Iris, 138, 286, 296, 359, 360, 361, 365, 401, 430
Kæmpferi, 506
sibirica, 506
versicolor, 45
Isochilus, 386
Ixia carminosa, 84
Ixora, 296
grandiflora, 504
J.
Jasione, 20, 114
Jasminaceæ, 137, 504
Jasminum, 137, 296, 360, 400
grandiflorum, 288, 505
hirsutum, 505
nudiflorum, 21
officinale, 21, 505
Sambac, 505
Jatropha Pohliana, 254, 256
Juglans, 244, 400
nigra, 396
regia, 66, 193
Juncaceæ, 115, 167, 169
Juncus, 115, 317
articulatus, 431
conglomeratus, 325
uliginosus, 107
Juniperus virginiana, 194
sinensis, 217
Jussicua, 81, 180
Justicia oxyphylla, 25
K.
Impatiens, 161, 231, 295, 299
Balsamina, 238, 502
Imperatoria, 82
Indigofera, 459
Inula, 20
Ionopsidium acaule, 362
Ipomœa, 296, 510
pandurata, 505
Iridaceæ, 138, 506
Irina, 64
glabra, 65
Iris, 138, 286, 296, 359, 360, 361, 365, 401, 430
Kæmpferi, 506
sibirica, 506
versicolor, 45
Isochilus, 386
Ixia carminosa, 84
Ixora, 296
grandiflora, 504
J.
Jasione, 20, 114
Jasminaceæ, 137, 504
Jasminum, 137, 296, 360, 400
grandiflorum, 288, 505
hirsutum, 505
nudiflorum, 21
officinale, 21, 505
Sambac, 505
Jatropha Pohliana, 254, 256
Juglans, 244, 400
nigra, 396
regia, 66, 193
Juncaceæ, 115, 167, 169
Juncus, 115, 317
articulatus, 431
conglomeratus, 325
uliginosus, 107
Juniperus virginiana, 194
sinensis, 217
Jussicua, 81, 180
Justicia oxyphylla, 25
K.
Page 661
Kerria, 295
Knautia arvensis, 20, 114
Kochia Scoparia, 430
L.
Labiatæ, 138, 429
Laburnum (see Cylisus), 65, 157, 189, 222, 226
Lacistema, 359
Lactuca, 114
sativa, 11, 20, 33, 44, 313
Lambertia, 365
Lampsana, 114
Lamium, 73, 238, 361
album, 62, 63, 66, 86, 409
amplexicaule, 404
purpureum, 66, 325, 404
Larix, 114
europæa, 21, 90, 245, 435
microcarpa, 192
Lathyrus latifolius, 262
tuberosus, 30
Laurus, 296, 362
nobilis, 506
Sassafras, 250, 331, 506
Lavatera trimestris, 20
Lebeckia, 459
Lecythis, 149, 180
Leguminosæ, 48, 71, 73, 106, 113, 122, 137, 139, 146, 147, 149, 272, 276, 429, 434, 444
Leitneria floridana, 194
Leontice, 125
Leontodon, 17, 20, 44, 163, 243, 442
Lepidium, 148, 364
sativum, 57
Lepyrodia hermaphrodita, 197
Leskea sericea, 49
Leucanthemum, 86
Leucobryum giganteum, 194
Leucoium, 150, 296
æstivum, 84, 138, 350
vernum, 350, 506
Lilium, 73, 106, 115, 296, 367, 375, 421
auratum, 73, 289, 400
bulbiferum, 45
candidum, 21, 286, 325, 375, 507
cruentum, 21
lancifolium, 35, 400, 443
Knautia arvensis, 20, 114
Kochia Scoparia, 430
L.
Labiatæ, 138, 429
Laburnum (see Cylisus), 65, 157, 189, 222, 226
Lacistema, 359
Lactuca, 114
sativa, 11, 20, 33, 44, 313
Lambertia, 365
Lampsana, 114
Lamium, 73, 238, 361
album, 62, 63, 66, 86, 409
amplexicaule, 404
purpureum, 66, 325, 404
Larix, 114
europæa, 21, 90, 245, 435
microcarpa, 192
Lathyrus latifolius, 262
tuberosus, 30
Laurus, 296, 362
nobilis, 506
Sassafras, 250, 331, 506
Lavatera trimestris, 20
Lebeckia, 459
Lecythis, 149, 180
Leguminosæ, 48, 71, 73, 106, 113, 122, 137, 139, 146, 147, 149, 272, 276, 429, 434, 444
Leitneria floridana, 194
Leontice, 125
Leontodon, 17, 20, 44, 163, 243, 442
Lepidium, 148, 364
sativum, 57
Lepyrodia hermaphrodita, 197
Leskea sericea, 49
Leucanthemum, 86
Leucobryum giganteum, 194
Leucoium, 150, 296
æstivum, 84, 138, 350
vernum, 350, 506
Lilium, 73, 106, 115, 296, 367, 375, 421
auratum, 73, 289, 400
bulbiferum, 45
candidum, 21, 286, 325, 375, 507
cruentum, 21
lancifolium, 35, 400, 443
Page 662
longiflorum, 310
Martagon, 21, 286, 325, 507
tigrinum, 306, 310
Linaria, 137, 229, 230, 233, 296, 316, 361, 365, 367, 405
æruginea, 238
chalepensis, 238
Cymbalaria, 238
decumbens, 238
Elatine, 238
origanifolia, 238
Pelisseriana, 238
pilosa, 238
purpurea, 20, 44, 238
spuria, 238
triphylla, 238
vulgaris, 162, 226, 234, 235, 238, 316, 374, 505
triornithophora, 238
Linum, 335
Linum usitatissimum, 20
Liquidambar, 362
Listera ovata, 398
Lobeliaceæ, 72, 114
Lobelia, 211, 424
Lolium, 86, 113, 115, 351
perenne, 61, 169, 325
Lonicera, 38, 44, 73, 82, 226, 271, 281, 296, 297
brachypoda, 358
Caprifolium, 408
Periclymenum, 66, 226, 251, 256, 262, 263, 338, 379, 404, 406, 504
Xylosteum, 252, 358
Lopezia, 298, 410
Lotus, 113, 295, 360, 510
corniculatus, 104, 377, 436, 502
uliginosus, 363
Lowea, 396
Lunaria, 136, 364
Lupinus, 106, 165, 226, 280
polyphyllus, 238
Lycaste Skinneri, 93, 95, 291
Lychnis, 113, 137, 148, 295
chalcedonica, 501
Coronaria, 107, 510
dioica, 67, 252, 262, 280, 404, 464
flos cuculi, 501
sylvestris, 252, 501
vespertina, 501
Viscaria, 501
Lycium, 365, 367
Martagon, 21, 286, 325, 507
tigrinum, 306, 310
Linaria, 137, 229, 230, 233, 296, 316, 361, 365, 367, 405
æruginea, 238
chalepensis, 238
Cymbalaria, 238
decumbens, 238
Elatine, 238
origanifolia, 238
Pelisseriana, 238
pilosa, 238
purpurea, 20, 44, 238
spuria, 238
triphylla, 238
vulgaris, 162, 226, 234, 235, 238, 316, 374, 505
triornithophora, 238
Linum, 335
Linum usitatissimum, 20
Liquidambar, 362
Listera ovata, 398
Lobeliaceæ, 72, 114
Lobelia, 211, 424
Lolium, 86, 113, 115, 351
perenne, 61, 169, 325
Lonicera, 38, 44, 73, 82, 226, 271, 281, 296, 297
brachypoda, 358
Caprifolium, 408
Periclymenum, 66, 226, 251, 256, 262, 263, 338, 379, 404, 406, 504
Xylosteum, 252, 358
Lopezia, 298, 410
Lotus, 113, 295, 360, 510
corniculatus, 104, 377, 436, 502
uliginosus, 363
Lowea, 396
Lunaria, 136, 364
Lupinus, 106, 165, 226, 280
polyphyllus, 238
Lycaste Skinneri, 93, 95, 291
Lychnis, 113, 137, 148, 295
chalcedonica, 501
Coronaria, 107, 510
dioica, 67, 252, 262, 280, 404, 464
flos cuculi, 501
sylvestris, 252, 501
vespertina, 501
Viscaria, 501
Lycium, 365, 367
Page 663
europæum, 250
Lycopersicum, see Solanum
esculentum, 389
Lysimachia, 119
Ephemerum, 271, 281
nummularia, 505
vulgaris, 20, 87
Lythrum, 335
Salicaria, 374
M.
Mælenia, 403
Mæsa, 145, 371
Magnoliaceæ, 122
Magnolia, 288, 388, 440
Campbelli, 427
fuscata, 304, 310, 427
Malpighiaceæ, 137, 334, 403, 404, 406
Malus, 78, 79, 388, 389, see Pyrus
Malvaceæ, 137, 149, 288, 292, 295, 362, 395
Malva, 295
crispa, 448
moschata, 501
rotundifolia, 501
sylvestris, 252
Mangifera, 159, 369
Marchantia, 174
Marcgraavia, 23
umbellata, 332
Masdevallia, 27, 95
Mathiola, 136, 148, 295
annua, 361, 500
incana, 20, 38, 44, 68, 299, 500
glabrata, 500
Matricaria Parthenium, 281
Maxillaria, 383
Medicago, 137, 149, 262, 295, 364, 502
lupulina, 404, 432
maculata, 218, 271
Melastoma, 74, 125, 424
Melia Azedarach, 21
Melianthus major, 298, 407
Melilotus, 137, 139, 147, 149, 262
arvensis, 374
leucantha, 432
macrorhiza, 271
Lycopersicum, see Solanum
esculentum, 389
Lysimachia, 119
Ephemerum, 271, 281
nummularia, 505
vulgaris, 20, 87
Lythrum, 335
Salicaria, 374
M.
Mælenia, 403
Mæsa, 145, 371
Magnoliaceæ, 122
Magnolia, 288, 388, 440
Campbelli, 427
fuscata, 304, 310, 427
Malpighiaceæ, 137, 334, 403, 404, 406
Malus, 78, 79, 388, 389, see Pyrus
Malvaceæ, 137, 149, 288, 292, 295, 362, 395
Malva, 295
crispa, 448
moschata, 501
rotundifolia, 501
sylvestris, 252
Mangifera, 159, 369
Marchantia, 174
Marcgraavia, 23
umbellata, 332
Masdevallia, 27, 95
Mathiola, 136, 148, 295
annua, 361, 500
incana, 20, 38, 44, 68, 299, 500
glabrata, 500
Matricaria Parthenium, 281
Maxillaria, 383
Medicago, 137, 149, 262, 295, 364, 502
lupulina, 404, 432
maculata, 218, 271
Melastoma, 74, 125, 424
Melia Azedarach, 21
Melianthus major, 298, 407
Melilotus, 137, 139, 147, 149, 262
arvensis, 374
leucantha, 432
macrorhiza, 271
Page 664
officinalis, 404
Melittis, 361
Mentha, 238, 361, 459
aquatica, 325
viridis, 325
Mercurialis, 62, 194, 365
annua, 198
perennis, 66
Mesembryanthemum, 26
Metrosideros, 103
Miconia, 355
Mimosa, 365
Lophantha, 31
Mimulus, 73, 284, 296
luteus, 505
Mnium serratum, 49
Mirabilis, 296, 418
Jalapa, 506
Modecca, 326
Mollugo Cerviana, 398
Momordica Elaterium, 20
Monarda fistulosa, 298
Morus, 193, 354, 459
Mozinna, 399
peltata, 194
Musa, 245, 407
Muscari, 408, 461
comosum, 340, 467
Musci, 174
Mussænda, 249, 283, 429
Myosotis cæspitosa, 281
palustris, 375
scorpioides, 20
Myosurus, 440
Myristica moschata, 194
Myrtaceæ, 80, 149, 362, 395
Myrtus, 295, 310
communis, 503
N.
Narcissus, 21, 67, 138, 286, 296, 360, 365, 443
aureus, 506
biflorus, 34, 45, 506
chrysanthus, 38, 45
concolor, 506
Cypri, 506
Melittis, 361
Mentha, 238, 361, 459
aquatica, 325
viridis, 325
Mercurialis, 62, 194, 365
annua, 198
perennis, 66
Mesembryanthemum, 26
Metrosideros, 103
Miconia, 355
Mimosa, 365
Lophantha, 31
Mimulus, 73, 284, 296
luteus, 505
Mnium serratum, 49
Mirabilis, 296, 418
Jalapa, 506
Modecca, 326
Mollugo Cerviana, 398
Momordica Elaterium, 20
Monarda fistulosa, 298
Morus, 193, 354, 459
Mozinna, 399
peltata, 194
Musa, 245, 407
Muscari, 408, 461
comosum, 340, 467
Musci, 174
Mussænda, 249, 283, 429
Myosotis cæspitosa, 281
palustris, 375
scorpioides, 20
Myosurus, 440
Myristica moschata, 194
Myrtaceæ, 80, 149, 362, 395
Myrtus, 295, 310
communis, 503
N.
Narcissus, 21, 67, 138, 286, 296, 360, 365, 443
aureus, 506
biflorus, 34, 45, 506
chrysanthus, 38, 45
concolor, 506
Cypri, 506
Page 665
Jonquilla, 506
incomparabilis, 38, 45, 301, 506
italicus, 506
lobularis, 506
major, 154
montanus, 301
poculiformis, 506
poeticus, 24, 33, 301, 506
pseudo-narcissus, 506
Tazetta, 45, 300, 506
Telamonius, 506
Nasturtium, 136, 271
amphibium, 181
Nelumbium, 295
speciosum, 500
Nepeta diffusa, 238
Nepenthes, 328, 473
Nephrodium molle, 447
Nerium Oleander, 62, 66, 296, 301, 377
odorum, 505
Neuropeltis, 32
Nicandra physaloides, 458
Nicotiana, 23, 31, 73
rustica, 281
Nigella, 220, 262, 295, 297, 364, 374, 399
damascena, 226, 269, 271, 286, 375, 500
Nuytsia, 371
Nymphæaceæ, 148
Nymphæa, 50, 141, 143, 148, 285, 295
alba, 162
dentata, 256, 261
guineensis, 170
Lotus, 277, 280
O.
Octadenia, 364
Odontoglossum Alexandræ, 387, 403
Uro Skinneri, 463
Œnanthe, 82, 114, 360, 365, 457
crocata, 80, 359, 457
Œnothera, 44, 367
striata, 252
Olea europœa, 21, 157, 420
Onagraceæ, 81, 137, 406, 503
Oncidium, 67, 114, 227
abortivum, 462
incomparabilis, 38, 45, 301, 506
italicus, 506
lobularis, 506
major, 154
montanus, 301
poculiformis, 506
poeticus, 24, 33, 301, 506
pseudo-narcissus, 506
Tazetta, 45, 300, 506
Telamonius, 506
Nasturtium, 136, 271
amphibium, 181
Nelumbium, 295
speciosum, 500
Nepeta diffusa, 238
Nepenthes, 328, 473
Nephrodium molle, 447
Nerium Oleander, 62, 66, 296, 301, 377
odorum, 505
Neuropeltis, 32
Nicandra physaloides, 458
Nicotiana, 23, 31, 73
rustica, 281
Nigella, 220, 262, 295, 297, 364, 374, 399
damascena, 226, 269, 271, 286, 375, 500
Nuytsia, 371
Nymphæaceæ, 148
Nymphæa, 50, 141, 143, 148, 285, 295
alba, 162
dentata, 256, 261
guineensis, 170
Lotus, 277, 280
O.
Octadenia, 364
Odontoglossum Alexandræ, 387, 403
Uro Skinneri, 463
Œnanthe, 82, 114, 360, 365, 457
crocata, 80, 359, 457
Œnothera, 44, 367
striata, 252
Olea europœa, 21, 157, 420
Onagraceæ, 81, 137, 406, 503
Oncidium, 67, 114, 227
abortivum, 462
Page 666
bicolor, 45
Cebolleta, 107
Oncidium cucullatum, 91
heteranthum, 224
ornithorhyncum, 43
Ononis minutissima, 404
monophylla, 396
Opercularia, 38
Ophrys, 114
apifera, 360, 398
aranifera, 35, 42, 45, 238, 298, 383, 384, 385, 386, 398
insectifera, 27, 301, 382
fucifera, 506
Opuntia, 81, 113, 149, 180
fragilis, 107, 178
monacantha, 178
Salmiana, 107, 178, 179
Orchidaceæ, 27, 34, 42, 91, 112, 114, 128, 138, 150, 153, 209, 223, 290, 345, 360, 366, 367,
380-387, 397
Orchis, 138, 150, 153, 154, 227, 296
conopsea, 238
latifolia, 238
mascula, 153, 154, 238, 299, 387, 506
militaris, 387
Morio, 238, 382, 384, 387, 506
palustris, 366
papilionacea, 238
pyramidalis, 128, 238, 506
sambucina, 63, 66
simia, 238
Ornithogalum, 114, 360, 365, 367
longebracteatum, 171
Orobanchaceæ, 137
Orobanche, 73, 137, 209, 296, 505
gracilis, 442
Orobus, 295
vernus, 502
viscosus, 502
Ouvirandra, 458
Oxalidaceæ, 403
Oxalis, 295, 329, 404, 459
Acetosella, 404
carnea, 502
P.
Pæonia, 295, 300, 364, 399, 407
Cebolleta, 107
Oncidium cucullatum, 91
heteranthum, 224
ornithorhyncum, 43
Ononis minutissima, 404
monophylla, 396
Opercularia, 38
Ophrys, 114
apifera, 360, 398
aranifera, 35, 42, 45, 238, 298, 383, 384, 385, 386, 398
insectifera, 27, 301, 382
fucifera, 506
Opuntia, 81, 113, 149, 180
fragilis, 107, 178
monacantha, 178
Salmiana, 107, 178, 179
Orchidaceæ, 27, 34, 42, 91, 112, 114, 128, 138, 150, 153, 209, 223, 290, 345, 360, 366, 367,
380-387, 397
Orchis, 138, 150, 153, 154, 227, 296
conopsea, 238
latifolia, 238
mascula, 153, 154, 238, 299, 387, 506
militaris, 387
Morio, 238, 382, 384, 387, 506
palustris, 366
papilionacea, 238
pyramidalis, 128, 238, 506
sambucina, 63, 66
simia, 238
Ornithogalum, 114, 360, 365, 367
longebracteatum, 171
Orobanchaceæ, 137
Orobanche, 73, 137, 209, 296, 505
gracilis, 442
Orobus, 295
vernus, 502
viscosus, 502
Ouvirandra, 458
Oxalidaceæ, 403
Oxalis, 295, 329, 404, 459
Acetosella, 404
carnea, 502
P.
Pæonia, 295, 300, 364, 399, 407
Page 667
Moutan, 500
officinalis, 261
paradoxa, 500
tenuiflora, 500
Palmaceæ, 155
Panax, 331
Papaver, 289, 295, 297, 473
bracteatum, 28, 304, 310, 500
nudicaule, 310
orientale, 250
Papaver Rhœas, 429, 458, 500
somniferum, 28, 305, 310, 500
Papaveraceæ, 99, 500
Papilionaceæ, 397, 502, see Leguminosæ
Paris quadrifolia, 31, 358, 360, 367, 396
Paritium, 137
Parnassia, 364
Paronychia serpyllifolia, 473
Parthenium inodorum, 243
Passiflora, 137, 201, 301, 365, 463, 473
cœrulea, 185
palmata, 185
quadrangularis, 181
Passifloraceæ, 137
Pastinaca, 82
sativa, 272
Pavia, 137
Pedicularis, 238
euphrasioides, 238
sylvatica, 44, 223, 226, 238
Peganum, 363
Pelargonium, 22, 23, 25, 63, 107, 113, 208, 221, 225, 295, 336, 360, 373, 389, 418, 419, 434
grandiflorum, 86
inquinans, 30, 226
zonale, 106, 226, 501
Peltaria, 137
alliacea, 250, 251, 252
Pentstemon, 226, 455
Pereskia, 107, 113, 149
Bleo, 179
Perilla, 459
Persica (Amygdalus)
vulgaris, 44, 250, 252, 503
Petalostylis, 296
Petunia, 254, 256, 291, 296
nyctaginiflora, 505
violacea, 34, 44, 250, 253, 505
Phaius grandiflorus, 107, 509, 510
officinalis, 261
paradoxa, 500
tenuiflora, 500
Palmaceæ, 155
Panax, 331
Papaver, 289, 295, 297, 473
bracteatum, 28, 304, 310, 500
nudicaule, 310
orientale, 250
Papaver Rhœas, 429, 458, 500
somniferum, 28, 305, 310, 500
Papaveraceæ, 99, 500
Papilionaceæ, 397, 502, see Leguminosæ
Paris quadrifolia, 31, 358, 360, 367, 396
Paritium, 137
Parnassia, 364
Paronychia serpyllifolia, 473
Parthenium inodorum, 243
Passiflora, 137, 201, 301, 365, 463, 473
cœrulea, 185
palmata, 185
quadrangularis, 181
Passifloraceæ, 137
Pastinaca, 82
sativa, 272
Pavia, 137
Pedicularis, 238
euphrasioides, 238
sylvatica, 44, 223, 226, 238
Peganum, 363
Pelargonium, 22, 23, 25, 63, 107, 113, 208, 221, 225, 295, 336, 360, 373, 389, 418, 419, 434
grandiflorum, 86
inquinans, 30, 226
zonale, 106, 226, 501
Peltaria, 137
alliacea, 250, 251, 252
Pentstemon, 226, 455
Pereskia, 107, 113, 149
Bleo, 179
Perilla, 459
Persica (Amygdalus)
vulgaris, 44, 250, 252, 503
Petalostylis, 296
Petunia, 254, 256, 291, 296
nyctaginiflora, 505
violacea, 34, 44, 250, 253, 505
Phaius grandiflorus, 107, 509, 510
Page 668
Phalænopsis, 238
amabilis, 227
equestris, 231, 238
Schilleriana, 224
Pharbitis, 73
Phaseolus, 21, 298, 364
Philadelphaceæ, 113, 137, 149
Philadelphus, 113, 137, 142, 149, 295, 364, 367
coronarius, 503
speciosus, 178, 298
Philyra brasiliensis, 198
Phleum, 115, 138
phalaroides, 169
pratense, 325
Phlomis, 138
biloba, 72
fruticosa, 119, 238
Phlox, 73
Phœnix dactylifera, 318
Phylica, 325
Phyllanthus longifolius, 198
Phyllarthron, 64, 396
Phyteuma odoratum, 271
orbiculare, 20
spicatum, 166, 252
Phytolacca, 20
Picea, see Abies
Lowii, 203
nobilis, 203
Nordmanniana, 203
Webbiana, 203
Picrasma, 509
Pimpinella magna, 66
Saxifragra, 66
Pinckneya, 429
Pinus, 11, 19, 114, 318, 346, 471, (see Abies, Larix, Cedrus)
alba, 192
pinea, 90, 218
Pinaster, 13, 21
sylvestris, 21, 349
Pistacia Lentiscus, 197
Pisum, 137, 295
sativum, 27, 30, 31, 95, 166, 302, 372, 432, 472, 502, 510
Platycodon, 296
grandiflorum, 504
Plantaginaceæ, 114
Plantago, 114, 136
Coronopus, 458
amabilis, 227
equestris, 231, 238
Schilleriana, 224
Pharbitis, 73
Phaseolus, 21, 298, 364
Philadelphaceæ, 113, 137, 149
Philadelphus, 113, 137, 142, 149, 295, 364, 367
coronarius, 503
speciosus, 178, 298
Philyra brasiliensis, 198
Phleum, 115, 138
phalaroides, 169
pratense, 325
Phlomis, 138
biloba, 72
fruticosa, 119, 238
Phlox, 73
Phœnix dactylifera, 318
Phylica, 325
Phyllanthus longifolius, 198
Phyllarthron, 64, 396
Phyteuma odoratum, 271
orbiculare, 20
spicatum, 166, 252
Phytolacca, 20
Picea, see Abies
Lowii, 203
nobilis, 203
Nordmanniana, 203
Webbiana, 203
Picrasma, 509
Pimpinella magna, 66
Saxifragra, 66
Pinckneya, 429
Pinus, 11, 19, 114, 318, 346, 471, (see Abies, Larix, Cedrus)
alba, 192
pinea, 90, 218
Pinaster, 13, 21
sylvestris, 21, 349
Pistacia Lentiscus, 197
Pisum, 137, 295
sativum, 27, 30, 31, 95, 166, 302, 372, 432, 472, 502, 510
Platycodon, 296
grandiflorum, 504
Plantaginaceæ, 114
Plantago, 114, 136
Coronopus, 458
Page 669
lanceolata, 61, 104, 108, 110, 111, 243
media, 20, 111
major, 65, 108, 109, 111, 243, 372, 374
maritima, 108
Platanus, 50
Plectranthus fruticosus, 238
Plumbaginaceæ, 114
Plumbago, 360
Poa alpina, 169
annua, 169
bulbosa, 169
pratensis, 169
trivialis, 169
Podalyria myrtillifolia, 44
Podocarpus, 426
Podophyllum, 295
peltatum, 500
Podospermum laciniatum, 247, 250
Poggendorffia, 294
Pogonia ophioglossoides, 238, 386
Poinsettia, 340
Polemonium cœruleum, 20, 66, 253, 305, 310, 404
Polianthes, 296
Polygala vulgaris, 399
Polygonaceæ, 114, 138, 150
Polygonatum anceps, 42, 45
multifolium, 23, 31
Polygonum, 114, 365
orientale, 31, 390
viviparum, 106, 169
Polypodium anomalum, 190
Pomaceæ, 70, 71, 77, 79, 142, 405, 503
Pomaderris elliptica, 355
Pomax, 38
Populus, 202, 309
alba, 66
Portulaca, 295
Potamogeton, 329
bifolium, 434
Potentilla, 139, 140, 149, 295, 374, 438
alpestris, 503
argentea, 280
anserina, 503
major, 374
nepalensis, 250, 252, 275, 280
reptans, 503
Tormentilla, 503
Poterium, 113
media, 20, 111
major, 65, 108, 109, 111, 243, 372, 374
maritima, 108
Platanus, 50
Plectranthus fruticosus, 238
Plumbaginaceæ, 114
Plumbago, 360
Poa alpina, 169
annua, 169
bulbosa, 169
pratensis, 169
trivialis, 169
Podalyria myrtillifolia, 44
Podocarpus, 426
Podophyllum, 295
peltatum, 500
Podospermum laciniatum, 247, 250
Poggendorffia, 294
Pogonia ophioglossoides, 238, 386
Poinsettia, 340
Polemonium cœruleum, 20, 66, 253, 305, 310, 404
Polianthes, 296
Polygala vulgaris, 399
Polygonaceæ, 114, 138, 150
Polygonatum anceps, 42, 45
multifolium, 23, 31
Polygonum, 114, 365
orientale, 31, 390
viviparum, 106, 169
Polypodium anomalum, 190
Pomaceæ, 70, 71, 77, 79, 142, 405, 503
Pomaderris elliptica, 355
Pomax, 38
Populus, 202, 309
alba, 66
Portulaca, 295
Potamogeton, 329
bifolium, 434
Potentilla, 139, 140, 149, 295, 374, 438
alpestris, 503
argentea, 280
anserina, 503
major, 374
nepalensis, 250, 252, 275, 280
reptans, 503
Tormentilla, 503
Poterium, 113
Page 670
Sanguisorba, 107
polygamum, 281
Primula, 71, 73, 114, 121, 138, 150, 296, 335, 360, 365, 367
acaulis (see vulgaris), 45, 248, 250, 308, 310, 377, 504
Auricula, 45, 133, 271, 282, 504
denticulata, 20, 504
elatior, 45, 250, 504
calycanthema, 283
imperialis, 103
officinalis, 250, 283
prænitens, 271, 504 (see sinensis)
sinensis, 24, 45, 95, 98, 103, 133, 253, 256, 262, 263, 271, 281, 297, 314, 315, 389,
429, 441, 449
variabilis, 104
veris, 17, 20, 35, 39, 45, 105, 164
villosa, 504
vulgaris, 17, 70, 106 (see acaulis and officinalis)
Primulaceæ, 98, 114, 118, 121, 133, 138, 150, 262, 339, 366, 504
Prismatocarpus, 114, 150, 178
Prunella vulgaris, 226
Prunus, 137, 203, 295, 300, 360, 364, 367 (see Cerasus, Amygdalus,
Persica)
Armeniaca, 44
Cerasus, 74
domestica, 366, 464
Lauro Cerasus, 21, 446
spinosa, 44
Prunus sylvestris, 21
Pseudostachyum polymorphum, 168
Psittacanthus, 371
Ptelea, 364
Pterandra, 290
Pteridophyllum, 331
Pterisanthes, 328
Pteris quadriaurita, 333
Punica Granatum, 21, 295, 325
Pyrethrum, 114
inodorum, 431
Parthenium, 339
Pyrus, 82, 107, 113, 121, 137, 149, 203
communis (Pear), 36, 66, 162, 178, 422, 423, 503
dioica, 304
japonica, 161
Malus (Apple), 44, 78, 79, 166, 210, 250, 282, 295, 310, 325, 327, 375, 406, 420, 503
spectabilis, 508
torminalis, 325
polygamum, 281
Primula, 71, 73, 114, 121, 138, 150, 296, 335, 360, 365, 367
acaulis (see vulgaris), 45, 248, 250, 308, 310, 377, 504
Auricula, 45, 133, 271, 282, 504
denticulata, 20, 504
elatior, 45, 250, 504
calycanthema, 283
imperialis, 103
officinalis, 250, 283
prænitens, 271, 504 (see sinensis)
sinensis, 24, 45, 95, 98, 103, 133, 253, 256, 262, 263, 271, 281, 297, 314, 315, 389,
429, 441, 449
variabilis, 104
veris, 17, 20, 35, 39, 45, 105, 164
villosa, 504
vulgaris, 17, 70, 106 (see acaulis and officinalis)
Primulaceæ, 98, 114, 118, 121, 133, 138, 150, 262, 339, 366, 504
Prismatocarpus, 114, 150, 178
Prunella vulgaris, 226
Prunus, 137, 203, 295, 300, 360, 364, 367 (see Cerasus, Amygdalus,
Persica)
Armeniaca, 44
Cerasus, 74
domestica, 366, 464
Lauro Cerasus, 21, 446
spinosa, 44
Prunus sylvestris, 21
Pseudostachyum polymorphum, 168
Psittacanthus, 371
Ptelea, 364
Pterandra, 290
Pteridophyllum, 331
Pterisanthes, 328
Pteris quadriaurita, 333
Punica Granatum, 21, 295, 325
Pyrethrum, 114
inodorum, 431
Parthenium, 339
Pyrus, 82, 107, 113, 121, 137, 149, 203
communis (Pear), 36, 66, 162, 178, 422, 423, 503
dioica, 304
japonica, 161
Malus (Apple), 44, 78, 79, 166, 210, 250, 282, 295, 310, 325, 327, 375, 406, 420, 503
spectabilis, 508
torminalis, 325
Page 671
Q.
Page 672
Quercus, 51
Cerris, 66
pubescens, 66
R.
Raphanus sativus, 161, 252, 327, 360, 369
Ranunculaceæ, 113, 122, 136, 148, 195, 246, 339, 410, 499
Ranunculus, 19, 55, 113, 119, 258, 288, 295, 296, 297, 360, 367, 407
aconitifolius, 500
acris, 67, 250, 436, 500
aquatilis, 458
asiaticas, 500
auricomus, 195, 285, 397, 406, 461, 469
bulbosus, 17, 20, 44, 195, 406, 500
bullatus, 500
Ficaria, 70, 96, 195, 261, 368, 406
fluitans, 433, 437
gramineus, 500
Lingua, 44, 67
Philonotis, 20, 339
tripartitus, 20, 46
repens, 23, 252, 261, 500
Resedaceæ, 137, 148
Reseda, 67, 137, 141, 148
lutea, 252, 271
Luteola, 61, 112, 399
odorata, 20, 44, 280
Phyteuma, 252, 261
Restiaceæ, 115, 167, 189, 350, 463
Restio, 115, 194
Rhamnus catharticus, 463
Frangula, 280
Rhinanthus crista galli, 238
Rhodiola rosea, 197
Rhodora, 73
Rhododendron, 72, 73, 91, 133, 137, 176, 289, 290, 296, 453
indicum, 504
linearilobum, 72
Rhus Cotinus, 62, 66, 348, 409, 467, 472
Toxicodendron, 66
Ribes, 296
nigrum, 66, 296
sanguineum, 503
Richardia æthiopica, 337, 358
Ricinus communis, 198, 300
Cerris, 66
pubescens, 66
R.
Raphanus sativus, 161, 252, 327, 360, 369
Ranunculaceæ, 113, 122, 136, 148, 195, 246, 339, 410, 499
Ranunculus, 19, 55, 113, 119, 258, 288, 295, 296, 297, 360, 367, 407
aconitifolius, 500
acris, 67, 250, 436, 500
aquatilis, 458
asiaticas, 500
auricomus, 195, 285, 397, 406, 461, 469
bulbosus, 17, 20, 44, 195, 406, 500
bullatus, 500
Ficaria, 70, 96, 195, 261, 368, 406
fluitans, 433, 437
gramineus, 500
Lingua, 44, 67
Philonotis, 20, 339
tripartitus, 20, 46
repens, 23, 252, 261, 500
Resedaceæ, 137, 148
Reseda, 67, 137, 141, 148
lutea, 252, 271
Luteola, 61, 112, 399
odorata, 20, 44, 280
Phyteuma, 252, 261
Restiaceæ, 115, 167, 189, 350, 463
Restio, 115, 194
Rhamnus catharticus, 463
Frangula, 280
Rhinanthus crista galli, 238
Rhodiola rosea, 197
Rhodora, 73
Rhododendron, 72, 73, 91, 133, 137, 176, 289, 290, 296, 453
indicum, 504
linearilobum, 72
Rhus Cotinus, 62, 66, 348, 409, 467, 472
Toxicodendron, 66
Ribes, 296
nigrum, 66, 296
sanguineum, 503
Richardia æthiopica, 337, 358
Ricinus communis, 198, 300
Page 673
Ricotiana, 364
Robinia, 365
Pseudacacia, 44, 204, 317, 325, 396
Rosaceæ, 70, 71, 77, 80, 113, 121, 122, 137, 149, 246, 274, 405, 406, 502
Rosa, 21, 77, 79, 82, 113, 121, 130, 137, 139, 149, 151, 152, 162, 176, 184, 185, 186, 204,
246, 247, 250, 257, 258, 262, 271, 274, 288, 289, 295, 367, 368, 394, 473, 502
alpina, 209
arvensis, 307, 310, 404
Banksiæ, 502
berberifolia, 396
canina, 502
centifolia, 31, 502
Carolina, 502
cinnamomea, 502
damascena, 502
diversifolia, 280
Eglanteria, 502
gallica, 31, 502
indica, 502
moschata, 502
nivea, 502
pimpinellifolia, 502
rubiginosa, 502
spinosissima, 502
sulphurea, 502
Rubiaceæ, 45, 80
Rubia tinctorum, 322, 325
Rubus, 137, 252, 279, 399, 406, 429, 465
arcticus, 67
cæsius, 281, 285, 502
corylifolius, 502
fruticosus, 66, 281, 374, 380, 461, 502
Idæus, 396
rosifolius, 502
Rudbeckia, 82, 114, 127
Ruellia clandestina, 403, 404
Rumex, 45, 138, 150, 325
arifolius, 278, 281
crispus, 304, 310, 448
scutatus, 278, 281, 431
Ruscus, 328, 470
aculeatus, 60, 318, 348, 445
Russellia juncea, 20
Rutaceæ, 137, 149
Ruta, 363, 367, 371
S.
Robinia, 365
Pseudacacia, 44, 204, 317, 325, 396
Rosaceæ, 70, 71, 77, 80, 113, 121, 122, 137, 149, 246, 274, 405, 406, 502
Rosa, 21, 77, 79, 82, 113, 121, 130, 137, 139, 149, 151, 152, 162, 176, 184, 185, 186, 204,
246, 247, 250, 257, 258, 262, 271, 274, 288, 289, 295, 367, 368, 394, 473, 502
alpina, 209
arvensis, 307, 310, 404
Banksiæ, 502
berberifolia, 396
canina, 502
centifolia, 31, 502
Carolina, 502
cinnamomea, 502
damascena, 502
diversifolia, 280
Eglanteria, 502
gallica, 31, 502
indica, 502
moschata, 502
nivea, 502
pimpinellifolia, 502
rubiginosa, 502
spinosissima, 502
sulphurea, 502
Rubiaceæ, 45, 80
Rubia tinctorum, 322, 325
Rubus, 137, 252, 279, 399, 406, 429, 465
arcticus, 67
cæsius, 281, 285, 502
corylifolius, 502
fruticosus, 66, 281, 374, 380, 461, 502
Idæus, 396
rosifolius, 502
Rudbeckia, 82, 114, 127
Ruellia clandestina, 403, 404
Rumex, 45, 138, 150, 325
arifolius, 278, 281
crispus, 304, 310, 448
scutatus, 278, 281, 431
Ruscus, 328, 470
aculeatus, 60, 318, 348, 445
Russellia juncea, 20
Rutaceæ, 137, 149
Ruta, 363, 367, 371
S.
Page 674
Sagina, 397
procumbens, 501
Sagittaria, 296, 329
latifolia, 507
sagittifolia, 325, 507
Salisburia adiantifolia, 61
Salix, 166, 168, 244, 310, 326, 349
babylonica, 202, 257, 262, 299, 309, 326
calyculata, 29
capræa, 271, 299
cinerea, 41, 45, 189, 299
fragilis, 357
monandra, 29
nigricans, 299
pendula, 357
repens, 194
silesiaca, 299
vitellina, 21
Salpiglossis straminea, 44
Salvia, 226
officinalis, 66
pratensis, 36
Verbenaca, 87
Sambucus, 296, 359, 360, 365, 367
nigra, 21, 55, 66, 164, 325, 396
racemosa, 66
Sanguinaria, 295
canadensis, 500
Sanguisorba, 113
officinalis, 107
Santalaceæ, 122, 138, 150
Sapindaceæ, 137, 448
Saponaria, 28, 67, 76, 149, 295, 297, 300, 360, 379
officinalis, 97, 301, 501
Sarothamnus, 360, 363
Scoparius, 363
Saxifraga, 33, 44, 106, 113, 296, 390
crassifolia, 307, 310
decipiens, 288
foliosa, 281
granulata, 298, 504
irrigua, 20
longifolia, 404
Saxifraga mutata, 20
Saxifragaceæ, 80, 81
Scabiosa, 35, 106, 114, 138, 141, 296, 297, 429
agrestis, 252
arvensis, 325
procumbens, 501
Sagittaria, 296, 329
latifolia, 507
sagittifolia, 325, 507
Salisburia adiantifolia, 61
Salix, 166, 168, 244, 310, 326, 349
babylonica, 202, 257, 262, 299, 309, 326
calyculata, 29
capræa, 271, 299
cinerea, 41, 45, 189, 299
fragilis, 357
monandra, 29
nigricans, 299
pendula, 357
repens, 194
silesiaca, 299
vitellina, 21
Salpiglossis straminea, 44
Salvia, 226
officinalis, 66
pratensis, 36
Verbenaca, 87
Sambucus, 296, 359, 360, 365, 367
nigra, 21, 55, 66, 164, 325, 396
racemosa, 66
Sanguinaria, 295
canadensis, 500
Sanguisorba, 113
officinalis, 107
Santalaceæ, 122, 138, 150
Sapindaceæ, 137, 448
Saponaria, 28, 67, 76, 149, 295, 297, 300, 360, 379
officinalis, 97, 301, 501
Sarothamnus, 360, 363
Scoparius, 363
Saxifraga, 33, 44, 106, 113, 296, 390
crassifolia, 307, 310
decipiens, 288
foliosa, 281
granulata, 298, 504
irrigua, 20
longifolia, 404
Saxifraga mutata, 20
Saxifragaceæ, 80, 81
Scabiosa, 35, 106, 114, 138, 141, 296, 297, 429
agrestis, 252
arvensis, 325
Page 675
atropurpurea, 62, 66
Columbaria, 252, 281
Scandix, 125
Schismatopera distichophylla, 194
Schœnodorus, 364 adnot
Schœnus cephalotes, 245
Schœnodon, 365
Sciadopitys verticillata, 352
Scilla, 296
autumnalis, 507
nutans, 507
Scirpus lacustris, 325
Scolopendrium D'Urvillei, 332
vulgare, 64, 314, 326, 328, 459, 467
Scorzonera, 19
octangularis, 247, 250
Scrophularia, 114, 455, 463
aquatica, 226, 238, 262, 281
arguta, 334
nodosa, 44, 106, 281
Scrophulariaceæ, 114, 121, 137, 150, 222, 505
Secale cereale, 55, 115
Sechium edule, 81, 202
Sedum cristatum, 18, 20
reflexum, 20
Selenipedium caudatum, 224, 227
Selinum, 82, 114
caruifolium, 281
Sempervivum, 105, 113, 204, 509
montanum, 310
sediforme, 164
tectorum, 292 adnot, 308, 310
Senecio, 44, 114
vulgaris, 247, 250, 252, 339, 432, 437, 439, 457
Serissa, 296, 377, 504
Sesamum, 365
indicum, 238
Seseli, 113, 281
coloratum, 166
Sideritis, 238, 365
Silaus, 113, 365
Silene, 44, 137, 148, 295, 301
conica, 68
Otites, 464
inflata, 501
Sinapis, 57, 136, 270, 295, 371
arvensis, 181, 250, 261, 264, 271, 280, 501
Sinningia purpurea, 431
Columbaria, 252, 281
Scandix, 125
Schismatopera distichophylla, 194
Schœnodorus, 364 adnot
Schœnus cephalotes, 245
Schœnodon, 365
Sciadopitys verticillata, 352
Scilla, 296
autumnalis, 507
nutans, 507
Scirpus lacustris, 325
Scolopendrium D'Urvillei, 332
vulgare, 64, 314, 326, 328, 459, 467
Scorzonera, 19
octangularis, 247, 250
Scrophularia, 114, 455, 463
aquatica, 226, 238, 262, 281
arguta, 334
nodosa, 44, 106, 281
Scrophulariaceæ, 114, 121, 137, 150, 222, 505
Secale cereale, 55, 115
Sechium edule, 81, 202
Sedum cristatum, 18, 20
reflexum, 20
Selenipedium caudatum, 224, 227
Selinum, 82, 114
caruifolium, 281
Sempervivum, 105, 113, 204, 509
montanum, 310
sediforme, 164
tectorum, 292 adnot, 308, 310
Senecio, 44, 114
vulgaris, 247, 250, 252, 339, 432, 437, 439, 457
Serissa, 296, 377, 504
Sesamum, 365
indicum, 238
Seseli, 113, 281
coloratum, 166
Sideritis, 238, 365
Silaus, 113, 365
Silene, 44, 137, 148, 295, 301
conica, 68
Otites, 464
inflata, 501
Sinapis, 57, 136, 270, 295, 371
arvensis, 181, 250, 261, 264, 271, 280, 501
Sinningia purpurea, 431
Page 676
Sisymbrium, 136
officinale, 250, 261, 271, 280
Sisymbrium tenuifolium, 252
Slateria, 74
Solanaceæ, 150, 430, 505
Solanum, 73, 125, 150, 296, 360, 365, 367, 370, 424, 453
amazonicum, 430
Dulcamara, 66, 288, 430, 504
esculentum, 74
cornutum, 430
Lycopersicum, 38, 44, 74, 171, 391, 442
tridynamum, 430
tuberosum, 54, 142, 288, 333, 420
Vespertilio, 430
Sophora, 55
Spartianthus, 295
junceus, 502
Spartium, 295
junceum, 18, 21
Scoparium, 21
Spathiphyllum, 245
Spilanthes, 138, 365
oleracea, 44
Spinacia oleracea, 31, 197
Spiræa, 21, 137, 295
Filipendula, 503
oblongifolia, 252
prunifolia, 503
Reevesii, 503
strobilacea, 503
Ulmaria, 503
Spiranthes, 319
Splachnum vasculosum, 49
Stachys, 138, 339, adnot
germanica, 310
lanata, 375
sylvatica, 74, 253, 271, 281
Stackhousia juncea, 334
Stangeria paradoxa, 172
Stapelia, 20
Staphylea pinnata, 30
Stauntonia latifolia, 194
Stellaria, 148, 404, 406
media, 271, 279, 280
Stellatæ, 396, 408
Sterculiaceæ, 362, 464
Sterculia platanifolia, 20, 256
Sternbergia, 296
officinale, 250, 261, 271, 280
Sisymbrium tenuifolium, 252
Slateria, 74
Solanaceæ, 150, 430, 505
Solanum, 73, 125, 150, 296, 360, 365, 367, 370, 424, 453
amazonicum, 430
Dulcamara, 66, 288, 430, 504
esculentum, 74
cornutum, 430
Lycopersicum, 38, 44, 74, 171, 391, 442
tridynamum, 430
tuberosum, 54, 142, 288, 333, 420
Vespertilio, 430
Sophora, 55
Spartianthus, 295
junceus, 502
Spartium, 295
junceum, 18, 21
Scoparium, 21
Spathiphyllum, 245
Spilanthes, 138, 365
oleracea, 44
Spinacia oleracea, 31, 197
Spiræa, 21, 137, 295
Filipendula, 503
oblongifolia, 252
prunifolia, 503
Reevesii, 503
strobilacea, 503
Ulmaria, 503
Spiranthes, 319
Splachnum vasculosum, 49
Stachys, 138, 339, adnot
germanica, 310
lanata, 375
sylvatica, 74, 253, 271, 281
Stackhousia juncea, 334
Stangeria paradoxa, 172
Stapelia, 20
Staphylea pinnata, 30
Stauntonia latifolia, 194
Stellaria, 148, 404, 406
media, 271, 279, 280
Stellatæ, 396, 408
Sterculiaceæ, 362, 464
Sterculia platanifolia, 20, 256
Sternbergia, 296
Page 677
lutea, 506
Stratiotes aloides, 406
Strelitzia juncea, 459
regina, 25
Streptocarpus Rexii, 15, 44, 226, 227, 461
Strophanthus, 326
Suaeda, 365
fruticosa, 430
maritima, 20, 21
Suregada, 198
Symphoricarpus racemosus, 66
Symphyomyrtus, 38
Symphytum, 71, 138, 365
officinale, 253, 262, 263, 271
Zeyheri, 271
Syringa, 296, 360, 367
persica, 44, 61, 66, 284, 505
vulgaris, 79, 505
T.
Tabernæmontana, 296
coronaria, 504
Tacsonia pinnatistipula, 294
Tamus communis, 21
Taraxacum, 164 (see Leontodon)
Taxodium, 114
distichum, 444
Taxus baccata, 11, 21, 90
Tetragonia, 113
expansa, 142, 180
Tetragoniaceæ, 149
Teucrium campanulatum, 226, 233, 238
Thalictrum, 205, 500
minus, 307, 310
Thea, 295, 502
Thelymitra, 224
Thesium, 121, 123, 138, 150
Thlaspi arvense, 250, 271
bursa-pastoris, 252, 428
Thuja occidentalis, 319
orientalis, 21
Thymus Serpyllum, 325, 405, 406
Thysselinum, 137, 365
palustre, 437, 457
Tigridia, 360, 361, 365
Pavonia, 35, 462
Stratiotes aloides, 406
Strelitzia juncea, 459
regina, 25
Streptocarpus Rexii, 15, 44, 226, 227, 461
Strophanthus, 326
Suaeda, 365
fruticosa, 430
maritima, 20, 21
Suregada, 198
Symphoricarpus racemosus, 66
Symphyomyrtus, 38
Symphytum, 71, 138, 365
officinale, 253, 262, 263, 271
Zeyheri, 271
Syringa, 296, 360, 367
persica, 44, 61, 66, 284, 505
vulgaris, 79, 505
T.
Tabernæmontana, 296
coronaria, 504
Tacsonia pinnatistipula, 294
Tamus communis, 21
Taraxacum, 164 (see Leontodon)
Taxodium, 114
distichum, 444
Taxus baccata, 11, 21, 90
Tetragonia, 113
expansa, 142, 180
Tetragoniaceæ, 149
Teucrium campanulatum, 226, 233, 238
Thalictrum, 205, 500
minus, 307, 310
Thea, 295, 502
Thelymitra, 224
Thesium, 121, 123, 138, 150
Thlaspi arvense, 250, 271
bursa-pastoris, 252, 428
Thuja occidentalis, 319
orientalis, 21
Thymus Serpyllum, 325, 405, 406
Thysselinum, 137, 365
palustre, 437, 457
Tigridia, 360, 361, 365
Pavonia, 35, 462
Page 678
Tilia asplenifolia, 66, 459
europæa, 22, 30, 65
parvifolia, 66
Tiliaceæ, 137, 362
Tithonia, 26
Tofieldia, 296
calyculata, 300
Torenia scabra, 406
Torilis, 82, 149
Anthriscus, 256, 271, 281, 339, 406, 441, 442
Tradescantia, 245, 296, 360, 365, 367, 404
virginica, 88, 507
Tragopogon, 19, 20, 73, 114, 431
orientale, 250
pratense, 247, 250, 442, 457
Trapa natans, 202, 364
Trichostomum rigidulum, 49
Trifolium, 106, 113, 121, 137, 139, 149, 295, 360, 364
Trifolium hybridum, 262, 399, 406, 407
pratense, 20
repens, 20, 23, 68, 70, 98, 145, 146, 226, 231, 238, 250, 252, 256, 260, 262, 265, 271,
276, 279, 356, 368, 397, 399, 406, 407, 434, 436, 438, 502
resupinatum, 20
Trillium grandiflorum, 507, 510
Trinia vulgaris, 405
Triphasia aurantiaca, 369
Triticum, 86, 115, 350, 391
repens, 106, 325
vulgare, 55, 113
Triumfetta, 137, 252, 259, 260, 262, 265, 271, 280
Trollius europæus, 66, 295, 500
Tropæolaceæ, 149, 501
Tropæolum, 149, 398, 283
majus, 20, 222, 225, 226, 232, 238, 251, 252, 254, 256, 271, 280, 295, 310, 406, 442,
501
minus, 501
Tulipa, 35, 45, 67, 75, 84, 85, 138, 262, 300, 302, 348, 359, 360, 361, 365, 367, 388, 390, 421
Gesneriana, 31, 250, 310, 315, 442, 507
sylvestris, 507
Turritis, 271
glabra, 252
Typha, 189
U.
Ulex, 295, 360
europæus, 377, 502
europæa, 22, 30, 65
parvifolia, 66
Tiliaceæ, 137, 362
Tithonia, 26
Tofieldia, 296
calyculata, 300
Torenia scabra, 406
Torilis, 82, 149
Anthriscus, 256, 271, 281, 339, 406, 441, 442
Tradescantia, 245, 296, 360, 365, 367, 404
virginica, 88, 507
Tragopogon, 19, 20, 73, 114, 431
orientale, 250
pratense, 247, 250, 442, 457
Trapa natans, 202, 364
Trichostomum rigidulum, 49
Trifolium, 106, 113, 121, 137, 139, 149, 295, 360, 364
Trifolium hybridum, 262, 399, 406, 407
pratense, 20
repens, 20, 23, 68, 70, 98, 145, 146, 226, 231, 238, 250, 252, 256, 260, 262, 265, 271,
276, 279, 356, 368, 397, 399, 406, 407, 434, 436, 438, 502
resupinatum, 20
Trillium grandiflorum, 507, 510
Trinia vulgaris, 405
Triphasia aurantiaca, 369
Triticum, 86, 115, 350, 391
repens, 106, 325
vulgare, 55, 113
Triumfetta, 137, 252, 259, 260, 262, 265, 271, 280
Trollius europæus, 66, 295, 500
Tropæolaceæ, 149, 501
Tropæolum, 149, 398, 283
majus, 20, 222, 225, 226, 232, 238, 251, 252, 254, 256, 271, 280, 295, 310, 406, 442,
501
minus, 501
Tulipa, 35, 45, 67, 75, 84, 85, 138, 262, 300, 302, 348, 359, 360, 361, 365, 367, 388, 390, 421
Gesneriana, 31, 250, 310, 315, 442, 507
sylvestris, 507
Turritis, 271
glabra, 252
Typha, 189
U.
Ulex, 295, 360
europæus, 377, 502
Page 679
Ulmus americana, 66
campestris, 31, 52, 62, 157, 325, 353, 427
Umbelliferæ, 37, 71, 73, 80, 107, 113, 121, 127, 131, 132, 137, 139, 140, 143, 149, 150, 162,
244, 257, 261, 339, 358, 395, 405, 406, 407, 437, 439
Uredo candida, 279
Urticaceæ, 114
Urtica dioica, 62, 66, 194
Usteria, 283
V.
Vaccinium, 68
Valantia cruciata, 44
Valerianaceæ, 114
Valeriana, 114, 165, 419
dioica, 325
officinalis, 325
Valisneria spiralis, 319, 433
Verbascum, 73, 116, 137, 225, 253, 430
australe, 35
nigrum, 226, 251
phlomoides, 281
Thapsus, 404
Verbena, 67, 68, 506
Veronica, 60, 150, 296, 334, 360, 361, 365, 375, 505
austriaca, 66
Beccabunga, 468
Chamædrys, 442
latifolia, 459
longifolia, 325
spicata, 325, 374
Viburnum, 44, 296, 417
Opulus, 473
Vicia, 30, 365, 369, 472
Faba, 434
Vinca, 137, 296
herbacea, 389
major, 505
minor, 44, 99, 358, 390, 505
rosea, 31
Viola, 23, 137, 225, 229, 289, 295, 297, 405
grandiflora, 501
hirta, 226, 238
odorata, 20, 94, 220, 226, 238, 286, 404, 429, 501, 508
tricolor, 461, 501
Violaceæ, 137, 334, 403, 406, 500
Viscum album, 56, 369, 509
campestris, 31, 52, 62, 157, 325, 353, 427
Umbelliferæ, 37, 71, 73, 80, 107, 113, 121, 127, 131, 132, 137, 139, 140, 143, 149, 150, 162,
244, 257, 261, 339, 358, 395, 405, 406, 407, 437, 439
Uredo candida, 279
Urticaceæ, 114
Urtica dioica, 62, 66, 194
Usteria, 283
V.
Vaccinium, 68
Valantia cruciata, 44
Valerianaceæ, 114
Valeriana, 114, 165, 419
dioica, 325
officinalis, 325
Valisneria spiralis, 319, 433
Verbascum, 73, 116, 137, 225, 253, 430
australe, 35
nigrum, 226, 251
phlomoides, 281
Thapsus, 404
Verbena, 67, 68, 506
Veronica, 60, 150, 296, 334, 360, 361, 365, 375, 505
austriaca, 66
Beccabunga, 468
Chamædrys, 442
latifolia, 459
longifolia, 325
spicata, 325, 374
Viburnum, 44, 296, 417
Opulus, 473
Vicia, 30, 365, 369, 472
Faba, 434
Vinca, 137, 296
herbacea, 389
major, 505
minor, 44, 99, 358, 390, 505
rosea, 31
Viola, 23, 137, 225, 229, 289, 295, 297, 405
grandiflora, 501
hirta, 226, 238
odorata, 20, 94, 220, 226, 238, 286, 404, 429, 501, 508
tricolor, 461, 501
Violaceæ, 137, 334, 403, 406, 500
Viscum album, 56, 369, 509
Page 680
Vitaceæ, 137
Vitex incisa, 238
Vitis, 417
vinifera, 20, 29, 34, 44, 66, 137, 157, 182, 183, 211, 280, 374, 422, 424
W.
Wedelia perfoliata, 442
Weigela rosea, 358
Welwitschia, 162, adnot
Wildenovia, 115, 167, 168
Wistaria, 364
sinensis, 226
X.
Xanthosoma appendiculatum, 31
Xanthoxylum, 21
Xylophylla, 328
Y.
Yucca, 361
flexilis, 361
Z.
Zamia, 170
Zea Mays, 21, 113, 136, 191, 197, 300, 310, 350, 369, 466
Zingiber Zerumbet, 224, 227
Zinnia, 44
elegans, 20
Zygopetalum maxillare, 398
Vitex incisa, 238
Vitis, 417
vinifera, 20, 29, 34, 44, 66, 137, 157, 182, 183, 211, 280, 374, 422, 424
W.
Wedelia perfoliata, 442
Weigela rosea, 358
Welwitschia, 162, adnot
Wildenovia, 115, 167, 168
Wistaria, 364
sinensis, 226
X.
Xanthosoma appendiculatum, 31
Xanthoxylum, 21
Xylophylla, 328
Y.
Yucca, 361
flexilis, 361
Z.
Zamia, 170
Zea Mays, 21, 113, 136, 191, 197, 300, 310, 350, 369, 466
Zingiber Zerumbet, 224, 227
Zinnia, 44
elegans, 20
Zygopetalum maxillare, 398
Page 681
ERRATA.
The reader is requested to make the following corrections:—
Page 182. Fig. 94 should be 94*.
Page 194. The reference 3 applies not to the nutmeg but to the hop, figured
at p. 193.
Page 309. Fig. 165 legend—for Sempervivun tecotorum read Sempervivum
tectorum.
PRINTED BY J. E. ADLARD, BARTHOLOMEW CLOSE.
Transcriber's Notes:
Page xx and 202:
208. Passage of pinnate to palmate leaves in horse-chesnut 439
The reader is requested to make the following corrections:—
Page 182. Fig. 94 should be 94*.
Page 194. The reference 3 applies not to the nutmeg but to the hop, figured
at p. 193.
Page 309. Fig. 165 legend—for Sempervivun tecotorum read Sempervivum
tectorum.
PRINTED BY J. E. ADLARD, BARTHOLOMEW CLOSE.
Transcriber's Notes:
Page xx and 202:
208. Passage of pinnate to palmate leaves in horse-chesnut 439
Page 682
'chesnut may be old spelling for chestnut?'. Changed as most are spelled
chestnut.
Page 65:
Naturforscherversammlung
Changed Naturvorschefversamlung to Naturforscherversammlung to match
other occurrence. (See footnote 528).
Page 145:
So far as the andræcium is concerned, the stamens
Changed to andrœcium to match other occurrences.
Page 149:
Echinocactus changed from Echinocatus to match other occurrences.
Page 397:
The species mentioned are Ambrina ambrosiodes,
Changed to ambrosioides to match index page.
Page 502:
Medicago sp., ? ., Europe.
The ? mark replaces a blank in original for a missing attribution.
Page 503:
Onagrariæ changed to Onagraceæ to match other occurrences, especially
the index referrence.
Footnote 126: 'Neue Denkschriften der allgemeine Schweizerischen
Gesellschaft,'
Perhaps this should be: allgemeinen. Unchanged.
Index:
Alströmeria, 319, 326
chestnut.
Page 65:
Naturforscherversammlung
Changed Naturvorschefversamlung to Naturforscherversammlung to match
other occurrence. (See footnote 528).
Page 145:
So far as the andræcium is concerned, the stamens
Changed to andrœcium to match other occurrences.
Page 149:
Echinocactus changed from Echinocatus to match other occurrences.
Page 397:
The species mentioned are Ambrina ambrosiodes,
Changed to ambrosioides to match index page.
Page 502:
Medicago sp., ? ., Europe.
The ? mark replaces a blank in original for a missing attribution.
Page 503:
Onagrariæ changed to Onagraceæ to match other occurrences, especially
the index referrence.
Footnote 126: 'Neue Denkschriften der allgemeine Schweizerischen
Gesellschaft,'
Perhaps this should be: allgemeinen. Unchanged.
Index:
Alströmeria, 319, 326
Page 683
Changed to Alstrœmeria to match referenced pages.
Dipsacaceæ, 86, 107, 114, 138
All dipsaceæ changed to dipsacaceæ to match index and current spelling.
Errata changes listed were made.
Inconsistent hyphenation:
co-existent and coexistent
Lauro-cerasus and Laurocerasus
mid-rib and midrib
outgrowth and out-growth
Dipsacaceæ, 86, 107, 114, 138
All dipsaceæ changed to dipsacaceæ to match index and current spelling.
Errata changes listed were made.
Inconsistent hyphenation:
co-existent and coexistent
Lauro-cerasus and Laurocerasus
mid-rib and midrib
outgrowth and out-growth
Page 684
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