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Title: Evolution in Modern Thought

Contributor: J. Arthur Thomson
William Bateson
Célestin Charles Alfred Bouglé
J. B. Bury
Ernst Haeckel
Harald Høffding
C. Lloyd Morgan
Gustav Albert Schwalbe
P. N. Waggett
August Weismann

Release date: August 29, 2007 [eBook #22430]

Language: English

Other information and formats: www.gutenberg.org/ebooks/22430

Credits: Produced by Marilynda Fraser-Cunliffe, Sankar Viswanathan,
and the Online Distributed Proofreading Team at
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*** START OF THE PROJECT GUTENBERG EBOOK EVOLUTION IN
MODERN THOUGHT ***

EVOLUTION IN MODERN
THOUGHT

BY HAECKEL, THOMSON, WEISMANN
AND OTHERS

THE MODERN LIBRARY

PUBLISHERS :: :: NEW YORK

CONTENTS

PAGE
I Darwin's Predecessors 1
J. Arthur Thomson, Professor of Natural
History in the University of Aberdeen
II The Selection Theory 23
August Weismann, Professor of Zoology in
the University of Freiburg (Baden)
III Heredity and Variation in Modern Lights 87
W. Bateson, Professor of Biology in the
University of Cambridge
IV "The Descent of Man" 111
G. Schwalbe, Professor of Anatomy in the
University of Strassburg
V Charles Darwin as an Anthropologist 146
Ernst Haeckel, Professor of Zoology in the
University of Jena
VI Mental Factors in Evolution 166
C. Lloyd Morgan, Professor of Psychology
at University College, Bristol
VII The Influence of the Conception of
Evolution on Modern Philosophy 197
H. Höffding, Professor of Philosophy in the
University of Copenhagen
VIII The Influence of Darwin Upon Religious
Thought 223
Rev. P. H. Waggett
IX Darwinism and History 246

J. B. Bury, Regious Professor of Modern
History in the University of Cambridge
X Darwinism and Sociology 264
C. Bouglé, Professor of Social Philosophy
in the University of Toulouse, and Deputy-
Professor at the Sorbonne, Paris

EVOLUTION IN MODERN THOUGHT

I

DARWIN'S PREDECESSORS
By J. Arthur Thomson

Professor of Natural History in the University of Aberdeen

In seeking to discover Darwin's relation to his predecessors it is useful to
distinguish the various services which he rendered to the theory of organic
evolution.
(I) As everyone knows, the general idea of the Doctrine of Descent is that
the plants and animals of the present day are the lineal descendants of
ancestors on the whole somewhat simpler, that these again are descended
from yet simpler forms, and so on backwards towards the literal "Protozoa"
and "Protophyta" about which we unfortunately know nothing. Now no one
supposes that Darwin originated this idea, which in rudiment at least is as
old as Aristotle. What Darwin did was to make it current intellectual coin.
He gave it a form that commended itself to the scientific and public
intelligence of the day, and he won widespread conviction by showing with
consummate skill that it was an effective formula to work with, a key which
no lock refused. In a scholarly, critical, and pre-eminently fair-minded way,
admitting difficulties and removing them, foreseeing objections and
forestalling them, he showed that the doctrine of descent supplied a modal
interpretation of how our present-day fauna and flora have come to be.
(II) In the second place, Darwin applied the evolution-idea to particular
problems, such as the descent of man, and showed what a powerful organon
it is, introducing order into masses of uncorrelated facts, interpreting
enigmas both of structure and function, both bodily and mental, and, best of
all, stimulating and guiding further investigation. But here again it cannot
be claimed that Darwin was original. The problem of the descent or ascent
of man, and other particular cases of evolution, had attracted not a few
naturalists before Darwin's day, though no one [except Herbert Spencer in
the psychological domain (1855)] had come near him in precision and
thoroughness of inquiry.

(III) In the third place, Darwin contributed largely to a knowledge of the
factors in the evolution-process, especially by his analysis of what occurs in
the case of domestic animals and cultivated plants, and by his elaboration of
the theory of Natural Selection which Alfred Russel Wallace independently
stated at the same time, and of which there had been a few previous
suggestions of a more or less vague description. It was here that Darwin's
originality was greatest, for he revealed to naturalists the many different
forms—often very subtle—which natural selection takes, and with the
insight of a disciplined scientific imagination he realised what a mighty
engine of progress it has been and is.
(IV) As an epoch-marking contribution, not only to Ætiology but to Natural
History in the widest sense, we rank the picture which Darwin gave to the
world of the web of life, that is to say, of the inter-relations and linkages in
Nature. For the Biology of the individual—if that be not a contradiction in
terms—no idea is more fundamental than that of the correlation of organs,
but Darwin's most characteristic contribution was not less fundamental,—it
was the idea of the correlation of organisms. This, again, was not novel; we
find it in the works of naturalists like Christian Conrad Sprengel, Gilbert
White, and Alexander von Humboldt, but the realisation of its full import
was distinctly Darwinian.
As Regards the General Idea of Organic Evolution
While it is true, as Prof. H. F. Osborn puts it, that "'Before and after Darwin'
will always be the ante et post urbem conditam of biological history," it is
also true that the general idea of organic evolution is very ancient. In his
admirable sketch From the Greeks to Darwin,[1] Prof. Osborn has shown
that several of the ancient philosophers looked upon Nature as a gradual
development and as still in process of change. In the suggestions of
Empedocles, to take the best instance, there were "four sparks of truth,—
first, that the development of life was a gradual process; second, that plants
were evolved before animals; third, that imperfect forms were gradually
replaced (not succeeded) by perfect forms; fourth, that the natural cause of
the production of perfect forms was the extinction of the imperfect."[2] But
the fundamental idea of one stage giving origin to another was absent. As
the blue Ægean teemed with treasures of beauty and threw many upon its
shores, so did Nature produce like a fertile artist what had to be rejected as

well as what was able to survive, but the idea of one species emerging out
of another was not yet conceived.
Aristotle's views of Nature[3] seem to have been more definitely
evolutionist than those of his predecessors, in this sense, at least, that he
recognised not only an ascending scale, but a genetic series from polyp to
man and an age-long movement towards perfection. "It is due to the
resistance of matter to form that Nature can only rise by degrees from lower
to higher types." "Nature produces those things which, being continually
moved by a certain principle contained in themselves, arrive at a certain
end."
To discern the outcrop of evolution-doctrine in the long interval between
Aristotle and Bacon seems to be very difficult, and some of the instances
that have been cited strike one as forced. Epicurus and Lucretius, often
called poets of evolution, both pictured animals as arising directly out of the
earth, very much as Milton's lion long afterwards pawed its way out. Even
when we come to Bruno who wrote that "to the sound of the harp of the
Universal Apollo (the World Spirit), the lower organisms are called by
stages to higher, and the lower stages are connected by intermediate forms
with the higher," there is great room, as Prof. Osborn points out,[4] for
difference of opinion as to how far he was an evolutionist in our sense of
the term.
The awakening of natural science in the sixteenth century brought the
possibility of a concrete evolution theory nearer, and in the early
seventeenth century we find evidences of a new spirit—in the embryology
of Harvey and the classifications of Ray. Besides sober naturalists there
were speculative dreamers in the sixteenth and seventeenth centuries who
had at least got beyond static formulae, but, as Professor Osborn points out,
[5] "it is a very striking fact, that the basis of our modern methods of
studying the Evolution problem was established not by the early naturalists
nor by the speculative writers, but by the Philosophers." He refers to Bacon,
Descartes, Leibnitz, Hume, Kant, Lessing, Herder, and Schelling. "They
alone were upon the main track of modern thought. It is evident that they
were groping in the dark for a working theory of the Evolution of life, and it
is remarkable that they clearly perceived from the outset that the point to
which observation should be directed was not the past but the present

mutability of species, and further, that this mutability was simply the
variation of individuals on an extended scale."
Bacon seems to have been one of the first to think definitely about the
mutability of species, and he was far ahead of his age in his suggestion of
what we now call a Station of Experimental Evolution. Leibnitz discusses
in so many words how the species of animals may be changed and how
intermediate species may once have linked those that now seem
discontinuous. "All natural orders of beings present but a single chain"....
"All advances by degrees in Nature, and nothing by leaps." Similar
evolutionist statements are to be found in the works of the other
"philosophers," to whom Prof. Osborn refers, who were, indeed, more
scientific than the naturalists of their day. It must be borne in mind that the
general idea of organic evolution—that the present is the child of the past—
is in great part just the idea of human history projected upon the natural
world, differentiated by the qualification that the continuous "Becoming"
has been wrought out by forces inherent in the organisms themselves and in
their environment.
A reference to Kant[6] should come in historical order after Buffon, with
whose writings he was acquainted, but he seems, along with Herder and
Schelling, to be best regarded as the culmination of the evolutionist
philosophers—of those at least who interested themselves in scientific
problems. In a famous passage he speaks of "the agreement of so many
kinds of animals in a certain common plan of structure" ... an "analogy of
forms" which "strengthens the supposition that they have an actual blood-
relationship, due to derivation from a common parent." He speaks of "the
great Family of creatures, for as a Family we must conceive it, if the above-
mentioned continuous and connected relationship has a real foundation."
Prof. Osborn alludes to the scientific caution which led Kant, biology being
what it was, to refuse to entertain the hope "that a Newton may one day
arise even to make the production of a blade of grass comprehensible,
according to natural laws ordained by no intention." As Prof. Haeckel finely
observes, Darwin rose up as Kant's Newton.[7]
The scientific renaissance brought a wealth of fresh impressions and some
freedom from the tyranny of tradition, and the twofold stimulus stirred the
speculative activity of a great variety of men from old Claude Duret of

Moulins, of whose weird transformism (1609) Dr. Henry de Varigny[8]
gives us a glimpse, to Lorenz Oken (1779-1851) whose writings are such
mixtures of sense and nonsense that some regard him as a far-seeing
prophet and others as a fatuous follower of intellectual will-o'-the-wisps.
Similarly, for De Maillet, Maupertuis, Diderot, Bonnet, and others, we must
agree with Professor Osborn that they were not actually in the main
Evolution movement. Some have been included in the roll of honour on
very slender evidence, Robinet for instance, whose evolutionism seems to
us extremely dubious.[9]
The first naturalist to give a broad and concrete expression to the
evolutionist doctrine of descent was Buffon (1707-1788), but it is
interesting to recall the fact that his contemporary Linnæus (1707-1778),
protagonist of the counter-doctrine of the fixity of species,[10] went the
length of admitting (in 1762) that new species might arise by inter-crossing.
Buffon's position among the pioneers of the evolution-doctrine is weakened
by his habit of vacillating between his own conclusions and the orthodoxy
of the Sorbonne, but there is no doubt that he had firm grasp of the general
idea of "l'enchaînment des êtres."
Erasmus Darwin (1731-1802), probably influenced by Buffon, was another
firm evolutionist, and the outline of his argument in the Zoonomia[11] might
serve in part at least to-day. "When we revolve in our minds the
metamorphoses of animals, as from the tadpole to the frog; secondly, the
changes produced by artificial cultivation, as in the breeds of horses, dogs,
and sheep; thirdly, the changes produced by conditions of climate and of
season, as in the sheep of warm climates being covered with hair instead of
wool, and the hares and partridges of northern climates becoming white in
winter: when, further, we observe the changes of structure produced by
habit, as seen especially in men of different occupations; or the changes
produced by artificial mutilation and prenatal influences, as in the crossing
of species and production of monsters; fourth, when we observe the
essential unity of plan in all warm-blooded animals,—we are led to
conclude that they have been alike produced from a similar living
filament".... "From thus meditating upon the minute portion of time in
which many of the above changes have been produced, would it be too bold
to imagine, in the great length of time since the earth began to exist,
perhaps millions of years before the commencement of the history of

mankind, that all warm-blooded animals have arisen from one living
filament?"... "This idea of the gradual generation of all things seems to have
been as familiar to the ancient philosophers as to the modern ones, and to
have given rise to the beautiful hieroglyphic figure of the πρω̃τον ὠὁν, or
first great egg, produced by night, that is, whose origin is involved in
obscurity, and animated by Ἔρως, that is, by Divine Love; from whence
proceeded all things which exist."
Lamarck (1744-1829) seems to have become an evolutionist independently
of Erasmus Darwin's influence, though the parallelism between them is
striking. He probably owed something to Buffon, but he developed his
theory along a different line. Whatever view be held in regard to that theory
there is no doubt that Lamarck was a thorough-going evolutionist. Professor
Haeckel speaks of the Philosophie Zoologique as "the first connected and
thoroughly logical exposition of the theory of descent."[12]
Besides the three old masters, as we may call them, Buffon, Erasmus
Darwin, and Lamarck, there were other quite convinced pre-Darwinian
evolutionists. The historian of the theory of descent must take account of
Treviranus whose Biology or Philosophy of Animate Nature is full of
evolutionary suggestions; of Etienne Geoffroy St. Hilaire, who in 1830,
before the French Academy of Sciences, fought with Cuvier, the fellow-
worker of his youth, an intellectual duel on the question of descent; of
Goethe, one of the founders of morphology and the greatest poet of
Evolution—who, in his eighty-first year, heard the tidings of Geoffrey St.
Hilaire's defeat with an interest which transcended the political anxieties of
the time; and of many others who had gained with more or less confidence
and clearness a new outlook on Nature. It will be remembered that Darwin
refers to thirty-four more or less evolutionist authors in his Historical
Sketch, and the list might be added to. Especially when we come near to
1858 do the numbers increase, and one of the most remarkable, as also most
independent champions of the evolution-idea before that date was Herbert
Spencer, who not only marshalled the arguments in a very forcible way in
1852, but applied the formula in detail in his Principles of Psychology in
1855.[13]
It is right and proper that we should shake ourselves free from all creationist
appreciations of Darwin, and that we should recognise the services of pre-

Darwinian evolutionists who helped to make the time ripe, yet one cannot
help feeling that the citation of them is apt to suggest two fallacies. It may
suggest that Darwin simply entered into the labours of his predecessors,
whereas, as a matter of fact, he knew very little about them till after he had
been for years at work. To write, as Samuel Butler did, "Buffon planted,
Erasmus Darwin and Lamarck watered, but it was Mr. Darwin who said
'That fruit is ripe,' and shook it into his lap" ... seems to us a quite
misleading version of the facts of the case. The second fallacy which the
historical citation is a little apt to suggest is that the filiation of ideas is a
simple problem. On the contrary, the history of an idea, like the pedigree of
an organism, is often very intricate, and the evolution of the evolution-idea
is bound up with the whole progress of the world. Thus in order to interpret
Darwin's clear formulation of the idea of organic evolution and his
convincing presentation of it, we have to do more than go back to his
immediate predecessors, such as Buffon, Erasmus Darwin, and Lamarck;
we have to inquire into the acceptance of evolutionary conceptions in
regard to other orders of facts, such as the earth and the solar system;[14] we
have to realise how the growing success of scientific interpretation along
other lines gave confidence to those who refused to admit that there was
any domain from which science could be excluded as a trespasser; we have
to take account of the development of philosophical thought, and even of
theological and religious movements; we should also, if we are wise
enough, consider social changes. In short, we must abandon the idea that we
can understand the history of any science as such, without reference to
contemporary evolution in other departments of activity.
While there were many evolutionists before Darwin, few of them were
expert naturalists and few were known outside a small circle; what was of
much more importance was that the genetic view of Nature was insinuating
itself in regard to other than biological orders of facts, here a little and there
a little, and that the scientific spirit had ripened since the days when Cuvier
laughed Lamarck out of court. How was it that Darwin succeeded where
others had failed? Because, in the first place, he had clear visions
—"pensées de la jeunesse, executées par l'âge mûr"—which a University
curriculum had not made impossible, which the Beagle voyage made vivid,
which an unrivalled British doggedness made real—visions of the web of
life, of the fountain of change within the organism, of the struggle for

existence and its winnowing, and of the spreading genealogical tree.
Because, in the second place, he put so much grit into the verification of his
visions, putting them to the proof in an argument which is of its kind—
direct demonstration being out of the question—quite unequalled. Because,
in the third place, he broke down the opposition which the most scientific
had felt to the seductive modal formula of evolution by bringing forward a
more plausible theory of the process than had been previously suggested.
Nor can one forget, since questions of this magnitude are human and not
merely academic, that he wrote so that all men could understand.
As Regards the Factors of Evolution
It is admitted by all who are acquainted with the history of biology that the
general idea of organic evolution as expressed in the Doctrine of Descent
was quite familiar to Darwin's grandfather and to others before and after
him, as we have briefly indicated. It must also be admitted that some of
these pioneers of evolutionism did more than apply the evolution-idea as a
modal formula of becoming, they began to inquire into the factors in the
process. Thus there were pre-Darwinian theories of evolution, and to these
we must now briefly refer.[15]
In all biological thinking we have to work with the categories Organism—
Function—Environment, and theories of evolution may be classified in
relation to these. To some it has always seemed that the fundamental fact is
the living organism,—a creative agent, a striving will, a changeful Proteus,
selecting its environment, adjusting itself to it, self-differentiating and self-
adaptive. The necessity of recognising the importance of the organism is
admitted by all Darwinians who start with inborn variations, but it is open
to question whether the whole truth of what we might call the Goethian
position is exhausted in the postulate of inherent variability.
To others it has always seemed that the emphasis should be laid on
Function,—on use and disuse, on doing and not doing. Practice makes
perfect; c'est à force de forger qu'on devient forgeron. This is one of the
fundamental ideas of Lamarckism; to some extent it met with Darwin's
approval; and it finds many supporters to-day. One of the ablest of these—
Mr. Francis Darwin—has recently given strong reasons for combining a

modernised Lamarckism with what we usually regard as sound Darwinism.
[16]

To others it has always seemed that the emphasis should be laid on the
Environment, which wakes the organism to action, prompts it to change,
makes dints upon it, moulds it, prunes it, and finally, perhaps, kills it. It is
again impossible to doubt that there is truth in this view, for even if
environmentally induced "modifications" be not transmissible,
environmentally induced "variations" are; and even if the direct influence of
the environment be less important than many enthusiastic supporters of this
view—may we call them Buffonians—think, there remains the indirect
influence which Darwinians in part rely on,—the eliminative process. Even
if the extreme view be held that the only form of discriminate elimination
that counts is inter-organismal competition, this might be included under
the rubric of the animate environment.
In many passages Buffon[17] definitely suggested that environmental
influences—especially of climate and food—were directly productive of
changes in organisms, but he did not discuss the question of the
transmissibility of the modifications so induced, and it is difficult to gather
from his inconsistent writings what extent of transformation he really
believed in. Prof. Osborn says of Buffon: "The struggle for existence, the
elimination of the least-perfected species, the contest between the fecundity
of certain species and their constant destruction, are all clearly expressed in
various passages." He quotes two of these:[18]
"Le cours ordinaire de la nature vivante, est en général toujours constant,
toujours le même; son mouvement, toujours régulier, roule sur deux points
inébranlables: l'un, la fécondité sans bornes donnée à toutes les espèces;
l'autre, les obstacles sans nombre qui réduisent cette fécondité à une mesure
déterminée et ne laissent en tout temps qu'à peu près la même quantité
d'individus de chaque espèce" ... "Les espèces les moins parfaites, les plus
délicates, les plus pesantes, les moins agissantes, les moins armées, etc., ont
déjà disparu ou disparaîtront.".
Erasmus Darwin[19] had a firm grip of the "idea of the gradual formation
and improvement of the Animal world," and he had his theory of the
process. No sentence is more characteristic than this: "All animals undergo

transformations which are in part produced by their own exertions, in
response to pleasures and pains, and many of these acquired forms or
propensities are transmitted to their posterity." This is Lamarckism before
Lamarck, as his grandson pointed out. His central idea is that wants
stimulate efforts and that these result in improvements which subsequent
generations make better still. He realised something of the struggle for
existence and even pointed out that this advantageously checks the rapid
multiplication. "As Dr. Krause points out, Darwin just misses the
connection between this struggle and the Survival of the Fittest."[20]
Lamarck[21] (1744-1829) seems to have thought out his theory of evolution
without any knowledge of Erasmus Darwin's which it closely resembled.
The central idea of his theory was the cumulative inheritance of functional
modifications. "Changes in environment bring about changes in the habits
of animals. Changes in their wants necessarily bring about parallel changes
in their habits. If new wants become constant or very lasting, they form new
habits, the new habits involve the use of new parts, or a different use of old
parts, which results finally in the production of new organs and the
modification of old ones." He differed from Buffon in not attaching
importance, as far as animals are concerned, to the direct influence of the
environment, "for environment can effect no direct change whatever upon
the organisation of animals," but in regard to plants he agreed with Buffon
that external conditions directly moulded them.
Treviranus[22] (1776-1837), whom Huxley ranked beside Lamarck, was on
the whole Buffonian, attaching chief importance to the influence of a
changeful environment both in modifying and in eliminating, but he was
also Goethian, for instance in his idea that species like individuals pass
through periods of growth, full bloom, and decline. "Thus, it is not only the
great catastrophes of Nature which have caused extinction, but the
completion of cycles of existence, out of which new cycles have begun." A
characteristic sentence is quoted by Prof. Osborn: "In every living being
there exists a capability of an endless variety of form-assumption; each
possesses the power to adapt its organisation to the changes of the outer
world, and it is this power, put into action by the change of the universe,
that has raised the simple zoophytes of the primitive world to continually
higher stages of organisation, and has introduced a countless variety of
species into animate Nature."

Goethe[23] (1749-1832), who knew Buffon's work but not Lamarck's, is
peculiarly interesting as one of the first to use the evolution-idea as a
guiding hypothesis, e.g. in the interpretation of vestigial structures in man,
and to realise that organisms express an attempt to make a compromise
between specific inertia and individual change. He gave the finest
expression that science has yet known—if it has known it—of the kernel-
idea of what is called "bathmism," the idea of an "inherent growth-force"—
and at the same time he held that "the way of life powerfully reacts upon all
form" and that the orderly growth of form "yields to change from externally
acting causes."
Besides Buffon, Erasmus Darwin, Lamarck, Treviranus, and Goethe, there
were other "pioneers of evolution," whose views have been often discussed
and appraised. Étienne Geoffroy Saint-Hilaire (1772-1884), whose work
Goethe so much admired, was on the whole Buffonian, emphasising the
direct action of the changeful milieu. "Species vary with their environment,
and existing species have descended by modification from earlier and
somewhat simpler species." He had a glimpse of the selection idea, and
believed in mutations or sudden leaps—induced in the embryonic condition
by external influences. The complete history of evolution-theories will
include many instances of guesses at truth which were afterwards
substantiated, thus the geographer von Buch (1773-1853) detected the
importance of the Isolation factor on which Wagner, Romanes, Gulick and
others have laid great stress, but we must content ourselves with recalling
one other pioneer, the author of the Vestiges of Creation (1844), a work
which passed through ten editions in nine years and certainly helped to
harrow the soil for Darwin's sowing. As Darwin said, "it did excellent
service in this country in calling attention to the subject, in removing
prejudice, and in thus preparing the ground for the reception of analogous
views."[24] Its author, Robert Chambers (1802-1871) was in part a
Buffonian—maintaining that environment moulded organisms adaptively,
and in part a Goethian—believing in an inherent progressive impulse which
lifted organisms from one grade of organisation to another.
As Regards Natural Selection
The only thinker to whom Darwin was directly indebted, so far as the
theory of Natural Selection is concerned, was Malthus, and we may once

more quote the well-known passage in the Autobiography: "In October,
1838, that is, fifteen months after I had begun my systematic enquiry, I
happened to read for amusement 'Malthus on Population,' and being well
prepared to appreciate the struggle for existence which everywhere goes on
from long-continued observation of the habits of animals and plants, it at
once struck me that under these circumstances favourable variations would
tend to be preserved, and unfavourable ones to be destroyed. The result of
this would be the formation of new species."[25]
Although Malthus gives no adumbration of the idea of Natural Selection in
his exposition of the eliminative processes which go on in mankind, the
suggestive value of his essay is undeniable, as is strikingly borne out by the
fact that it gave to Alfred Russel Wallace also "the long-sought clue to the
effective agent in the evolution of organic species."[26] One day in Ternate
when he was resting between fits of fever, something brought to his
recollection the work of Malthus which he had read twelve years before. "I
thought of his clear exposition of 'the positive checks to increase'—disease,
accidents, war, and famine—which keep down the population of savage
races to so much lower an average than that of more civilized peoples. It
then occurred to me that these causes or their equivalents are continually
acting in the case of animals also; and as animals usually breed much more
rapidly than does mankind, the destruction every year from these causes
must be enormous in order to keep down the numbers of each species, since
they evidently do not increase regularly from year to year, as otherwise the
world would long ago have been densely crowded with those that breed
most quickly. Vaguely thinking over the enormous and constant destruction
which this implied, it occurred to me to ask the question, Why do some die
and some live? And the answer was clearly, that on the whole the best fitted
live. From the effects of disease the most healthy escaped; from enemies the
strongest, the swiftest, or the most cunning; from famine the best hunters or
those with the best digestion; and so on. Then it suddenly flashed upon me
that this self-acting process would necessarily improve the race, because in
every generation the inferior would inevitably be killed off and the superior
would remain—that is, the fittest would survive."[27] We need not apologise
for this long quotation, it is a tribute to Darwin's magnanimous colleague,
the Nestor of the evolutionist camp,—and it probably indicates the line of
thought which Darwin himself followed. It is interesting also to recall the

fact that in 1852, when Herbert Spencer wrote his famous Leader article on
"The Development Hypothesis" in which he argued powerfully for the
thesis that the whole animate world is the result of an age-long process of
natural transformation, he wrote for The Westminster Review another
important essay, "A Theory of Population deduced from the General Law of
Animal Fertility," towards the close of which he came within an ace of
recognising that the struggle for existence was a factor in organic evolution.
At a time when pressure of population was practically interesting men's
minds, Darwin, Wallace, and Spencer were being independently led from a
social problem to a biological theory. There could be no better illustration,
as Prof. Patrick Geddes has pointed out, of the Comtian thesis that science
is a "social phenomenon."
Therefore, as far more important than any further ferreting out of vague
hints of Natural Selection in books which Darwin never read, we would
indicate by a quotation the view that the central idea in Darwinism is
correlated with contemporary social evolution. "The substitution of Darwin
for Paley as the chief interpreter of the order of nature is currently regarded
as the displacement of an anthropomorphic view by a purely scientific one:
a little reflection, however, will show that what has actually happened has
been merely the replacement of the anthropomorphism of the eighteenth
century by that of the nineteenth. For the place vacated by Paley's
theological and metaphysical explanation has simply been occupied by that
suggested to Darwin and Wallace by Malthus in terms of the prevalent
severity of industrial competition, and those phenomena of the struggle for
existence which the light of contemporary economic theory has enabled us
to discern, have thus come to be temporarily exalted into a complete
explanation of organic progress."[28] It goes without saying that the idea
suggested by Malthus was developed by Darwin into a biological theory
which was then painstakingly verified by being used as an interpretative
formula, and that the validity of a theory so established is not affected by
what suggested it, but the practical question which this line of thought
raises in the mind is this: if Biology did thus borrow with such splendid
results from social theory, why should we not more deliberately repeat the
experiment?
Darwin was characteristically frank and generous in admitting that the
principle of Natural Selection had been independently recognised by Dr. W.

C. Wells in 1813 and by Mr. Patrick Matthew in 1831, but he had no
knowledge of these anticipations when he published the first edition of The
Origin of Species. Wells, whose "Essay on Dew" is still remembered, read
in 1813 before the Royal Society a short paper entitled "An Account of a
White Female, part of whose skin resembles that of a Negro" (published in
1818). In this communication, as Darwin said, "he observes, firstly, that all
animals tend to vary in some degree, and, secondly, that agriculturists
improve their domesticated animals by selection; and then, he adds, but
what is done in this latter case 'by art, seems to be done with equal efficacy,
though more slowly, by nature, in the formation of varieties of mankind,
fitted for the country which they inhabit.'"[29] Thus Wells had the clear idea
of survival dependent upon a favourable variation, but he makes no more
use of the idea and applies it only to man. There is not in the paper the least
hint that the author ever thought of generalising the remarkable sentence
quoted above.
Of Mr. Patrick Matthew, who buried his treasure in an appendix to a work
on Naval Timber and Arboriculture, Darwin said that "he clearly saw the
full force of the principle of natural selection." In 1860 Darwin wrote—very
characteristically—about this to Lyell: "Mr. Patrick Matthew publishes a
long extract from his work on Naval Timber and Arboriculture, published in
1831, in which he briefly but completely anticipates the theory of Natural
Selection. I have ordered the book, as some passages are rather obscure, but
it is certainly, I think, a complete but not developed anticipation. Erasmus
always said that surely this would be shown to be the case some day.
Anyhow, one may be excused in not having discovered the fact in a work
on Naval Timber."[30]
De Quatrefages and De Varigny have maintained that the botanist Naudin
stated the theory of evolution by natural selection in 1852. He explains very
clearly the process of artificial selection, and says that in the garden we are
following Nature's method. "We do not think that Nature has made her
species in a different fashion from that in which we proceed ourselves in
order to make our variations." But, as Darwin said, "he does not show how
selection acts under nature." Similarly it must be noted in regard to several
pre-Darwinian pictures of the struggle for existence (such as Herder's, who
wrote in 1790 "All is in struggle ... each one for himself" and so on), that a
recognition of this is only the first step in Darwinism.

Profs. E. Perrier and H. F. Osborn have called attention to a remarkable
anticipation of the selection-idea which is to be found in the speculations of
Étienne Geoffroy Saint-Hilaire (1825-1828) on the evolution of modern
Crocodilians from the ancient Teleosaurs. Changing environment induced
changes in the respiratory system and far-reaching consequences followed.
The atmosphere, acting upon the pulmonary cells, brings about
"modifications which are favourable or destructive ('funestes'); these are
inherited, and they influence all the rest of the organisation of the animal
because if these modifications lead to injurious effects the animals which
exhibit them perish and are replaced by others of a somewhat different
form, a form changed so as to be adapted to (à la convenance) the new
environment."
Prof. E. B. Poulton[31] has shown that the anthropologist James Cowles
Prichard (1786-1848) must be included even in spite of himself among the
precursors of Darwin. In some passages of the second edition of his
Researches into the Physical History of Mankind (1826), he certainly talks
evolution and anticipates Prof. Weismann in denying the transmission of
acquired characters. He is, however, sadly self-contradictory and his
evolutionism weakens in subsequent editions—the only ones that Darwin
saw. Prof. Poulton finds in Prichard's work a recognition of the operation of
Natural Selection. "After inquiring how it is that 'these varieties are
developed and preserved in connexion with particular climates and
differences of local situation,' he gives the following very significant
answer: 'One cause which tends to maintain this relation is obvious.
Individuals and families, and even whole colonies perish and disappear in
climates for which they are, by peculiarity of constitution, not adapted. Of
this fact proofs have been already mentioned.'" Mr. Francis Darwin and
Prof. A. C. Seward discuss Prichard's "anticipations" in More Letters of
Charles Darwin, Vol. I. p. 43, and come to the conclusion that the
evolutionary passages are entirely neutralised by others of an opposite
trend. There is the same difficulty with Buffon.
Hints of the idea of Natural Selection have been detected elsewhere. James
Watt,[32] for instance, has been reported as one of the anticipators (1851).
But we need not prolong the inquiry further, since Darwin did not know of
any anticipations until after he had published the immortal work of 1859,
and since none of those who got hold of the idea made any use of it. What

Darwin did was to follow the clue which Malthus gave him, to realise, first
by genius and afterwards by patience, how the complex and subtle struggle
for existence works out a natural selection of those organisms which vary in
the direction of fitter adaptation to the conditions of their life. So much
success attended his application of the Selection-formula that for a time he
regarded Natural Selection as almost the sole factor in evolution, variations
being pre-supposed; gradually, however, he came to recognise that there
was some validity in the factors which had been emphasised by Lamarck
and by Buffon, and in his well known summing up in the sixth edition of
the Origin he says of the transformation of species: "This has been effected
chiefly through the natural selection of numerous successive, slight,
favourable variations; aided in an important manner by the inherited effects
of the use and disuse of parts; and in an unimportant manner, that is, in
relation to adaptive structures, whether past or present, by the direct action
of external conditions, and by variations which seem to us in our ignorance
to arise spontaneously."
To sum up: the idea of organic evolution, older than Aristotle, slowly
developed from the stage of suggestion to the stage of verification, and the
first convincing verification was Darwin's; from being an a priori
anticipation it has become an interpretation of nature, and Darwin is still the
chief interpreter; from being a modal interpretation it has advanced to the
rank of a causal theory, the most convincing part of which men will never
cease to call Darwinism.

FOOTNOTES:
[1] Columbia University Biological Series, Vol. I. New York and London, 1894.
We must acknowledge our great indebtedness to this fine piece of work.
[2] op. cit. p. 41.
[3] See G. J. Romanes, "Aristotle as a Naturalist," Contemporary Review, Vol.
lix. p. 275, 1891; G. Pouchet, La Biologie Aristotélique, Paris, 1885; E. Zeller, A
History of Greek Philosophy, London, 1881, and "Ueber die griechischen
Vorgänger Darwin's," Abhandl. Berlin Akad. 1878, pp. 111-124.
[4] op. cit. p. 81.
[5] op. cit. p. 87.

[6] See Brock, "Die Stellung Kant's zur Deszendenztheorie," Biol. Centralbl.
viii. 1889, pp. 641-648. Fritz Schultze, Kant und Darwin, Jena, 1875.
[7] Mr. Alfred Russel Wallace writes: "We claim for Darwin that he is the
Newton of natural history, and that, just so surely as that the discovery and
demonstration by Newton of the law of gravitation established order in place of
chaos and laid a sure foundation for all future study of the starry heavens, so
surely has Darwin, by his discovery of the law of natural selection and his
demonstration of the great principle of the preservation of useful variations in
the struggle for life, not only thrown a flood of light on the process of
development of the whole organic world, but also established a firm foundation
for all future study of nature" (Darwinism, London, 1889, p. 9). See also Prof.
Karl Pearson's Grammar of Science (2nd edit.), London, 1900, p. 32. See
Osborn, op. cit. p. 100.
[8] Experimental Evolution. London, 1892. Chap. I. p. 14.
[9] See J. Arthur Thomson, The Science of Life. London, 1899, Chap. XVI.
"Evolution of Evolution Theory."
[10] See Carus Sterne (Ernst Krause), Die allgemeine Weltanschauung in ihrer
historischen Entwickelung. Stuttgart, 1889. Chapter entitled "Beständigkeit oder
Veränderlichkeit der Naturwesen."
[11] Zoonomia, or the Laws of Organic Life, 2 vols. London, 1794; Osborn, op.
cit. p. 145.
[12] See Alpheus S. Packard, Lamarck, the Founder of Evolution, His Life and
Work, with Translations of his writings on Organic Evolution. London, 1901.
[13] See Edward Clodd, Pioneers of Evolution, London, p. 161, 1897.
[14] See Chapter ix. "The Genetic View of Nature" in J. T. Merz's History of
European Thought in the Nineteenth Century, Vol. 2, Edinburgh and London,
1903.
[15] See Prof. W. A. Locy's Biology and its Makers. New York, 1908. Part II.
"The Doctrine of Organic Evolution."
[16] Presidential Address to the British Association meeting at Dublin in 1908.
[17] See in particular Samuel Butler, Evolution Old and New, London, 1879; J.
L. de Lanessan, "Buffon et Darwin," Revue Scientifique, XLIII. pp. 385-391,
425-432, 1889.
[18] op. cit. p. 136.
[19] See Ernest Krause and Charles Darwin, Erasmus Darwin, London, 1879.
[20] Osborn, op. cit. p. 142.
[21] See E. Perrier, La Philosophie Zoologique avant Darwin, Paris, 1884; A. de
Quatrefages, Darwin et ses Précurseurs Français, Paris, 1870; Packard, op. cit.;
also Claus, Lamarck als Begründer der Descendenzlehre, Wien, 1888; Haeckel,

Natural History of Creation, Eng. transl. London, 1879; Lang, Zur
Charakteristik der Forschungswege von Lamarck und Darwin, Jena, 1889.
[22] See Huxley's article "Evolution in Biology," Encyclopaedia Britannica (9th
edit.), 1879, pp. 744-751, and Sully's article, "Evolution in Philosophy," ibid. pp.
751-772.
[23] See Haeckel, Die Naturanschauung von Darwin, Goethe und Lamarck,
Jena, 1882.
[24] Origin of Species (6th edit.), p. xvii.
[25] The Life and Letters of Charles Darwin, Vol. 1. p. 83. London, 1887.
[26] A. R. Wallace, My Life, a Record of Events and Opinions, London, 1905,
Vol. 1, p. 232.
[27] My Life, Vol. 1. p. 361.
[28] P. Geddes. article "Biology." Chambers's Encyclopaedia.
[29] Origin of Species (6th edit.), p. xv.
[30] Life and Letters, ii, p. 301.
[31] Science Progress, New Series, Vol. 1. 1897. "A Remarkable Anticipation of
Modern Views on Evolution." See also Chap. vi. in Essays on Evolution, Oxford,
1908.
[32] See Prof. Patrick Geddes's article "Variation and Selection," Encyclopaedia
Britannica (9th edit.) 1888.

II

THE SELECTION THEORY
By August Weismann

Professor of Zoology in the University of Freiburg (Baden)

I. The Idea of Selection

Many and diverse were the discoveries made by Charles Darwin in the
course of a long and strenuous life, but none of them has had so far-
reaching an influence on the science and thought of his time as the theory of
selection. I do not believe that the theory of evolution would have made its
way so easily and so quickly after Darwin took up the cudgels in favour of
it if he had not been able to support it by a principle which was capable of
solving, in a simple manner, the greatest riddle that living nature presents to
us,—I mean the purposiveness of every living form relative to the
conditions of its life and its marvellously exact adaptation to these.
Everyone knows that Darwin was not alone in discovering the principle of
selection, and that the same idea occurred simultaneously and
independently to Alfred Russel Wallace. At the memorable meeting of the
Linnean Society on 1st July, 1858, two papers were read (communicated by
Lyell and Hooker) both setting forth the same idea of selection. One was
written by Charles Darwin in Kent, the other by Alfred Wallace in Ternate,
in the Malay Archipelago. It was a splendid proof of the magnanimity of
these two investigators, that they thus in all friendliness and without envy,
united in laying their ideas before a scientific tribunal: their names will
always shine side by side as two of the brightest stars in the scientific sky.
The idea of selection set forth by the two naturalists was at the time
absolutely new, but it was also so simple that Huxley could say of it later,
"How extremely stupid not to have thought of that." As Darwin was led to
the general doctrine of descent, not through the labours of his predecessors
in the early years of the century, but by his own observations, so it was in

regard to the principle of selection. He was struck by the innumerable cases
of adaptation, as, for instance, that of the woodpeckers and tree-frogs to
climbing, or the hooks and feather-like appendages of seeds, which aid in
the distribution of plants, and he said to himself that an explanation of
adaptations was the first thing to be sought for in attempting to formulate a
theory of evolution.
But since adaptations point to changes which have been undergone by the
ancestral forms of existing species, it is necessary, first of all, to inquire
how far species in general are variable. Thus Darwin's attention was
directed in the first place to the phenomenon of variability, and the use man
has made of this, from very early times, in the breeding of his domesticated
animals and cultivated plants. He inquired carefully how breeders set to
work, when they wished to modify the structure and appearance of a species
to their own ends, and it was soon clear to him that selection for breeding
purposes played the chief part.
But how was it possible that such processes should occur in free nature?
Who is here the breeder, making the selection, choosing out one individual
to bring forth offspring and rejecting others? That was the problem that for
a long time remained a riddle to him.
Darwin himself relates how illumination suddenly came to him. He had
been reading, for his own pleasure, Malthus' book on Population, and, as he
had long known from numerous observations, that every species gives rise
to many more descendants than ever attain to maturity, and that, therefore,
the greater number of the descendants of a species perish without
reproducing, the idea came to him that the decision as to which member of
a species was to perish and which was to attain to maturity and reproduction
might not be a matter of chance, but might be determined by the
constitution of the individuals themselves, according as they were more or
less fitted for survival. With this idea the foundation of the theory of
selection was laid.
In artificial selection the breeder chooses out for pairing only such
individuals as possess the character desired by him in a somewhat higher
degree than the rest of the race. Some of the descendants inherit this
character, often in a still higher degree, and if this method be pursued

throughout several generations, the race is transformed in respect of that
particular character.
Natural selection depends on the same three factors as artificial selection:
on variability, inheritance, and selection for breeding, but this last is here
carried out not by a breeder but by what Darwin called the "struggle for
existence." This last factor is one of the special features of the Darwinian
conception of nature. That there are carnivorous animals which take heavy
toll in every generation of the progeny of the animals on which they prey,
and that there are herbivores which decimate the plants in every generation
had long been known, but it is only since Darwin's time that sufficient
attention has been paid to the facts that, in addition to this regular
destruction, there exists between the members of a species a keen
competition for space and food, which limits multiplication, and that
numerous individuals of each species perish because of unfavourable
climatic conditions. The "struggle for existence," which Darwin regarded as
taking the place of the human breeder in free nature, is not a direct struggle
between carnivores and their prey, but is the assumed competition for
survival between individuals of the same species, of which, on an average,
only those survive to reproduce which have the greatest power of
resistance, while the others, less favourably constituted, perish early. This
struggle is so keen, that, within a limited area, where the conditions of life
have long remained unchanged, of every species, whatever be the degree of
fertility, only two, on an average, of the descendants of each pair survive;
the others succumb either to enemies, or to disadvantages of climate, or to
accident. A high degree of fertility is thus not an indication of the special
success of a species, but of the numerous dangers that have attended its
evolution. Of the six young brought forth by a pair of elephants in the
course of their lives only two survive in a given area; similarly, of the
millions of eggs which two thread-worms leave behind them only two
survive. It is thus possible to estimate the dangers which threaten a species
by its ratio of elimination, or, since this cannot be done directly, by its
fertility.
Although a great number of the descendants of each generation fall victims
to accident, among those that remain it is still the greater or less fitness of
the organism that determines the "selection for breeding purposes," and it
would be incomprehensible if, in this competition, it were not ultimately,

that is, on an average, the best equipped which survive, in the sense of
living long enough to reproduce.
Thus the principle of natural selection is the selection of the best for
reproduction, whether the "best" refers to the whole constitution, to one or
more parts of the organism, or to one or more stages of development. Every
organ, every part, every character of an animal, fertility and intelligence
included, must be improved in this manner, and be gradually brought up in
the course of generations to its highest attainable state of perfection. And
not only may improvement of parts be brought about in this way, but new
parts and organs may arise, since, through the slow and minute steps of
individual or "fluctuating" variations, a part may be added here or dropped
out there, and thus something new is produced.
The principle of selection solved the riddle as to how what was purposive
could conceivably be brought about without the intervention of a directing
power, the riddle which animate nature presents to our intelligence at every
turn, and in face of which the mind of a Kant could find no way out, for he
regarded a solution of it as not to be hoped for. For, even if we were to
assume an evolutionary force that is continually transforming the most
primitive and the simplest forms of life into ever higher forms, and the
homogeneity of primitive times into the infinite variety of the present, we
should still be unable to infer from this alone how each of the numberless
forms adapted to particular conditions of life should have appeared
precisely at the right moment in the history of the earth to which their
adaptations were appropriate, and precisely at the proper place in which all
the conditions of life to which they were adapted occurred: the humming-
birds at the same time as the flowers; the trichina at the same time as the
pig; the bark-coloured moth at the same time as the oak, and the wasp-like
moth at the same time as the wasp which protects it. Without processes of
selection we should be obliged to assume a "pre-established harmony" after
the famous Leibnitzian model, by means of which the clock of the evolution
of organisms is so regulated as to strike in exact synchronism with that of
the history of the earth! All forms of life are strictly adapted to the
conditions of their life, and can persist under these conditions alone.
There must therefore be an intrinsic connection between the conditions and
the structural adaptations of the organism, and, since the conditions of life

cannot be determined by the animal itself, the adaptations must be called
forth by the conditions.
The selection theory teaches us how this is conceivable, since it enables us
to understand that there is a continual production of what is non-purposive
as well as of what is purposive, but the purposive alone survives, while the
non-purposive perishes in the very act of arising. This is the old wisdom
taught long ago by Empedocles.
II. The Lamarckian Principle
Lamarck, as is well known, formulated a definite theory of evolution at the
beginning of the nineteenth century, exactly fifty years before the Darwin-
Wallace principle of selection was given to the world. This brilliant
investigator also endeavoured to support his theory by demonstrating forces
which might have brought about the transformations of the organic world in
the course of the ages. In addition to other factors, he laid special emphasis
on the increased or diminished use of the parts of the body, assuming that
the strengthening or weakening which takes place from this cause during
the individual life, could be handed on to the offspring, and thus intensified
and raised to the rank of a specific character. Darwin also regarded this
Lamarckian principle, as it is now generally called, as a factor in evolution,
but he was not fully convinced of the transmissibility of acquired
characters.
As I have here to deal only with the theory of selection, I need not discuss
the Lamarckian hypothesis, but I must express my opinion that there is
room for much doubt as to the coöperation of this principle in evolution.
Not only is it difficult to imagine how the transmission of functional
modifications could take place, but, up to the present time, notwithstanding
the endeavours of many excellent investigators, not a single actual proof of
such inheritance has been brought forward. Semon's experiments on plants
are, according to the botanist Pfeffer, not to be relied on, and even the
recent, beautiful experiments made by Dr. Kammerer on salamanders,
cannot, as I hope to show elsewhere, be regarded as proof, if only because
they do not deal at all with functional modifications, that is, with
modifications brought about by use, and it is to these alone that the
Lamarckian principle refers.

III. Objections to the Theory of Selection
(a) Saltatory evolution
The Darwinian doctrine of evolution depends essentially on the cumulative
augmentation of minute variations in the direction of utility. But can such
minute variations, which are undoubtedly continually appearing among the
individuals of the same species, possess any selection-value; can they
determine which individuals are to survive, and which are to succumb; can
they be increased by natural selection till they attain to the highest
development of a purposive variation?
To many this seems so improbable that they have urged a theory of
evolution by leaps from species to species. Kölliker, in 1872, compared the
evolution of species with the processes which we can observe in the
individual life in cases of alternation of generations. But a polyp only gives
rise to a medusa because it has itself arisen from one, and there can be no
question of a medusa ever having arisen suddenly and de novo from a
polyp-bud, if only because both forms are adapted in their structure as a
whole, and in every detail to the conditions of their life. A sudden origin, in
a natural way, of numerous adaptations is inconceivable. Even the
degeneration of a medusoid from a free-swimming animal to a mere brood-
sac (gonophore) is not sudden and saltatory, but occurs by imperceptible
modifications throughout hundreds of years, as we can learn from the
numerous stages of the process of degeneration persisting at the same time
in different species.
If, then, the degeneration to a simple brood-sac takes place only by very
slow transitions, each stage of which may last for centuries, how could the
much more complex ascending evolution possibly have taken place by
sudden leaps? I regard this argument as capable of further extension, for
wherever in nature we come upon degeneration, it is taking place by minute
steps and with a slowness that makes it not directly perceptible, and I
believe that this in itself justifies us in concluding that the same must be
true of ascending evolution. But in the latter case the goal can seldom be
distinctly recognised while in cases of degeneration the starting-point of the
process can often be inferred, because several nearly related species may
represent different stages.

In recent years Bateson in particular has championed the idea of saltatory,
or so-called discontinuous evolution, and has collected a number of cases in
which more or less marked variations have suddenly appeared. These are
taken for the most part from among domesticated animals which have been
bred and crossed for a long time, and it is hardly to be wondered at that
their much mixed and much influenced germ-plasm should, under certain
conditions, give rise to remarkable phenomena, often indeed producing
forms which are strongly suggestive of monstrosities, and which would
undoubtedly not survive in free nature, unprotected by man. I should regard
such cases as due to an intensified germinal selection—though this is to
anticipate a little—and from this point of view it cannot be denied that they
have a special interest. But they seem to me to have no significance as far
as the transformation of species is concerned, if only because of the extreme
rarity of their occurrence.
There are, however, many variations which have appeared in a sudden and
saltatory manner, and some of these Darwin pointed out and discussed in
detail: the copper beech, the weeping trees, the oak with "fern-like leaves,"
certain garden-flowers, etc. But none of them have persisted in free nature,
or evolved into permanent types.
On the other hand, wherever enduring types have arisen, we find traces of a
gradual origin by successive stages, even if, at first sight, their origin may
appear to have been sudden. This is the case with seasonal Dimorphism, the
first known cases of which exhibited marked differences between the two
generations, the winter and the summer brood. Take for instance the much
discussed and studied form Vanessa (Araschnia) levana-prorsa. Here the
differences between the two forms are so great and so apparently
disconnected, that one might almost believe it to be a sudden mutation,
were it not that old transition-stages can be called forth by particular
temperatures, and we know other butterflies, as for instance our Garden
Whites, in which the differences between the two generations are not nearly
so marked; indeed, they are so little apparent that they are scarcely likely to
be noticed except by experts. Thus here again there are small initial steps,
some of which, indeed, must be regarded as adaptations, such as the green-
sprinkled or lightly tinted under-surface which gives them a deceptive
resemblance to parsley or to Cardamine leaves.

Even if saltatory variations do occur, we cannot assume that these have ever
led to forms which are capable of survival under the conditions of wild life.
Experience has shown that in plants which have suddenly varied the power
of persistence is diminished. Korschinsky attributes to them weaknesses of
organisation in general; "they bloom late, ripen few of their seeds, and show
great sensitiveness to cold." These are not the characters which make for
success in the struggle for existence.
We must briefly refer here to the views—much discussed in the last decade
—of H. de Vries, who believes that the roots of transformation must be
sought for in saltatory variations arising from internal causes, and
distinguishes such mutations, as he has called them, from ordinary
individual variations, in that they breed true, that is, with strict in-breeding
they are handed on pure to the next generation. I have elsewhere
endeavoured to point out the weaknesses of this theory,[33] and I am the less
inclined to return to it here that it now appears[34] that the far-reaching
conclusions drawn by de Vries from his observations on the Evening
Primrose, Oenothera lamarckiana, rest upon a very insecure foundation.
The plant from which de Vries saw numerous "species"—his "mutations"—
arise was not, as he assumed, a wild species that had been introduced to
Europe from America, but was probably a hybrid form which was first
discovered in the Jardin des Plantes in Paris, and which does not appear to
exist anywhere in America as a wild species.
This gives a severe shock to the "Mutation theory," for the other actually
wild species with which de Vries experimented showed no "mutations" but
yielded only negative results.
Thus we come to the conclusion that Darwin[35] was right in regarding
transformations as taking place by minute steps, which, if useful, are
augmented in the course of innumerable generations, because their
possessors more frequently survive in the struggle for existence.
(b) Selection-value of the initial steps
Is it possible that the insignificant deviations which we know as "individual
variations" can form the beginning of a process of selection? Can they
decide which is to perish and which to survive? To use a phrase of
Romanes, can they have selection-value?

Darwin himself answered this question, and brought together many
excellent examples to show that differences, apparently insignificant
because very small, might be of decisive importance for the life of the
possessor. But it is by no means enough to bring forward cases of this kind,
for the question is not merely whether finished adaptations have selection-
value, but whether the first beginnings of these, and whether the small, I
might almost say minimal increments, which have led up from these
beginnings to the perfect adaptation, have also had selection-value. To this
question even one who, like myself, has been for many years a convinced
adherent of the theory of selection, can only reply: We must assume so, but
we cannot prove it in any case. It is not upon demonstrative evidence that
we rely when we champion the doctrine of selection as a scientific truth; we
base our argument on quite other grounds. Undoubtedly there are many
apparently insignificant features, which can nevertheless be shown to be
adaptations—for instance, the thickness of the basin-shaped shell of the
limpets that live among the breakers on the shore. There can be no doubt
that the thickness of these shells, combined with their flat forms, protects
the animals from the force of the waves breaking upon them,—but how
have they become so thick? What proportion of thickness was sufficient to
decide that of two variants of a limpet one should survive, the other be
eliminated? We can say nothing more than that we infer from the present
state of the shell, that it must have varied in regard to differences in shell-
thickness, and that these differences must have had selection-value,—no
proof therefore, but an assumption which we must show to be convincing.
For a long time the marvellously complex radiate and lattice-work
skeletons of Radiolarians were regarded as a mere outflow of "Nature's
infinite wealth of form," as an instance of a purely morphological character
with no biological significance. But recent investigations have shown that
these, too, have an adaptive significance (Häcker). The same thing has been
shown by Schütt in regard to the lowly unicellular plants, the Peridineae,
which abound alike on the surface of the ocean and in its depths. It has been
shown that the long skeletal processes which grow out from these
organisms have significance not merely as a supporting skeleton, but also as
an extension of the superficial area, which increases the contact with the
water-particles, and prevents the floating organisms from sinking. It has
been established that the processes are considerably shorter in the colder

layers of the ocean, and that they may be twelve times as long[36] in the
warmer layers, thus corresponding to the greater or smaller amount of
friction which takes place in the denser and less dense layers of the water.
The Peridineae of the warmer ocean layers have thus become long-rayed,
those of the colder layers short-rayed, not through the direct effect of
friction on the protoplasm, but through processes of selection, which
favoured the longer rays in warm water, since they kept the organism afloat,
while those with short rays sank and were eliminated. If we put the question
as to selection-value in this case, and ask how great the variations in the
length of processes must be in order to possess selection-value; what can
we answer except that these variations must have been minimal, and yet
sufficient to prevent too rapid sinking and consequent elimination? Yet this
very case would give the ideal opportunity for a mathematical calculation of
the minimal selection-value, although of course it is not feasible from lack
of data to carry out the actual calculation.
But even in organisms of more than microscopic size there must frequently
be minute, even microscopic differences which set going the process of
selection, and regulate its progress to the highest possible perfection.
Many tropical trees possess thick, leathery leaves, as a protection against
the force of the tropical raindrops. The direct influence of the rain cannot be
the cause of this power of resistance, for the leaves, while they were still
thin, would simply have been torn to pieces. Their toughness must therefore
be referred to selection, which would favour the trees with slightly thicker
leaves, though we cannot calculate with any exactness how great the first
stages of increase in thickness must have been. Our hypothesis receives
further support from the fact that, in many such trees, the leaves are drawn
out into a beak-like prolongation (Stahl and Haberlandt) which facilitates
the rapid falling off of the rain water, and also from the fact that the leaves,
while they are still young, hang limply down in bunches which offer the
least possible resistance to the rain. Thus there are here three adaptations
which can only be interpreted as due to selection. The initial stages of these
adaptations must undoubtedly have had selection-value.
But even in regard to this case we are reasoning in a circle, not giving
"proofs," and no one who does not wish to believe in the selection-value of

the initial stages can be forced to do so. Among the many pieces of
presumptive evidence a particularly weighty one seems to me to be the
smallness of the steps of progress which we can observe in certain cases, as
for instance in leaf-imitation among butterflies, and in mimicry generally.
The resemblance to a leaf, for instance of a particular Kallima, seems to us
so close as to be deceptive, and yet we find in another individual, or it may
be in many others, a spot added which increases the resemblance, and
which could not have become fixed unless the increased deceptiveness so
produced had frequently led to the overlooking of its much persecuted
possessor. But if we take the selection-value of the initial stages for granted,
we are confronted with the further question which I myself formulated
many years ago: How does it happen that the necessary beginnings of a
useful variation are always present? How could insects which live upon or
among green leaves become all green, while those that live on bark become
brown? How have the desert animals become yellow and the Arctic animals
white? Why were the necessary variations always present? How could the
green locust lay brown eggs, or the privet caterpillar develop white and
lilac-coloured lines on its green skin?
It is of no use answering to this that the question is wrongly formulated[37]
and that it is the converse that is true; that the process of selection takes
place in accordance with the variations that present themselves. This
proposition is undeniably true, but so also is another, which apparently
negatives it: the variation required has in the majority of cases actually
presented itself. Selection cannot solve this contradiction; it does not call
forth the useful variation, but simply works upon it. The ultimate reason
why one and the same insect should occur in green and in brown, as often
happens in caterpillars and locusts, lies in the fact that variations towards
brown presented themselves, and so also did variations towards green: the
kernel of the riddle lies in the varying, and for the present we can only say,
that small variations in different directions present themselves in every
species. Otherwise so many different kinds of variations could not have
arisen. I have endeavoured to explain this remarkable fact by means of the
intimate processes that must take place within the germ-plasm, and I shall
return to the problem when dealing with "germinal selection."
We have, however, to make still greater demands on variation, for it is not
enough that the necessary variation should occur in isolated individuals,

because in that case there would be small prospect of its being preserved,
notwithstanding its utility. Darwin at first believed, that even single
variations might lead to transformation of the species, but later he became
convinced that this was impossible, at least without the coöperation of other
factors, such as isolation and sexual selection.
In the case of the green caterpillars with bright longitudinal stripes,
numerous individuals exhibiting this useful variation must have been
produced to start with. In all higher, that is, multicellular organisms, the
germ-substance is the source of all transmissible variations, and this germ-
plasm is not a simple substance but is made up of many primary
constituents. The question can therefore be more precisely stated thus: How
does it come about that in so many cases the useful variations present
themselves in numbers just where they are required, the white oblique lines
in the leaf-caterpillar on the under surface of the body, the accompanying
coloured stripes just above them? And, further, how has it come about that
in grass caterpillars, not oblique but longitudinal stripes, which are more
effective for concealment among grass and plants, have been evolved? And
finally, how is it that the same Hawk-moth caterpillars, which to-day show
oblique stripes, possessed longitudinal stripes in Tertiary times? We can
read this fact from the history of their development, and I have before
attempted to show the biological significance of this change of colour.[38]
For the present I need only draw the conclusion that one and the same
caterpillar may exhibit the initial stages of both, and that it depends on the
manner in which these marking elements are intensified and combined by
natural selection whether whitish longitudinal or oblique stripes should
result. In this case then the "useful variations" were actually "always there,"
and we see that in the same group of Lepidoptera, e.g. species of
Sphingidae, evolution has occurred in both directions according to whether
the form lived among grass or on broad leaves with oblique lateral veins,
and we can observe even now that the species with oblique stripes have
longitudinal stripes when young, that is to say, while the stripes have no
biological significance. The white places in the skin which gave rise,
probably first as small spots, to this protective marking could be combined
in one way or another according to the requirements of the species. They
must therefore either have possessed selection-value from the first, or, if
this was not the case at their earliest occurrence, there must have been some

other factors which raised them to the point of selection-value. I shall return
to this in discussing germinal selection. But the case may be followed still
farther, and leads us to the same alternative on a still more secure basis.
Many years ago I observed in caterpillars of Smerinthus populi (the poplar
hawk-moth), which also possess white oblique stripes, that certain
individuals showed red spots above these stripes; these spots occurred only
on certain segments, and never flowed together to form continuous stripes.
In another species (Smerinthus tiliae) similar blood-red spots unite to form
a line-like coloured seam in the last stage of larval life, while in S. ocellata
rust-red spots appear in individual caterpillars, but more rarely than in S.
populi, and they show no tendency to flow together.
Thus we have here the origin of a new character, arising from small
beginnings, at least in S. tiliae, in which species the coloured stripes are a
normal specific character. In the other species, S. populi and S. ocellata, we
find the beginnings of the same variation, in one more rarely than in the
other, and we can imagine that, in the course of time, in these two species,
coloured lines over the oblique stripes will arise. In any case these spots are
the elements of variation, out of which coloured lines may be evolved, if
they are combined in this direction through the agency of natural selection.
In S. populi the spots are often small, but sometimes it seems as though
several had united to form large spots. Whether a process of selection in this
direction will arise in S. populi and S. ocellata, or whether it is now going
on cannot be determined, since we cannot tell in advance what biological
value the marking might have for these two species. It is conceivable that
the spots may have no selection-value as far as these species are concerned,
and may therefore disappear again in the course of phylogeny, or, on the
other hand, that they may be changed in another direction, for instance
towards imitation of the rust-red fungoid patches on poplar and willow
leaves. In any case we may regard the smallest spots as the initial stages of
variation, the larger as a cumulative summation of these. Therefore either
these initial stages must already possess selection-value, or, as I said before:
There must be some other reason for their cumulative summation. I should
like to give one more example, in which we can infer, though we cannot
directly observe, the initial stages.

All the Holothurians or sea-cucumbers have in the skin calcereous bodies of
different forms, usually thick and irregular, which make the skin tough and
resistant. In a small group of them—the species of Synapta—the calcareous
bodies occur in the form of delicate anchors of microscopic size. Up till
1897 these anchors, like many other delicate microscopic structures, were
regarded as curiosities, as natural marvels. But a Swedish observer,
Oestergren, has recently shown that they have a biological significance:
they serve the footless Synapta as auxiliary organs of locomotion, since,
when the body swells up in the act of creeping, they press firmly with their
tips, which are embedded in the skin, against the substratum on which the
animal creeps, and thus prevent slipping backwards. In other Holothurians
this slipping is made impossible by the fixing of the tube-feet. The anchors
act automatically, sinking their tips towards the ground when the
corresponding part of the body thickens, and returning to the original
position at an angle of 45 degrees to the upper surface when the part
becomes thin again. The arms of the anchor do not lie in the same plane as
the shaft, and thus the curve of the arms forms the outermost part of the
anchor, and offers no further resistance to the gliding of the animal. Every
detail of the anchor, the curved portion, the little teeth at the head, the arms,
etc., can be interpreted in the most beautiful way, above all the form of the
anchor itself, for the two arms prevent it from swaying round to the side.
The position of the anchors, too, is definite and significant; they lie
obliquely to the longitudinal axis of the animal, and therefore they act alike
whether the animal is creeping backwards or forwards. Moreover, the tips
would pierce through the skin if the anchors lay in the longitudinal
direction. Synapta burrows in the sand; it first pushes in the thin anterior
end, and thickens this again, thus enlarging the hole, then the anterior
tentacles displace more sand, the body is worked in a little farther, and the
process begins anew. In the first act the anchors are passive, but they begin
to take an active share in the forward movement when the body is
contracted again. Frequently the animal retains only the posterior end
buried in the sand, and then the anchors keep it in position, and make rapid
withdrawal possible.
Thus we have in these apparently random forms of the calcereous bodies,
complex adaptations in which every little detail as to direction, curve, and
pointing is exactly determined. That they have selection-value in their

present perfected form is beyond all doubt, since the animals are enabled by
means of them to bore rapidly into the ground and so to escape from
enemies. We do not know what the initial stages were, but we cannot doubt
that the little improvements, which occurred as variations of the originally
simple slimy bodies of the Holothurians, were preserved because they
already possessed selection-value for the Synaptidae. For such minute
microscopic structures whose form is so delicately adapted to the rôle they
have to play in the life of the animal, cannot have arisen suddenly and as a
whole, and every new variation of the anchor, that is, in the direction of the
development of the two arms, and every curving of the shaft which
prevented the tips from projecting at the wrong time, in short, every little
adaptation in the modelling of the anchor must have possessed selection-
value. And that such minute changes of form fall within the sphere of
fluctuating variations, that is to say, that they occur is beyond all doubt.
In many of the Synaptidae the anchors are replaced by calcareous rods bent
in the form of an S, which are said to act in the same way. Others, such as
those of the genus Ankyroderma, have anchors which project considerably
beyond the skin, and, according to Oestergren, serve "to catch plant-
particles and other substances" and so mask the animal. Thus we see that in
the Synaptidae the thick and irregular calcareous bodies of the Holothurians
have been modified and transformed in various ways in adaptation to the
footlessness of these animals, and to the peculiar conditions of their life,
and we must conclude that the earlier stages of these changes presented
themselves to the processes of selection in the form of microscopic
variations. For it is as impossible to think of any origin other than through
selection in this case as in the case of the toughness, and the "drip-tips" of
tropical leaves. And as these last could not have been produced directly by
the beating of the heavy raindrops upon them, so the calcareous anchors of
Synapta cannot have been produced directly by the friction of the sand and
mud at the bottom of the sea, and, since they are parts whose function is
passive the Lamarckian factor of use and disuse does not come into
question. The conclusion is unavoidable, that the microscopically small
variations of the calcareous bodies in the ancestral forms have been
intensified and accumulated in a particular direction, till they have led to the
formation of the anchor. Whether this has taken place by the action of
natural selection alone, or whether the laws of variation and the intimate

processes within the germ-plasm have coöperated will become clear in the
discussion of germinal selection. This whole process of adaptation has
obviously taken place within the time that has elapsed since this group of
sea-cucumbers lost their tube-feet, those characteristic organs of locomotion
which occur in no group except the Echinoderms, and yet have totally
disappeared in the Synaptidae. And after all what would animals that live in
sand and mud do with tube-feet?
(c) Coadaptation
Darwin pointed out that one of the essential differences between artificial
and natural selection lies in the fact that the former can modify only a few
characters, usually only one at a time, while Nature preserves in the
struggle for existence all the variations of a species, at the same time and in
a purely mechanical way, if they possess selection-value.
Herbert Spencer, though himself an adherent of the theory of selection,
declared in the beginning of the nineties that in his opinion the range of this
principle was greatly over-estimated, if the great changes which have taken
place in so many organisms in the course of ages are to be interpreted as
due to this process of selection alone, since no transformation of any
importance can be evolved by itself; it is always accompanied by a host of
secondary changes. He gives the familiar example of the Giant Stag of the
Irish peat, the enormous antlers of which required not only a much stronger
skull cap, but also greater strength of the sinews, muscles, nerves and bones
of the whole anterior half of the animal, if their mass was not to weigh
down the animal altogether. It is inconceivable, he says, that so many
processes of selection should take place simultaneously, and we are
therefore forced to fall back on the Lamarckian factor of the use and disuse
of functional parts. And how, he asks, could natural selection follow two
opposite directions of evolution in different parts of the body at the same
time, as for instance in the case of the kangaroo, in which the forelegs must
have become shorter, while the hind legs and the tail were becoming longer
and stronger?
Spencer's main object was to substantiate the validity of the Lamarckian
principle, the coöperation of which with selection had been doubted by
many. And it does seem as though this principle, if it operates in nature at

all, offers a ready and simple explanation of all such secondary variations.
Not only muscles, but nerves, bones, sinews, in short all tissues which
function actively, increase in strength in proportion as they are used, and
conversely they decrease when the claims on them diminish. All the parts,
therefore, which depend on the part that varied first, as for instance the
enlarged antlers of the Irish Elk, must have been increased or decreased in
strength, in exact proportion to the claims made upon them,—just as is
actually the case.
But beautiful as this explanation would be, I regard it as untenable, because
it assumes the transmissibility of functional modifications (so-called
"acquired" characters), and this is not only undemonstrable, but is scarcely
theoretically conceivable, for the secondary variations which accompany or
follow the first as correlative variations, occur also in cases in which the
animals concerned are sterile and therefore cannot transmit anything to
their descendants. This is true of worker bees, and particularly of ants, and I
shall here give a brief survey of the present state of the problem as it
appears to me.
Much has been written on both sides of this question since the published
controversy on the subject in the nineties between Herbert Spencer and
myself. I should like to return to the matter in detail, if the space at my
disposal permitted, because it seems to me that the arguments I advanced at
that time are equally cogent to-day, notwithstanding all the objections that
have since been urged against them. Moreover, the matter is by no means
one of subordinate interest; it is the very kernel of the whole question of the
reality and value of the principle of selection. For if selection alone does not
suffice to explain "harmonious adaptation" as I have called Spencer's
Coadaptation, and if we require to call in the aid of the Lamarckian factor it
would be questionable whether selection would explain any adaptations
whatever. In this particular case—of worker bees—the Lamarckian factor
may be excluded altogether, for it can be demonstrated that here at any rate
the effects of use and disuse cannot be transmitted.
But if it be asked why we are unwilling to admit the coöperation of the
Darwinian factor of selection and the Lamarckian factor, since this would
afford us an easy and satisfactory explanation of the phenomena, I answer:
Because the Lamarckian principle is fallacious, and because by accepting it

we close the way towards deeper insight. It is not a spirit of combativeness
or a desire for self-vindication that induces me to take the field once more
against the Lamarckian principle, it is the conviction that the progress of
our knowledge is being obstructed by the acceptance of this fallacious
principle, since the facile explanation it apparently affords prevents our
seeking after a truer explanation and a deeper analysis.
The workers in the various species of ants are sterile, that is to say, they
take no regular part in the reproduction of the species, although individuals
among them may occasionally lay eggs. In addition to this they have lost
the wings, and the receptaculum seminis, and their compound eyes have
degenerated to a few facets. How could this last change have come about
through disuse, since the eyes of workers are exposed to light in the same
way as are those of the sexual insects and thus in this particular case are not
liable to "disuse" at all? The same is true of the receptaculum seminis,
which can only have been disused as far as its glandular portion and its
stalk are concerned, and also of the wings, the nerves tracheae and
epidermal cells of which could not cease to function until the whole wing
had degenerated, for the chitinous skeleton of the wing does not function at
all in the active sense.
But, on the other hand, the workers in all species have undergone
modifications in a positive direction, as, for instance, the greater
development of brain. In many species large workers have evolved,—the
so-called soldiers, with enormous jaws and teeth, which defend the colony,
—and in others there are small workers which have taken over other special
functions, such as the rearing of the young Aphides. This kind of division of
the workers into two castes occurs among several tropical species of ants,
but it is also present in the Italian species, Colobopsis truncata. Beautifully
as the size of the jaws could be explained as due to the increased use made
of them by the "soldiers," or the enlarged brain as due to the mental
activities of the workers, the fact of the infertility of these forms is an
insurmountable obstacle to accepting such an explanation. Neither jaws nor
brain can have been evolved on the Lamarckian principle.
The problem of coadaptation is no easier in the case of the ant than in the
case of the Giant Stag. Darwin himself gave a pretty illustration to show
how imposing the difference between the two kinds of workers in one

species would seem if we translated it into human terms. In regard to the
Driver ants (Anomma) we must picture to ourselves a piece of work, "for
instance the building of a house, being carried on by two kinds of workers,
of which one group was five feet four inches high, the other sixteen feet
high."[39]
Although the ant is a small animal as compared with man or with the Irish
Elk, the "soldier" with its relatively enormous jaws is hardly less heavily
burdened than the Elk with its antlers, and in the ant's case, too, a
strengthening of the skeleton, of the muscles, the nerves of the head, and of
the legs must have taken place parallel with the enlargement of the jaws.
Harmonious adaptation (coadaptation) has here been active in a high
degree, and yet these "soldiers" are sterile! There thus remains nothing for it
but to refer all their adaptations, positive and negative alike, to processes of
selection which have taken place in the rudiments of the workers within the
egg and sperm-cells of their parents. There is no way out of the difficulty
except the one Darwin pointed out. He himself did not find the solution of
the riddle at once. At first he believed that the case of the workers among
social insects presented "the most serious special difficulty" in the way of
his theory of natural selection; and it was only after it had become clear to
him that it was not the sterile insects themselves but their parents that were
selected, according as they produced more or less well adapted workers,
that he was able to refer to this very case of the conditions among ants "in
order to show the power of natural selection."[40] He explains his view by a
simple but interesting illustration. Gardeners have produced, by means of
long continued artificial selection, a variety of Stock, which bears entirely
double, and therefore infertile flowers.[41] Nevertheless the variety
continues to be reproduced from seed, because, in addition to the double
and infertile flowers, the seeds always produce a certain number of single,
fertile blossoms, and these are used to reproduce the double variety. These
single and fertile plants correspond "to the males and females of an ant-
colony, the infertile plants, which are regularly produced in large numbers,
to the neuter workers of the colony."
This illustration is entirely apt, the only difference between the two cases
consisting in the fact that the variation in the flower is not a useful, but a
disadvantageous one, which can only be preserved by artificial selection on
the part of the gardener, while the transformations that have taken place

parallel with the sterility of the ants are useful, since they procure for the
colony an advantage in the struggle for existence, and they are therefore
preserved by natural selection. Even the sterility itself in this case is not
disadvantageous, since the fertility of the true females has at the same time
considerably increased. We may therefore regard the sterile forms of ants,
which have gradually been adapted in several directions to varying
functions, as a certain proof that selection really takes place in the germ-
cells of the fathers and mothers of the workers, and that special complexes
of primordia (ids) are present in the workers and in the males and females,
and these complexes contain the primordia of the individual parts
(determinants). But since all living entities vary, the determinants must also
vary, now in a favourable, now in an unfavourable direction. If a female
produces eggs, which contain favourably varying determinants in the
worker-ids, then these eggs will give rise to workers modified in the
favourable direction, and if this happens with many females, the colony
concerned will contain a better kind of worker than other colonies.
I digress here in order to give an account of the intimate processes, which,
according to my view, take place within the germ-plasm, and which I have
called "germinal selection." These processes are of importance since they
form the roots of variation, which in its turn is the root of natural selection.
I cannot here do more than give a brief outline of the theory in order to
show how the Darwin-Wallace theory of selection has gained support from
it.
With others, I regard the minimal amount of substance which is contained
within the nucleus of the germ-cells, in the form of rods, bands, or granules,
as the germ-substance or germ-plasm, and I call the individual granules ids.
There is always a multiplicity of such ids present in the nucleus, either
occurring individually, or united in the form of rods or bands
(chromosomes). Each id contains the primary constituents of a whole
individual, so that several ids are concerned in the development of a new
individual.
In every being of complex structure thousands of primary constituents must
go to make up a single id; these I call determinants, and I mean by this
name very small individual particles, far below the limits of microscopic
visibility, vital units which feed, grow, and multiply by division. These

determinants control the parts of the developing embryo,—in what manner
need not here concern us. The determinants differ among themselves, those
of a muscle are differently constituted from those of a nerve-cell or a
glandular cell, etc., and every determinant is in its turn made up of minute
vital units, which I call biophores, or the bearers of life. According to my
view, these determinants not only assimilate, like every other living unit,
but they vary in the course of their growth, as every living unit does; they
may vary qualitatively if the elements of which they are composed vary,
they may grow and divide more or less rapidly, and their variations give rise
to corresponding variations of the organ, cell, or cell-group which they
determine. That they are undergoing ceaseless fluctuations in regard to size
and quality seems to me the inevitable consequence of their unequal
nutrition; for although the germ-cell as a whole usually receives sufficient
nutriment, minute fluctuations in the amount carried to different parts
within the germ-plasm cannot fail to occur.
Now, if a determinant, for instance of a sensory cell, receives for a
considerable time more abundant nutriment than before, it will grow more
rapidly—become bigger, and divide more quickly, and, later, when the id
concerned develops into an embryo, this sensory cell will become stronger
than in the parents, possibly even twice as strong. This is an instance of a
hereditary individual variation, arising from the germ.
The nutritive stream which, according to our hypothesis, favours the
determinant N by chance, that is, for reasons unknown to us, may remain
strong for a considerable time, or may decrease again; but even in the latter
case it is conceivable that the ascending movement of the determinant may
continue, because the strengthened determinant now actively nourishes
itself more abundantly,—that is to say, it attracts the nutriment to itself, and
to a certain extent withdraws it from its fellow-determinants. In this way, it
may—as it seems to me—get into permanent upward movement, and attain
a degree of strength from which there is no falling back. Then positive or
negative selection sets in, favouring the variations which are advantageous,
setting aside those which are disadvantageous.
In a similar manner a downward variation of the determinants may take
place, if its progress be started by a diminished flow of nutriment. The
determinants which are weakened by this diminished flow will have less

affinity for attracting nutriment because of their diminished strength, and
they will assimilate more feebly and grow more slowly, unless chance
streams of nutriment help them to recover themselves. But, as will presently
be shown, a change of direction cannot take place at every stage of the
degenerative process. If a certain critical stage of downward progress be
passed, even favourable conditions of food-supply will no longer suffice
permanently to change the direction of the variation. Only two cases are
conceivable; if the determinant corresponds to a useful organ, only its
removal can bring back the germ-plasm to its former level; therefore
personal selection removes the id in question, with its determinants, from
the germ-plasm, by causing the elimination of the individual in the struggle
for existence. But there is another conceivable case; the determinants
concerned may be those of an organ which has become useless, and they
will then continue unobstructed, but with exceeding slowness, along the
downward path, until the organ becomes vestigial, and finally disappears
altogether.
The fluctuations of the determinants hither and thither may thus be
transformed into a lasting ascending or descending movement; and this is
the crucial point of these germinal processes.
This is not a fantastic assumption; we can read it in the fact of the
degeneration of disused parts. Useless organs are the only ones which are
not helped to ascend again by personal selection, and therefore in their
case alone can we form any idea of how the primary constituents behave,
when they are subject solely to intra-germinal forces.
The whole determinant system of an id, as I conceive it, is in a state of
continual fluctuation upwards and downwards. In most cases the
fluctuations will counteract one another, because the passive streams of
nutriment soon change, but in many cases the limit from which a return is
possible will be passed, and then the determinants concerned will continue
to vary in the same direction, till they attain positive or negative selection-
value. At this stage personal selection intervenes and sets aside the variation
if it is disadvantageous, or favours—that is to say, preserves—it if it is
advantageous. Only the determinant of a useless organ is uninfluenced by
personal selection, and, as experience shows, it sinks downwards; that is,
the organ that corresponds to it degenerates very slowly but uninterruptedly

till, after what must obviously be an immense stretch of time, it disappears
from the germ-plasm altogether.
Thus we find in the fact of the degeneration of disused parts the proof that
not all the fluctuations of a determinant return to equilibrium again, but
that, when the movement has attained to a certain strength, it continues in
the same direction. We have entire certainty in regard to this as far as the
downward progress is concerned, and we must assume it also in regard to
ascending variations, as the phenomena of artificial selection certainly
justify us in doing. If the Japanese breeders were able to lengthen the tail-
feathers of the cock to six feet, it can only have been because the
determinants of the tail-feathers in the germ-plasm had already struck out a
path of ascending variation, and this movement was taken advantage of by
the breeder, who continually selected for reproduction the individuals in
which the ascending variation was most marked. For all breeding depends
upon the unconscious selection of germinal variations.
Of course these germinal processes cannot be proved mathematically, since
we cannot actually see the play of forces of the passive fluctuations and
their causes. We cannot say how great these fluctuations are, and how
quickly or slowly, how regularly or irregularly they change. Nor do we
know how far a determinant must be strengthened by the passive flow of
the nutritive stream if it is to be beyond the danger of unfavourable
variations, or how far it must be weakened passively before it loses the
power of recovering itself by its own strength. It is no more possible to
bring forward actual proofs in this case than it was in regard to the
selection-value of the initial stages of an adaptation. But if we consider that
all heritable variations must have their roots in the germ-plasm, and further,
that when personal selection does not intervene, that is to say, in the case of
parts which have become useless, a degeneration of the part, and therefore
also of its determinant must inevitably take place; then we must conclude
that processes such as I have assumed are running their course within the
germ-plasm, and we can do this with as much certainty as we were able to
infer, from the phenomena of adaptation, the selection-value of their initial
stages. The fact of the degeneration of disused parts seems to me to afford
irrefutable proof that the fluctuations within the germ-plasm are the real
root of all hereditary variation, and the preliminary condition for the
occurrence of the Darwin-Wallace factor of selection. Germinal selection

supplies the stones out of which personal selection builds her temples and
palaces: adaptations. The importance for the theory of the process of
degeneration of disused parts cannot be over-estimated, especially when it
occurs in sterile animal forms, where we are free from the doubt as to the
alleged Lamarckian factor which is apt to confuse our ideas in regard to
other cases.
If we regard the variation of the many determinants concerned in the
transformation of the female into the sterile worker as having come about
through the gradual transformation of the ids into worker-ids, we shall see
that the germ-plasm of the sexual ants must contain three kinds of ids, male,
female, and worker ids, or if the workers have diverged into soldiers and
nest-builders, then four kinds. We understand that the worker-ids arose
because their determinants struck out a useful path of variation, whether
upward or downward, and that they continued in this path until the highest
attainable degree of utility of the parts determined was reached. But in
addition to the organs of positive or negative selection-value, there were
some which were indifferent as far as the success and especially the
functional capacity of the workers was concerned: wings, ovarian tubes,
receptaculum seminis, a number of the facets of the eye, perhaps even the
whole eye. As to the ovarian tubes it is is possible that their degeneration
was an advantage for the workers, in saving energy, and if so selection
would favour the degeneration; but how could the presence of eyes
diminish the usefulness of the workers to the colony? or the minute
receptaculum seminis, or even the wings? These parts have therefore
degenerated because they were of no further value to the insect. But if
selection did not influence the setting aside of these parts because they were
neither of advantage nor of disadvantage to the species, then the Darwinian
factor of selection is here confronted with a puzzle which it cannot solve
alone, but which at once becomes clear when germinal selection is added.
For the determinants of organs that have no further value for the organism,
must, as we have already explained, embark on a gradual course of
retrograde development.
In ants the degeneration has gone so far that there are no wing-rudiments
present in any species, as is the case with so many butterflies, flies, and
locusts, but in the larvae the imaginable discs of the wings are still laid
down. With regard to the ovaries, degeneration has reached different levels

in different species of ants, as has been shown by the researches of my
former pupil, Elizabeth Bickford. In many species there are twelve ovarian
tubes, and they decrease from that number to one; indeed, in one species no
ovarian tube at all is present. So much at least is certain from what has been
said, that in this case everything depends on the fluctuations of the elements
of the germ-plasm. Germinal selection, here as elsewhere, presents the
variations of the determinants, and personal selection favours or rejects
these, or,—if it be a question of organs which have become useless,—it
does not come into play at all, and allows the descending variation free
course.
It is obvious that even the problem of coadaptation in sterile animals can
thus be satisfactorily explained. If the determinants are oscillating upwards
and downwards in continual fluctuation, and varying more pronouncedly
now in one direction now in the other, useful variations of every
determinant will continually present themselves anew, and may, in the
course of generations, be combined with one another in various ways. But
there is one character of the determinants that greatly facilitates this
complex process of selection, that, after a certain limit has been reached,
they go on varying in the same direction. From this it follows that
development along a path once struck out may proceed without the
continual intervention of personal selection. This factor only operates, so to
speak, at the beginning, when it selects the determinants which are varying
in the right direction, and again at the end, when it is necessary to put a
check upon further variation. In addition to this, enormously long periods
have been available for all these adaptations, as the very gradual transition
stages between females and workers in many species plainly show, and thus
this process of transformation loses the marvellous and mysterious
character that seemed at the first glance to invest it, and takes rank, without
any straining, among the other processes of selection. It seems to me that,
from the facts that sterile animal forms can adapt themselves to new vital
functions, their superfluous parts degenerate, and the parts more used adapt
themselves in an ascending direction, those less used in a descending
direction, we must draw the conclusion that harmonious adaptation here
comes about without the coöperation of the Lamarckian principle. This
conclusion once established, however, we have no reason to refer the
thousands of cases of harmonious adaptation, which occur in exactly the

same way among other animals or plants, to a principle, the active
intervention of which in the transformation of species is nowhere proved.
We do not require it to explain the facts, and therefore we must not assume
it.
The fact of coadaptation, which was supposed to furnish the strongest
argument against the principle of selection, in reality yields the clearest
evidence in favour of it. We must assume it, because no other possibility of
explanation is open to us, and because these adaptations actually exist, that
is to say, have really taken place. With this conviction I attempted, as far
back as 1894, when the idea of germinal selection had not yet occurred to
me, to make "harmonious adaptation" (coadaptation) more easily
intelligible in some way or other, and so I was led to the idea, which was
subsequently expounded in detail by Baldwin, and Lloyd Morgan, and also
by Osborn, and Gulick as Organic Selection. It seemed to me that it was not
necessary that all the germinal variations required for secondary variations
should have occurred simultaneously, since, for instance, in the case of the
stag, the bones, muscles, sinews, and nerves would be incited by the
increasing heaviness of the antlers to greater activity in the individual life,
and so would be strengthened. The antlers can only have increased in size
by very slow degrees, so that the muscles and bones may have been able to
keep pace with their growth in the individual life, until the requisite
germinal variations presented themselves. In this way a disharmony
between the increasing weight of the antlers and the parts which support
and move them would be avoided, since time would be given for the
appropriate germinal variations to occur, and so to set agoing the hereditary
variation of the muscles, sinews and bones.[42]
I still regard this idea as correct, but I attribute less importance to "organic
selection" than I did at that time, in so far that I do not believe that it alone
could effect complex harmonious adaptations. Germinal selection now
seems to me to play the chief part in bringing about such adaptations.
Something the same is true of the principle I have called Panmixia. As I
became more and more convinced, in the course of years, that the
Lamarckian principle ought not to be called in to explain the dwindling of
disused parts, I believed that this process might be simply explained as due
to the cessation of the conservative effect of natural selection. I said to
myself that, from the moment in which a part ceases to be of use, natural

selection withdraws its hand from it, and then it must inevitably fall from
the height of its adaptiveness, because inferior variants would have as good
a chance of persisting as better ones, since all grades of fitness of the part in
question would be mingled with one another indiscriminately. This is
undoubtedly true, as Romanes pointed out ten years before I did, and this
mingling of the bad with the good probably does bring about a deterioration
of the part concerned. But it cannot account for the steady diminution,
which always occurs when a part is in process of becoming rudimentary,
and which goes on until it ultimately disappears altogether. The process of
dwindling cannot therefore be explained as due to panmixia alone: we can
only find a sufficient explanation in germinal selection.
IV. Derivatives of the Theory of Selection
The impetus in all directions given by Darwin through his theory of
selection has been an immeasurable one, and its influence is still felt. It falls
within the province of the historian of science to enumerate all the ideas
which, in the last quarter of the nineteenth century, grew out of Darwin's
theories, in the endeavour to penetrate more deeply into the problem of the
evolution of the organic world. Within the narrow limits to which this paper
is restricted, I cannot attempt to discuss any of these.
V. Arguments for the Reality of the Processes of Selection
(a) Sexual Selection
Sexual selection goes hand in hand with natural selection. From the very
first I have regarded sexual selection as affording an extremely important
and interesting corroboration of natural selection, but, singularly enough, it
is precisely against this theory that an adverse judgment has been
pronounced in so many quarters, and it is only quite recently, and probably
in proportion as the wealth of facts in proof of it penetrates into a wider
circle, that we seem to be approaching a more general recognition of this
side of the problem of adaptations. Thus Darwin's words in his preface to
the second edition (1874) of his book, The Descent of Man and Sexual
Selection, are being justified: "My conviction as to the operation of natural
selection remains unshaken," and further, "If naturalists were to become
more familiar with the idea of sexual selection, it would, I think, be
accepted to a much greater extent, and already it is fully and favourably

accepted by many competent judges." Darwin was able to speak thus
because he was already acquainted with an immense mass of facts, which,
taken together, yield overwhelming evidence of the validity of the principle
of sexual selection.
Natural selection chooses out for reproduction the individuals that are best
equipped for the struggle for existence, and it does so at every stage of
development; it thus improves the species in all its stages and forms. Sexual
selection operates only on individuals that are already capable of
reproduction, and does so only in relation to the attainment of reproduction.
It arises from the rivalry of one sex, usually the male, for the possession of
the other, usually the female. Its influence can therefore only directly affect
one sex, in that it equips it better for attaining possession of the other. But
the effect may extend indirectly to the female sex, and thus the whole
species may be modified, without, however, becoming any more capable of
resistance in the struggle for existence, for sexual selection only gives rise
to adaptations which are likely to give their possessor the victory over rivals
in the struggle for possession of the female, and which are therefore
peculiar to the wooing sex: the manifold "secondary sexual characters." The
diversity of these characters is so great that I cannot here attempt to give
anything approaching a complete treatment of them, but I should like to
give a sufficient number of examples to make the principle itself, in its
various modes of expression, quite clear.
One of the chief preliminary postulates of sexual selection is the unequal
number of individuals in the two sexes, for if every male immediately finds
his mate there can be no competition for the possession of the female.
Darwin has shown that, for the most part, the inequality between the sexes
is due simply to the fact that there are more males than females, and
therefore the males must take some pains to secure a mate. But the
inequality does not always depend on the numerical preponderance of the
males, it is often due to polygamy; for, if one male claims several females,
the number of females in proportion to the rest of the males will be reduced.
Since it is almost always the males that are the wooers, we must expect to
find the occurrence of secondary sexual characters chiefly among them, and
to find it especially frequent in polygamous species. And this is actually the
case.

If we were to try to guess—without knowing the facts—what means the
male animals make use of to overcome their rivals in the struggle for the
possession of the female, we might name many kinds of means, but it
would be difficult to suggest any which is not actually employed in some
animal group of other. I begin with the mere difference in strength, through
which the male of many animals is so sharply distinguished from the
female, as, for instance, the lion, walrus, "sea-elephant," and others. Among
these the males fight violently for the possession of the female, who falls to
the victor in the combat. In this simple case no one can doubt the operation
of selection, and there is just as little room for doubt as to the selection-
value of the initial stages of the variation. Differences in bodily strength are
apparent even among human beings, although in their case the struggle for
the possession of the female is no longer decided by bodily strength alone.
Combats between male animals are often violent and obstinate, and the
employment of the natural weapons of the species in this way has led to
perfecting of these, e.g. the tusks of the boar, the antlers of the stag, and the
enormous, antler-like jaws of the stag-beetle. Here again it is impossible to
doubt that variations in these organs presented themselves, and that these
were considerable enough to be decisive in combat, and so to lead to the
improvement of the weapon.
Among many animals, however, the females at first withdraw from the
males; they are coy, and have to be sought out, and sometimes held by
force. This tracking and grasping of the females by the males has given rise
to many different characters in the latter, as, for instance, the larger eyes of
the male bee, and especially of the males of the Ephemerids (May-flies),
some species of which show, in addition to the usual compound eyes, large,
so-called turban-eyes, so that the whole head is covered with seeing
surfaces. In these species the females are very greatly in the minority (1-
100), and it is easy to understand that a keen competition for them must
take place, and that, when the insects of both sexes are floating freely in the
air, an unusually wide range of vision will carry with it a decided
advantage. Here again the actual adaptations are in accordance with the
preliminary postulates of the theory. We do not know the stages through
which the eye has passed to its present perfected state, but, since the
number of simple eyes (facets) has become very much greater in the male
than in the female, we may assume that their increase is due to a gradual

duplication of the determinants of the ommatidium in the germ-plasm, as I
have already indicated in regard to sense-organs in general. In this case,
again, the selection-value of the initial stages hardly admits of doubt; better
vision directly secures reproduction.
In many cases the organ of smell shows a similar improvement. Many
lower Crustaceans (Daphnidae) have better developed organs of smell in the
male sex. The difference is often slight and amounts only to one or two
olfactory filaments, but certain species show a difference of nearly a
hundred of these filaments (Leptodora). The same thing occurs among
insects.
We must briefly consider the clasping or grasping organs which have
developed in the males among many lower Crustaceans, but here natural
selection plays its part along with sexual selection, for the union of the
sexes is an indispensable condition for the maintenance of the species, and
as Darwin himself pointed out, in many cases the two forms of selection
merge into each other. This fact has always seemed to me to be a proof of
natural selection, for, in regard to sexual selection, it is quite obvious that
the victory of the best-equipped could have brought about the improvement
only of the organs concerned, the factors in the struggle, such as the eye and
the olfactory organ.
We come now to the excitants; that is, to the group of sexual characters
whose origin through processes of selection has been most frequently called
in question. We may cite the love-calls produced by many male insects,
such as crickets and cicadas. These could only have arisen in animal groups
in which the female did not rapidly flee from the male, but was inclined to
accept his wooing from the first. Thus, notes like the chirping of the male
cricket serve to entice the females. At first they were merely the signal
which showed the presence of a male in the neighbourhood, and the female
was gradually enticed nearer and nearer by the continued chirping. The
male that could make himself heard to the greatest distance would obtain
the largest following, and would transmit the beginnings, and, later, the
improvement of his voice to the greatest number of descendants. But sexual
excitement in the female became associated with the hearing of the love-
call, and then the sound-producing organ of the male began to improve,
until it attained to the emission of the long-drawn-out soft notes of the

mole-cricket or the maenad-like cry of the cicadas. I cannot here follow the
process of development in detail, but will call attention to the fact that the
original purpose of the voice, the announcing of the male's presence,
became subsidiary, and the exciting of the female became the chief goal to
be aimed at. The loudest singers awakened the strongest excitement, and the
improvement resulted as a matter of course. I conceive of the origin of bird-
song in a somewhat similar manner, first as a means of enticing, then of
exciting the female.
One more kind of secondary sexual character must here be mentioned: the
odour which emanates from so many animals at the breeding season. It is
possible that this odour also served at first merely to give notice of the
presence of individuals of the other sex, but it soon became an excitant, and
as the individuals which caused the greatest degree of excitement were
preferred, it reached as high a pitch of perfection as was possible to it. I
shall confine myself here to the comparatively recently discovered
fragrance of butterflies. Since Fritz Müller found out that certain Brazilian
butterflies gave off fragrance "like a flower," we have become acquainted
with many such cases, and we now know that in all lands, not only many
diurnal Lepidoptera but nocturnal ones also give off a delicate odour, which
is agreeable even to man. The ethereal oil to which this fragrance is due is
secreted by the skin-cells, usually of the wing, as I showed soon after the
discovery of the scent-scales. This is the case in the males; the females have
no special scent-scales recognisable as such by their form, but they must,
nevertheless, give off an extremely delicate fragrance, although our
imperfect organ of smell cannot perceive it, for the males become aware of
the presence of a female, even at night, from a long distance off, and gather
round her. We may therefore conclude, that both sexes have long given
forth a very delicate perfume, which announced their presence to others of
the same species, and that in many species (not in all) these small
beginnings become, in the males, particularly strong scent-scales of
characteristic form (lute, brush, or lyre-shaped). At first these scales were
scattered over the surface of the wing, but gradually they concentrated
themselves, and formed broad, velvety bands, or strong, prominent brushes,
and they attained their highest pitch of evolution when they became
enclosed within pits or folds of the skin, which could be opened to let the
delicious fragrance stream forth suddenly towards the female. Thus in this

case also we see that characters, the original use of which was to bring the
sexes together, and so to maintain the species, have been evolved in the
males into means for exciting the female. And we can hardly doubt, that the
females are most readily enticed to yield to the butterfly that sends out the
strongest fragrance,—that is to say, that excites them to the highest degree.
It is a pity that our organs of smell are not fine enough to examine the
fragrance of male Lepidoptera in general, and to compare it with other
perfumes which attract these insects.[43] As far as we can perceive them
they resemble the fragrance of flowers, but there are Lepidoptera whose
scent suggests musk. A smell of musk is also given off by several plants: it
is a sexual excitant in the musk-deer, the musk-sheep, and the crocodile.
As far as we know, then, it is perfumes similar to those of flowers that the
male Lepidoptera give off in order to entice their mates and this is a further
indication that animals, like plants, can to a large extent meet the claims
made upon them by life, and produce the adaptations which are most
purposive,—a further proof, too, of my proposition that the useful
variations, so to speak, are always there. The flowers developed the
perfumes which entice their visitors, and the male Lepidoptera developed
the perfumes which entice and excite their mates.
There are many pretty little problems to be solved in this connection, for
there are insects, such as some flies, that are attracted by smells which are
unpleasant to us, like those from decaying flesh and carrion. But there are
also certain flowers, some orchids for instance, which give forth no very
agreeable odour, but one which is to us repulsive and disgusting; and we
should therefore expect that the males of such insects would give off a
smell unpleasant to us, but there is no case known to me in which this has
been demonstrated.
In cases such as we have discussed, it is obvious that there is no possible
explanation except through selection. This brings us to the last kind of
secondary sexual characters, and the one in regard to which doubt has been
most frequently expressed,—decorative colours and decorative forms, the
brilliant plumage of the male pheasant, the humming-birds, and the bird of
Paradise, as well as the bright colours of many species of butterfly, from the
beautiful blue of our little Lycaenidae to the magnificent azure of the large
Morphinae of Brazil. In a great many cases, though not by any means in all,

the male butterflies are "more beautiful" than the females, and in the
Tropics in particular they shine and glow in the most superb colours. I
really see no reason why we should doubt the power of sexual selection,
and I myself stand wholly on Darwin's side. Even though we certainly
cannot assume that the females exercise a conscious choice of the
"handsomest" mate, and deliberate like the judges in a court of justice over
the perfections of their wooers, we have no reason to doubt that distinctive
forms (decorative feathers), and colours have a particularly exciting effect
upon the female, just as certain odours have among animals of so many
different groups, including the butterflies. The doubts which existed for a
considerable time, as a result of fallacious experiments, as to whether the
colours of flowers really had any influence in attracting butterflies have
now been set at rest through a series of more careful investigations; we now
know that the colours of flowers are there on account of the butterflies, as
Sprengel first showed, and that the blossoms of Phanerogams are selected
in relation to them, as Darwin pointed out.
Certainly it is not possible to bring forward any convincing proof of the
origin of decorative colours through sexual selection, but there are many
weighty arguments in favour of it, and these form a body of presumptive
evidence so strong that it almost amounts to certainty.
In the first place, there is the analogy with other secondary sexual
characters. If the song of birds and the chirping of the cricket have been
evolved through sexual selection, if the penetrating odours of male animals,
—the crocodile, the musk-deer, the beaver, the carnivores, and, finally, the
flower-like fragrances of the butterflies have been evolved to their present
pitch in this way, why should decorative colours have arisen in some other
way? Why should the eye be less sensitive to specifically male colours and
other visible signs enticing to the female, than the olfactory sense to
specifically male odours, or the sense of hearing to specifically male
sounds? Moreover, the decorative feathers of birds are almost always spread
out and displayed before the female during courtship. I have elsewhere[44]
pointed out that decorative colouring and sweet-scentedness may replace
one another in Lepidoptera as well as in flowers, for just as some modestly
coloured flowers (mignonette and violet) have often a strong perfume,
while strikingly coloured ones are sometimes quite devoid of fragrance, so
we find that the most beautiful and gaily-coloured of our native

Lepidoptera, the species of Vanessa, have no scent-scales, while these are
often markedly developed in grey nocturnal Lepidoptera. Both attractions
may, however, be combined in butterflies, just as in flowers. Of course, we
cannot explain why both means of attraction should exist in one genus, and
only one of them in another, since we do not know the minutest details of
the conditions of life of the genera concerned. But from the sporadic
distribution of scent-scales in Lepidoptera, and from their occurrence or
absence in nearly related species, we may conclude that fragrance is a
relatively modern acquirement, more recent than brilliant colouring.
One thing in particular that stamps decorative colouring as a product of
selection is its gradual intensification by the addition of new spots, which
we can quite well observe, because in many cases the colours have been
first acquired by the males, and later transmitted to the females by
inheritance. The scent-scales are never thus transmitted, probably for the
same reason that the decorative colours of many birds are often not
transmitted to the females: because with these they would be exposed to too
great elimination by enemies. Wallace was the first to point out that in
species with concealed nests the beautiful feathers of the male occurred in
the female also, as in the parrots, for instance, but this is not the case in
species which brood on an exposed nest. In the parrots one can often
observe that the general brilliant colouring of the male is found in the
female, but that certain spots of colour are absent, and these have probably
been acquired comparatively recently by the male and have not yet been
transmitted to the female.
Isolation of the group of individuals which is in process of varying is
undoubtedly of great value in sexual selection, for even a solitary
conspicuous variation will become dominant much sooner in a small
isolated colony, than among a large number of members of a species.
Any one who agrees with me in deriving variations from germinal selection
will regard that process as an essential aid towards explaining the selection
of distinctive courtship-characters, such as coloured spots, decorative
feathers, horny outgrowths in birds and reptiles, combs, feather-tufts, and
the like, since the beginnings of these would be presented with relative
frequency in the struggle between the determinants within the germ-plasm.
The process of transmission of decorative feathers to the female results, as

Darwin pointed out and illustrated by interesting examples, in the colour-
transformation of a whole species, and this process, as the phyletically older
colouring of young birds shows, must, in the course of thousands of years,
have repeated itself several times in a line of descent.
If we survey the wealth of phenomena presented to us by secondary sexual
characters, we can hardly fail to be convinced of the truth of the principle of
sexual selection. And certainly no one who has accepted natural selection
should reject sexual selection, for, not only do the two processes rest upon
the same basis, but they merge into one another, so that it is often
impossible to say how much of a particular character depends on one and
how much on the other form of selection.
(b) Natural Selection
An actual proof of the theory of sexual selection is out of the question, if
only because we cannot tell when a variation attains to selection-value. It is
certain that a delicate sense of smell is of value to the male moth in his
search for the female, but whether the possession of one additional
olfactory hair, or of ten, or of twenty additional hairs leads to the success of
its possessor we are unable to tell. And we are groping even more in the
dark when we discuss the excitement caused in the female by agreeable
perfumes, or by striking and beautiful colours. That these do make an
impression is beyond doubt; but we can only assume that slight
intensifications of them give any advantage, and we must assume this since
otherwise secondary sexual characters remain inexplicable.
The same thing is true in regard to natural selection. It is not possible to
bring forward any actual proof of the selection-value of the initial stages,
and the stages in the increase of variations, as has been already shown. But
the selection-value of a finished adaptation can in many cases be
statistically determined. Cesnola and Poulton have made valuable
experiments in this direction. The former attached forty-five individuals of
the green, and sixty-five of the brown variety of the praying mantis (Mantis
religiosa), by a silk thread to plants, and watched them for seven days. The
insects which were on a surface of a colour Similar to their own remained
uneaten, while twenty-five green insects on brown parts of plants had all
disappeared in eleven days.

The experiments of Poulton and Sanders[45] were made with 600 pupae of
Vanessa urticae, the "tortoise-shell butterfly." The pupae were artificially
attached to nettles, tree-trunks, fences, walls, and to the ground, some at
Oxford, some at St. Helens in the Isle of Wight. In the course of a month
93% of the pupae at Oxford were killed, chiefly by small birds, while at St.
Helens 68% perished. The experiments showed very clearly that the colour
and character of the surface on which the pupa rests—and thus its own
conspicuousness—are of the greatest importance. At Oxford only the four
pupae which were fastened to nettles emerged; all the rest—on bark, stones
and the like—perished. At St. Helens the elimination was as follows: on
fences where the pupae were conspicuous, 92%; on bark, 66%; on walls,
54%; and among nettles, 57%. These interesting experiments confirm our
views as to protective coloration, and show further, that the ratio of
elimination in the species is a very high one, and that therefore selection
must be very keen.
We may say that the process of selection follows as a logical necessity from
the fulfilment of the three preliminary postulates of the theory: variability,
heredity, and the struggle for existence, with its enormous ratio of
elimination in all species. To this we must add a fourth factor, the
intensification of variations which Darwin established as a fact, and which
we are now able to account for theoretically on the basis of germinal
selection. It may be objected that there is considerable uncertainty about
this logical proof, because of our inability to demonstrate the selection-
value of the initial stages and the individual stages of increase. We have
therefore to fall back on presumptive evidence. This is to be found in the
interpretative value of the theory. Let us consider this point in greater detail.
In the first place it is necessary to emphasize what is often overlooked,
namely, that the theory not only explains the transformations of species, it
also explains their remaining the same; in addition to the principle of
varying, it contains within itself that of persisting. It is part of the essence
of selection, that it not only causes a part to vary till it has reached its
highest pitch of adaptation, but that it maintains it at this pitch. This
conserving influence of natural selection is of great importance, and was
early recognised by Darwin; it follows naturally from the principle of the
survival of the fittest.

We understand from this how it is that a species which has become fully
adapted to certain conditions of life ceases to vary, but remains "constant,"
as long as the conditions of life for it remain unchanged, whether this be for
thousands of years, or for whole geological epochs. But the most
convincing proof of the power of the principle of selection lies in the
innumerable multitude of phenomena which cannot be explained in any
other way. To this category belong all structures which are only passively of
advantage to the organism, because none of these can have arisen by the
alleged Lamarckian principle. These have been so often discussed that we
need do no more than indicate them here. Until quite recently the
sympathetic coloration of animals—for instance, the whiteness of Arctic
animals—was referred, at least in part, to the direct influence of external
factors, but the facts can best be explained by referring them to the
processes of selection, for then it is unnecessary to make the gratuitous
assumption that many species are sensitive to the stimulus of cold and that
others are not. The great majority of Arctic land-animals, mammals and
birds, are white, and this proves that they were all able to present the
variation which was most useful for them. The sable is brown, but it lives in
trees, where the brown colouring protects and conceals it more effectively.
The musk-sheep (Ovibos moschatus) is also brown, and contrasts sharply
with the ice and snow, but it is protected from beasts of prey by its
gregarious habit, and therefore it is of advantage to be visible from as great
a distance as possible. That so many species have been able to give rise to
white varieties does not depend on a special sensitiveness of the skin to the
influence of cold, but to the fact that Mammals and Birds have a general
tendency to vary towards white. Even with us, many birds—starlings,
blackbirds, swallows, etc.—occasionally produce white individuals, but the
white variety does not persist, because it readily falls a victim to the
carnivores. This is true of white fawns, foxes, deer, etc. The whiteness,
therefore, arises from internal causes, and only persists when it is useful. A
great many animals living in a green environment have become clothed in
green, especially insects, caterpillars, and Mantidae, both persecuted and
persecutors.
That it is not the direct effect of the environment which calls forth the green
colour is shown by the many kinds of caterpillar which rest on leaves and
feed on them, but are nevertheless brown. These feed by night and betake

themselves through the day to the trunk of the tree, and hide in the furrows
of the bark. We cannot, however, conclude from this that they were unable
to vary towards green, for there are Arctic animals which are white only in
winter and brown in summer (Alpine hare, and the ptarmigan of the Alps),
and there are also green leaf-insects which remain green only while they are
young and difficult to see on the leaf, but which become brown again in the
last stage of larval life, when they have outgrown the leaf. They then
conceal themselves by day, sometimes only among withered leaves on the
ground, sometimes in the earth itself. It is interesting that in one genus,
Chaerocampa, one species is brown in the last stage of larval life, another
becomes brown earlier, and in many species the last stage is not wholly
brown, a part remaining green. Whether this is a case of a double
adaptation, or whether the green is being gradually crowded out by the
brown, the fact remains that the same species, even the same individual, can
exhibit both variations. The case is the same with many of the leaf-like
Orthoptera, as, for instance, the praying mantis (Mantis religiosa) which we
have already mentioned.
But the best proofs are furnished by those of ten-cited cases in which the
insect bears a deceptive resemblance to another object. We now know many
such cases, such as the numerous imitations of green or withered leaves,
which are brought about in the most diverse ways, sometimes by mere
variations in the form of the insect and in its colour, sometimes by an
elaborate marking, like that which occurs in the Indian leaf-butterflies,
Kallima inachis. In the single butterfly-genus Anaea, in the woods of South
America, there are about a hundred species which are all gaily coloured on
the upper surface, and on the reverse side exhibit the most delicate imitation
of the colouring and pattern of a leaf, generally without any indication of
the leaf-ribs, but extremely deceptive nevertheless. Anyone who has seen
only one such butterfly may doubt whether many of the insignificant details
of the marking can really be of advantage to the insect. Such details are for
instance the apparent holes and splits in the apparently dry or half-rotten
leaf, which are usually due to the fact that the scales are absent on a circular
or oval patch so that the colourless wing-membrane lies bare, and one can
look through the spot as through a window. Whether the bird which is
seeking or pursuing the butterflies takes these holes for dewdrops, or for the
work of a devouring insect, does not affect the question; the mirror-like spot

undoubtedly increases the general deceptiveness, for the same thing occurs
in many leaf-butterflies, though not in all, and in some cases it is replaced
in quite a peculiar manner. In one species of Anaea (A. divina), the resting
butterfly looks exactly like a leaf out of the outer edge of which a large
semi-circular piece has been eaten, possibly by a caterpillar; but if we look
more closely it is obvious that there is no part of the wing absent, and that
the semi-circular piece is of a clear, pale yellow colour, while the rest of the
wing is of a strongly contrasted dark brown.
But the deceptive resemblance may be caused in quite a different manner. I
have often speculated as to what advantage the brilliant white C could give
to the otherwise dusky-coloured "Comma butterfly" (Grapta C. album).
Poulton's recent observations[46] have shown that this represents the
imitation of a crack such as is often seen in dry leaves, and is very
conspicuous because the light shines through it.
The utility obviously lies in presenting to the bird the very familiar picture
of a broken leaf with a clear shining slit, and we may conclude, from the
imitation of such small details, that the birds are very sharp observers and
that the smallest deviation from the usual arrests their attention and incites
them to closer investigation. It is obvious that such detailed—we might
almost say such subtle—deceptive resemblances could only have come
about in the course of long ages through the acquirement from time to time
of something new which heightened the already existing resemblance.
In face of facts like these there can be no question of chance and no one has
succeeded so far in finding any other explanation to replace that by
selection. For the rest, the apparent leaves are by no means perfect copies of
a leaf; many of them only represent the torn or broken piece, or the half or
two-thirds of a leaf, but then the leaves themselves frequently do not
present themselves to the eye as a whole, but partially concealed among
other leaves. Even those butterflies which, like the species of Kallima and
Anaea, represent the whole of a leaf with stalk, ribs, apex, and the whole
breadth, are not actual copies which would satisfy a botanist; there is often
much wanting. In Kallima the lateral ribs of the leaf are never all included
in the markings; there are only two or three on the left side and at more four
or five on the right, and in many individuals these are rather obscure, while
in others they are comparatively distinct. This furnishes us with fresh

evidence in favour of their origin through processes of selection, for a
botanically perfect picture could not arise in this way; there could only be a
fixing of such details as heightened the deceptive resemblance.
Our postulate of origin through selection also enables us to understand why
the leaf-imitation is on the lower surface of the wing in the diurnal
Lepidoptera, and on the upper surface in the nocturnal forms, corresponding
to the attitude of the wings in the resting position of the two groups.
The strongest of all proofs of the theory, however, is afforded by cases of
true "mimicry," those adaptations discovered by Bates in 1861, consisting in
the imitation of one species by another, which becomes more and more like
its model. The model is always a species that enjoys some special
protection from enemies, whether because it is unpleasant to taste, or
because it is in some way dangerous.
It is chiefly among insects and especially among butterflies that we find the
greatest number of such cases. Several of these have been minutely studied
and every detail has been investigated so that it is difficult to understand
how there can still be disbelief in regard to them. If the many and exact
observations which have been carefully collected and critically discussed
for instance by Poulton[47] were thoroughly studied the arguments which
are still frequently urged against mimicry would be found untenable; we
can hardly hope to find more convincing proof of the actuality of the
processes of selection than these cases put into our hands. The preliminary
postulates of the theory of mimicry have been disputed, for instance, that
diurnal butterflies are persecuted and eaten by birds, but observations
specially directed towards this point in India, Africa, America and Europe
have placed it beyond all doubt. If it were necessary I could myself furnish
an account of my own observations on this point.
In the same way it has been established by experiment and observation in
the field that in all the great regions of distribution there are butterflies
which are rejected by birds and lizards, their chief enemies, on account of
their unpleasant smell or taste. These butterflies are usually gaily and
conspicuously coloured and thus—as Wallace first interpreted it—are
furnished with an easily recognisable sign: a sign of unpalatableness or
warning colours. If they were not thus recognisable easily and from a

distance, they would frequently be pecked at by birds, and then rejected
because of their unpleasant taste; but as it is, the insect-eaters recognise
them at once as unpalatable booty and ignore them. Such immune[48]
species, wherever they occur, are imitated by other palatable species, which
thus acquire a certain degree of protection.
It is true that this explanation of the bright, conspicuous colours is only a
hypothesis, but its foundations—unpalatableness, and the liability of other
butterflies to be eaten,—are certain, and its consequences—the existence of
mimetic palatable forms—conform it in the most convincing manner. Of the
many cases now known I select one, which is especially remarkable, and
which has been thoroughly investigated, Papilla dardanus (merope), a
large, beautiful, diurnal butterfly which ranges from Abyssinia throughout
the whole of Africa to the south coast of Cape Colony.
The males of this form are everywhere almost the same in colour and in
form of wings, save for a few variations in the sparse black markings on the
pale yellow ground. But the females occur in several quite different forms
and colourings, and one of these only, the Abyssinian form, is like the male,
while the other three or four are mimetic, that is to say, they copy a butterfly
of quite a different family the Danaids, which are among the immune forms.
In each region the females have thus copied two or three different immune
species. There is much that is interesting to be said in regard to these
species, but it would be out of keeping with the general tenor of this paper
to give details of this very complicated case of polymorphism in P.
Dardanus. Anyone who is interested in the matter will find a full and exact
statement of the case in as far as we know it, in Poulton's Essays on
Evolution (pp. 373-375[49]). I need only add that three different mimetic
female forms have been reared from the eggs of a single female in South
Africa. The resemblance of the forms to their immune models goes so far
that even the details of the local forms of the models are copied by the
mimetic species.
It remains to be said that in Madagascar a butterfly,
Papilio meriones, occurs, of which both sexes are very similar in form and
markings to the non-mimetic male of P. dardanus, so that it probably
represents the ancestor of this latter species.

In face of such facts as these every attempt at another explanation must fail.
Similarly all the other details of the case fulfil the preliminary postulates of
selection, and leave no room for any other interpretation. That the males do
not take on the protective colouring is easily explained, because they are in
general more numerous, and the females are more important for the
preservation of the species, and must also live longer in order to deposit
their eggs. We find the same state of things in many other species, and in
one case (Elymnias undularis) in which the male is also mimetically
coloured, it copies quite a differently coloured immune species from the
model followed by the female. This is quite intelligible when we consider
that if there were too many false immune types, the birds would soon
discover that there were palatable individuals among those with unpalatable
warning colours. Hence the imitation of different immune species by
Papilio dardanus!
I regret that lack of space prevents my bringing forward more examples of
mimicry and discussing them fully. But from the case of Papilio dardanus
alone there is much to be learnt which is of the highest importance for our
understanding of transformations. It shows us chiefly what I once called,
somewhat strongly perhaps, the omnipotence of natural selection in answer
to an opponent who had spoken of its "inadequacy." We here see that one
and the same species is capable of producing four or five different patterns
of colouring and marking; thus the colouring and marking are not, as has
often been supposed, a necessary outcome of the specific nature of the
species, but a true adaptation, which cannot arise as a direct effect of
climatic conditions, but solely through what I may call the sorting out of the
variations produced by the species, according to their utility. That
caterpillars may be either green or brown is already something more than
could have been expected according to the old conception of species, but
that one and the same butterfly should be now pale yellow, with black; now
red with black and pure white; now deep black with large, pure white spots;
and again black with a large ocheous-yellow spot, and many small white
and yellow spots; that in one sub-species it may be tailed like the ancestral
form, and in another tailless like its Danaid model,—all this shows a far-
reaching capacity for variation and adaptation that we could never have
expected if we did not see the facts before us. How it is possible that the
primary colour-variations should thus be intensified and combined remains

a puzzle even now; we are reminded of the modern three-colour printing,—
perhaps similar combinations of the primary colours take place in this case;
in any case the direction of these primary variations is determined by the
artist whom we know as natural selection, for there is no other conceivable
way in which the model could affect the butterfly that is becoming more
and more like it. The same climate surrounds all four forms of female; they
are subject to the same conditions of nutrition. Moreover, Papilio dardanus
is by no means the only species of butterfly which exhibits different kinds
of colour-pattern on its wings. Many species of the Asiatic genus Elymnias
have on the upper surface a very good imitation of an immune Euploeine
(Danainae), often with a steel-blue ground-colour, while the under surface
is well concealed when the butterfly is at rest,—thus there are two kinds of
protective coloration each with a different meaning! The same thing may be
observed in many non-mimetic butterflies, for instance in all our species of
Vanessa, in which the under side shows a grey-brown or brownish-black
protective coloration, but we do not yet know with certainty what may be
the biological significance of the gaily coloured upper surface.
In general it may be said that mimetic butterflies are comparatively rare
species, but there are exceptions, for instance Limenitis archippus in North
America, of which the immune model (Danaida plexippus) also occurs in
enormous numbers.
In another mimicry-category the imitators are often more numerous than the
models, namely in the case of the imitation of dangerous insects by
harmless species. Bees and wasps are dreaded for their sting, and they are
copied by harmless flies of the genera Eristalis and Syrphus, and these
mimics often occur in swarms about flowering plants without damage to
themselves or to their models; they are feared and are therefore left
unmolested.
In regard also to the faithfulness of the copy the facts are quite in harmony
with the theory, according to which the resemblance must have arisen and
increased by degrees. We can recognise this in many cases, for even now
the mimetic species show very varying degrees of resemblance to their
immune model. If we compare, for instance, the many different imitators of
Danaida chrysippus we find that, with their brownish-yellow ground-
colour, and the position and size, and more or less sharp limitation of their

clear marginal spots, they have reached very different degrees of nearness
to their model. Or compare the female of Elymnias undularis with its model
Danaida genutia; there is a general resemblance, but the marking of the
Danaida is very roughly imitated in Elymnias.
Another fact that bears out the theory of mimicry is, that even when the
resemblance in colour-pattern is very great, the wing-venation, which is so
constant, and so important in determining the systematic position of
butterflies, is never affected by the variation. The pursuers of the butterfly
have no time to trouble about entomological intricacies.
I must not pass over a discovery of Poulton's which is of great theoretical
importance—that mimetic butterflies may reach the same effect by very
different means.[50] Thus the glass-like transparency of the wing of a certain
Ithomiine (Methona) and its Pierine mimic (Dismorphia orise) depends on
a diminution in the size of the scales; in the Danaine genus Itune it is due to
the fewness of the scales and in a third imitator, a moth (Castnia linus var.
heliconoides) the glass-like appearance of the wing is due neither to
diminution nor to absence of scales, but to their absolute colourlessness and
transparency, and to the fact that they stand upright. In another moth mimic
(Anthomyza) the arrangement of the transparent scales is normal. Thus it is
not some unknown external influence that has brought about the
transparency of the wing in these five forms, as has sometimes been
supposed. Nor is it a hypothetical internal evolutionary tendency, for all
three vary in a different manner. The cause of this agreement can only lie in
selection, which preserves and intensifies in each species the favourable
variations that present themselves. The great faithfulness of the copy is
astonishing in these cases, for it is not the whole wing which is transparent;
certain markings are black in colour, and these contrast sharply with the
glass-like ground. It is obvious that the pursuers of these butterflies must be
very sharp-sighted, for otherwise the agreement between the species could
never have been pushed so far. The less the enemies see and observe, the
more defective must the imitation be, and if they had been blind, no visible
resemblance between the species which required protection could ever have
arisen.
A seemingly irreconcilable contradiction to the mimicry theory is presented
in the following cases, which were known to Bates, who, however, never

succeeded in bringing them into line with the principle of mimicry.
In South America there are, as we have already said, many mimics of the
immune Ithomiinae (or as Bates called them Heliconidae). Among these
there occur not merely species which are edible, and thus require the
protection of a disguise, but others which are rejected on account of their
unpalatableness. How could the Ithomiine dress have developed in their
case, and of what use is it, since the species would in any case be immune?
In Eastern Brazil, for instance, there are four butterflies, which bear a most
confusing resemblance to one another in colour, marking, and form of wing,
and all four are unpalatable to birds. They belong to four different genera
and three sub-families, and we have to inquire: Whence came this
resemblance and what end does it serve? For a long time no satisfactory
answer could be found, but Fritz Müller,[51] seventeen years after Bates,
offered a solution to the riddle, when he pointed out that young birds could
not have an instinctive knowledge of the unpalatableness of the Ithomiines,
but must learn by experience which species were edible and which inedible.
Thus each young bird must have tasted at least one individual of each
inedible species and discovered its unpalatability, before it learnt to avoid,
and thus to spare the species. But if the four species resemble each other
very closely the bird will regard them all as of the same kind, and avoid
them all. Thus there developed a process of selection which resulted in the
survival of the Ithomiine-like individuals, and in so great an increase of
resemblance between the four species, that they are difficult to distinguish
one from another even in a collection. The advantage for the four species,
living side by side as they do e.g. in Bahia, lies in the fact that only one
individual from the mimicry-ring ("inedible association") need be tasted by
a young bird, instead of at least four individuals, as would otherwise be the
case. As the number of young birds is great, this makes a considerable
difference in the ratio of elimination. The four Brazilian species are Lycorea
halia (Danainae), Heliconius narcaea (eucrate) (Heliconinae), Melinaea
ethra, and Mechanitis lysimnia (Ithomiinae).
These interesting mimicry-rings (trusts), which have much significance for
the theory, have been the subject of numerous and careful investigations,
and at least their essential features are now fully established. Müller took
for granted, without making any investigations, that young birds only learn
by experience to distinguish between different kinds of victims. But Lloyd

Morgan's[52] experiments with young birds proved that this is really the
case, and at the same time furnished an additional argument against the
Lamarckian principle.
In addition to the mimicry-rings first observed in South America, others
have been described from Tropical India by Moore, and by Poulton and
Dixey from Africa, and we may expect to learn many more interesting facts
in this connection. Here again the preliminary postulates of the theory are
satisfied. And how much more that would lead to the same conclusion
might be added!
As in the case of mimicry many species have come to resemble one another
through processes of selection, so we know whole classes of phenomena in
which plants and animals have become adapted to one another, and have
thus been modified to a considerable degree. I refer particularly to the
relation between flowers and insects. Darwin has shown that the originally
inconspicuous blossoms of the phanerogams were transformed into flowers
through the visits of insects, and that, conversely, several large orders of
insects have been gradually modified by their association with flowers,
especially as regards the parts of their body actively concerned. Bees and
butterflies in particular have become what they are through their relation to
flowers. In this case again all that is apparently contradictory to the theory
can, on closer investigation, be beautifully interpreted in corroboration of it.
Selection can give rise only to what is of use to the organism actually
concerned, never to what is of use to some other organism, and we must
therefore expect to find that in flowers only characters of use to themselves
have arisen, never characters which are of use to insects only, and
conversely that in the insects characters useful to them and not merely to
the plants would have originated. For a long time it seemed as if an
exception to this rule existed in the case of the fertilisation of the yucca
blossoms by a little moth, Pronuba yuccasella. This little moth has a sickle-
shaped appendage to its mouth-parts which occurs hi no other
Lepidopteron, and which is used for pushing the yellow pollen into the
opening of the pistil, thus fertilising the flower. Thus it appears as if a new
structure, which is useful only to the plant, has arisen in the insect. But the
difficulty is solved as soon as we learn that the moth lays its eggs in the
fruit-buds of the Yucca, and that the larvae, when they emerge, feed on the
developing seeds. In effecting the fertilisation of the flower the moth is at

the same time making provision for its own offspring, since it is only after
fertilisation that the seeds begin to develop. There is thus nothing to prevent
our referring this structural adaptation in Pronuba yuccasella to processes
of selection, which have gradually transformed the maxillary palps of the
female into the sickle-shaped instrument for collecting the pollen, and
which have at the same time developed in the insect the instinct to press the
pollen into the pistil.
In this domain, then, the theory of selection finds nothing but corroboration,
and it would be impossible to substitute for it any other explanation, which
now that the facts are so well known, could be regarded as a serious rival to
it. That selection is a factor, and a very powerful factor in the evolution of
organisms, can no longer be doubted. Even although we cannot bring
forward formal proofs of it in detail, cannot calculate definitely the size of
the variations which present themselves, and their selection-value, cannot,
in short, reduce the whole process to a mathematical formula, yet we must
assume selection, because it is the only possible explanation applicable to
whole classes of phenomena, and because, on the other hand, it is made up
of factors which we know can be proved actually to exist, and which, if they
exist, must of logical necessity coöperate in the manner required by the
theory. We must accept it because the phenomena of evolution and
adaptation must have a natural basis, and because it is the only possible
explanation of them.[53]
Many people are willing to admit that selection explains adaptations, but
they maintain that only a part of the phenomena are thus explained, because
everything does not depend upon adaptation. They regard adaptation as, so
to speak, a special effort on the part of Nature, which she keeps in readiness
to meet particularly difficult claims of the external world on organisms. But
if we look at the matter more carefully we shall find that adaptations are by
no means exceptional, but that they are present everywhere in such
enormous numbers, that it would be difficult in regard to any structure
whatever, to prove that adaptation had not played a part in its evolution.
How often has the senseless objection been urged against selection that it
can create nothing, it can only reject. It is true that it cannot create either the
living substance or the variations of it; both must be given. But in rejecting
one thing it preserves another, intensifies it, combines it, and in this way

creates what is new. Everything in organisms depends on adaptation; that is
to say, everything must be admitted through the narrow door of selection,
otherwise it can take no part in the building up of the whole. But, it is
asked, what of the direct effect of external conditions, temperature,
nutrition, climate and the like? Undoubtedly these can give rise to
variations, but they too must pass through the door of selection, and if they
cannot do this they are rejected, eliminated from the constitution of the
species.
It may, perhaps, be objected that such external influences are often of a
compelling power, and that every animal must submit to them, and that thus
selection has no choice and can neither select nor reject. There may be such
cases; let us assume for instance that the effect of the cold of the Arctic
regions was to make all the mammals become black; the result would be
that they would all be eliminated by selection, and that no mammals would
be able to live there at all. But in most cases a certain percentage of animals
resists these strong influences, and thus selection secures a foothold on
which to work, eliminating the unfavourable variation, and establishing a
useful colouring, consistent with what is required for the maintenance of the
species.
Everything depends upon adaptation! We have spoken much of adaptation
in colouring, in connection with the examples brought into prominence by
Darwin, because these are conspicuous, easily verified, and at the same
time convincing for the theory of selection. But is it only desert and polar
animals whose colouring is determined through adaptation? Or the leaf-
butterflies, and the mimetic species, or the terrifying markings, and
"warning-colours" and a thousand other kinds of sympathetic colouring? It
is, indeed, never the colouring alone which makes up the adaptation; the
structure of the animal plays a part, often a very essential part, in the
protective disguise, and thus many variations may cooperate towards one
common end. And it is to be noted that it is by no means only external parts
that are changed; internal parts are always modified at the same time—for
instance, the delicate elements of the nervous system on which depend the
instinct of the insect to hold its wings, when at rest, in a perfectly definite
position, which, in the leaf-butterfly, has the effect of bringing the two
pieces on which the marking occurs on the anterior and posterior wing into
the same direction, and thus displaying as a whole the fine curve of the

midrib on the seeming leaf. But the wing-holding instinct is not regulated in
the same way in all leaf-butterflies; even our indigenous species of Vanessa,
with their protective ground-colouring, have quite a distinctive way of
holding their wings so that the greater part of the anterior wing is covered
by the posterior when the butterfly is at rest. But the protective colouring
appears on the posterior wing and on the tip of the anterior, to precisely the
distance to which it is left uncovered. This occurs, as Standfuss has shown,
in different degrees in our two most nearly allied species, the uncovered
portion being smaller in V. urticae than in V. polychloros. In this case, as in
most leaf-butterflies, the holding of the wing was probably the primary
character; only after that was thoroughly established did the protective
marking develop. In any case, the instinctive manner of holding the wings
is associated with the protective colouring, and must remain as it is if the
latter is to be effective. How greatly instincts may change, that is to say,
may be adapted, is shown by the case of the Noctuid "shark" moth, Xylina
vetusta. This form bears a most deceptive resemblance to a piece of rotten
wood, and the appearance is greatly increased by the modification of the
innate impulse to flight common to so many animals, which has here been
transformed into an almost contrary instinct. This moth does not fly away
from danger, but "feigns death," that is, it draws antennae, legs and wings
close to the body, and remains perfectly motionless. It may be touched,
picked up, and thrown down again, and still it does not move. This
remarkable instinct must surely have developed simultaneously with the
wood-colouring; at all events, both coöperating variations are now present,
and prove that both the external and the most minute internal structure have
undergone a process of adaptation.
The case is the same with all structural variations of animal parts, which are
not absolutely insignificant. When the insects acquired wings they must
also have acquired the mechanism with which to move them—the
musculature, and the nervous apparatus necessary for its automatic
regulation. All instincts depend upon compound reflex mechanisms and are
just as indispensable as the parts they have to set in motion, and all may
have arisen through processes of selection if the reasons which I have
elsewhere given for this view are correct.[54]
Thus there is no lack of adaptations within the organism, and particularly in
its most important and complicated parts, so that we may say that there is

no actively functional organ that has not undergone a process of adaptation
relative to its function and the requirements of the organism. Not only is
every gland structurally adapted, down to the very minutest histological
details, to its function, but the function is equally minutely adapted to the
needs of the body. Every cell in the mucous lining of the intestine is exactly
regulated in its relation to the different nutritive substances, and behaves in
quite a different way towards the fats, and towards nitrogenous substances,
or peptones.
I have elsewhere called attention to the many adaptations of the whale to
the surrounding medium, and have pointed out—what has long been
known, but is not universally admitted, even now—that in it a great number
of important organs have been transformed in adaptation to the peculiar
conditions of aquatic life, although the ancestors of the whale must have
lived, like other hair-covered mammals, on land. I cited a number of these
transformations—the fish-like form of the body, the hairlessness of the skin,
the transformation of the fore-limbs to fins, the disappearance of the hind-
limbs and the development of a tail fin, the layer of blubber under the skin,
which affords the protection from cold necessary to a warm-blooded
animal, the disappearance of the ear-muscles and the auditory passages, the
displacement of the external nares to the forehead for the greater security of
the breathing-hole during the brief appearance at the surface, and certain
remarkable changes in the respiratory and circulatory organs which enable
the animal to remain for a long time under water. I might have added many
more, for the list of adaptations in the whale to aquatic life is by no means
exhausted; they are found in the histological structure and in the minutest
combinations in the nervous system. For it is obvious that a tail-fin must be
used in quite a different way from a tail, which serves as a fly-brush in
hoofed animals, or as an aid to springing in the kangaroo or as a climbing
organ; it will require quite different reflex-mechanisms and nerve
combinations in the motor centres.
I used this example in order to show how unnecessary it is to assume a
special internal evolutionary power for the phylogenesis of species, for this
whole order of whales is, so to speak, made up of adaptations; it deviates in
many essential respects from the usual mammalian type, and all the
deviations are adaptations to aquatic life. But if precisely the most essential
features of the organisation thus depend upon adaptation, what is left for a

phyletic force to do, since it is these essential features of the structure it
would have to determine? There are few people now who believe in a
phyletic evolutionary power, which is not made up of the forces known to
us—adaptation and heredity—but the conviction that every part of an
organism depends upon adaptation has not yet gained a firm footing.
Nevertheless, I must continue to regard this conception as the correct one,
as I have long done.
I may be permitted one more example. The feather of a bird is a marvellous
structure, and no one will deny that as a whole it depends upon adaptation.
But what part of it does not depend upon adaptation? The hollow quill, the
shaft with its hard, thin, light cortex, and the spongy substance within it, its
square section compared with the round section of the quill, the flat barbs,
their short, hooked barbules which, in the flight-feathers, hook into one
another with just sufficient firmness to resist the pressure of the air at each
wing-beat, the lightness and firmness of the whole apparatus, the elasticity
of the vane, and so on. And yet all this belongs to an organ which is only
passively functional, and therefore can have nothing to do with the
Lamarckian principle. Nor can the feather have arisen through some
magical effect of temperature, moisture, electricity, or specific nutrition,
and thus selection is again our only anchor of safety.
But—it will be objected—the substance of which the feather consists, this
peculiar kind of horny substance, did not first arise through selection in the
course of the evolution of the birds, for it formed the covering of the scales
of their reptilian ancestors. It is quite true that a similar substance covered
the scales of the Reptiles, but why should it not have arisen among them
through selection? Or in what other way could it have arisen, since scales
are also passively useful parts? It is true that if we are only to call
adaptation what has been acquired by the species we happen to be
considering, there would remain a great deal that could not be referred to
selection; but we are postulating an evolution which has stretched back
through aeons, and in the course of which innumerable adaptations took
place, which had not merely ephemeral persistence in a genus, a family or a
class, but which was continued into whole Phyla of animals, with continual
fresh adaptations to the special conditions of each species, family, or class,
yet with persistence of the fundamental elements. Thus the feather, once
acquired, persisted in all birds, and the vertebral column, once gained by

adaptation in the lowest forms, has persisted in all the Vertebrates from
Amphioxus upwards, although with constant readaptation to the conditions
of each particular group. Thus everything we can see in animals is
adaptation, whether of to-day, or of yesterday, or of ages long gone by;
every kind of cell, whether glandular, muscular, nervous, epidermic, or
skeletal, is adapted to absolutely definite and specific functions, and every
organ which is composed of these different kinds of cells contains them in
the proper proportions, and in the particular arrangement which best serves
the function of the organ; it is thus adapted to its function.
All parts of the organism are tuned to one another, that is, they are adapted
to one another, and in the same way the organism as a whole is adapted to
the conditions of its life, and it is so at every stage of its evolution.
But all adaptations can be referred to selection; the only point that remains
doubtful is whether they all must be referred to it.
However that may be, whether the Lamarckian principle is a factor that has
coöperated with selection in evolution, or whether it is altogether fallacious,
the fact remains, that selection is the cause of a great part of the phyletic
evolution of organisms on our earth. Those who agree with me in rejecting
the Lamarckian principle will regard selection as the only guiding factor in
evolution, which creates what is new out of the transmissible variations, by
ordering and arranging these, selecting them in relation to their number and
size, as the architect does his building-stones so that a particular style must
result.[55] But the building-stones themselves, the variations, have their
basis in the influences which cause variation in those vital units which are
handed on from one generation to another, whether, taken together they
form the whole organism, as in Bacteria and other low forms of life, or only
a germ-substance, as in unicellular and multicellular organisms.

FOOTNOTES:
[33] Vorträge über Descendenztheorie, Jena, 1904, ii. 269. Eng. Transl. London,
1904, ii. p. 317.
[34] See Poulton, Essays on Evolution, Oxford, 1908. pp. xix-xxii.
[35] Origin of Species (6th edit), pp. 176 et seq.
[36] Chun, Reise der Valdivia, Leipzig, 1904.
[37] Plate, Selektionsprinzip u. Probleme der Artbildung (3rd edit.), Leipzig,
1908.
[38] Studien zur Descendenz-Theorie ii., "Die Enstehung der Zeichnung bei den
Schmetterlings-raupen," Leipzig, 1876.
[39] Origin of Species (6th edit.), p. 232.
[40] Origin of Species, p. 233; see also edit. 1, p. 242.
[41] Ibid. p. 230.
[42] The Effect of External Influences upon Development, Romanes Lecture,
Oxford, 1894.
[43] See Poulton, Essays on Evolution, 1908, pp. 316, 317.
[44] The Evolution Theory, London, 1904, i. p. 219.
[45] Report of the British Association (Bristol, 1898), London, 1899, pp. 906-
909.
[46] Proc. Ent. Soc., London, May 6, 1903.
[47] Essays on Evolution, 1889-1907, Oxford, 1908, passim, e.g. p. 269.
[48] The expression does not refer to all the enemies of this butterfly; against
ichneumon-flies, for instance, their unpleasant smell usually gives no protection.
[49] Professor Poulton has corrected some wrong descriptions which I had
unfortunately overlooked in the Plates of my book Vorträge über
Descendenztheorie, and which refer to Papilio dardanus (merope). These
mistakes are of no importance as far as an understanding of the mimicry-theory
is concerned, but I hope shortly to be able to correct them in a later edition.
[50] Journ. Linn. Soc. London (Zool.), Vol. xxvi. 1898, pp. 598-602.
[51] In Kosmos, 1879, p. 100.
[52] Habit and Instinct, London. 1896.
[53] This has been discussed in many of my earlier works. See for instance The
All-Sufficiency of Natural Selection, a reply to Herbert Spencer, London, 1893.

[54] The Evolution Theory, London, 1904, p. 144.
[55] Variation under Domestication, 1875, ii. pp. 426, 427.

III

HEREDITY AND VARIATION IN MODERN
LIGHTS
By W. Bateson, M.A., F.R.S.

Professor of Biology in the University of Cambridge

Darwin's work has the property of greatness in that it may be admired from
more aspects than one. For some the perception of the principle of Natural
Selection stands out as his most wonderful achievement to which all the rest
is subordinate. Others, among whom I would range myself, look up to him
rather as the first who plainly distinguished, collected, and comprehensively
studied that new class of evidence from which hereafter a true
understanding of the process of Evolution may be developed. We each
prefer our own standpoint of admiration; but I think that it will be in their
wider aspect that his labours will most command the veneration of
posterity.
A treatise written to advance knowledge may be read in two moods. The
reader may keep his mind passive, willing merely to receive the impress of
the writer's thought; or he may read with his attention strained and alert,
asking at every instant how the new knowledge can be used in a further
advance, watching continually for fresh footholds by which to climb higher
still. Of Shelley it has been said that he was a poet for poets: so Darwin was
a naturalist for naturalists. It is when his writings are used in the critical and
more exacting spirit with which we test the outfit for our own enterprise
that we learn their full value and strength. Whether we glance back and
compare his performance with the efforts of his predecessors, or look
forward along the course which modern research is disclosing, we shall
honour most in him not the rounded merit of finite accomplishment, but the
creative power by which he inaugurated a line of discovery endless in
variety and extension. Let us attempt thus to see his work in true
perspective between the past from which it grew, and the present which is
its consequence. Darwin attacked the problem of Evolution by reference to

facts of three classes: Variation; Heredity; Natural Selection. His work was
not as the laity suppose, a sudden and unheralded revelation, but the first
fruit of a long and hitherto barren controversy. The occurrence of variation
from type, and the hereditary transmission of such variation had of course
been long familiar to practical men, and inferences as to the possible
bearing of those phenomena on the nature of specific difference had been
from time to time drawn by naturalists. Maupertuis, for example, wrote:
"Ce qui nous reste à examiner, c'est comment d'un seul individu, il a pu
naître tant d'espèces si différentes." And again: "La Nature contient le fonds
de toutes ces variétés: mais le hasard ou l'art les mettent en œuvre. C'est
ainsi que ceux dont l'industrie s'applique à satisfaire le goût des curieux,
sont, pour ainsi dire, créateurs d'espèces nouvelles."[56]
Such passages, of which many (though few so emphatic) can be found in
eighteenth century writers, indicate a true perception of the mode of
Evolution. The speculations hinted at by Buffon,[57] developed by Erasmus
Darwin, and independently proclaimed above all by Lamarck, gave to the
doctrine of descent a wide renown. The uniformitarian teaching which Lyell
deduced from geological observation had gained acceptance. The facts of
geographical distribution[58] had been shown to be obviously inconsistent
with the Mosaic legend. Prichard, and Lawrence, following the example of
Blumenbach, had successfully demonstrated that the races of Man could be
regarded as different forms of one species, contrary to the opinion up till
then received. These treatises all begin, it is true, with a profound obeisance
to the sons of Noah, but that performed, they continue on strictly modern
lines. The question of the mutability of species was thus prominently raised.
Those who rate Lamarck no higher than did Huxley in his contemptuous
phrase "buccinator tantum," will scarcely deny that the sound of the
trumpet had carried far, or that its note was clear. If then there were few
who had already turned to evolution with positive conviction, all scientific
men must at least have known that such views had been promulgated; and
many must, as Huxley says, have taken up his own position of "critical
expectancy."[59]
Why, then, was it, that Darwin succeeded where the rest had failed? The
cause of that success was twofold. First, and obviously, in the principle of
Natural Selection he had a suggestion which would work. It might not go

the whole way, but it was true as far as it went. Evolution could thus in
great measure be fairly represented as a consequence of demonstrable
processes. Darwin seldom endangers the mechanism he devised by putting
on it strains much greater than it can bear. He at least was under no illusion
as to the omnipotence of Selection; and he introduces none of the forced
pleading which in recent years has threatened to discredit that principle.
For example, in the latest text of the Origin[60] we find him saying:
"But as my conclusions have lately been much misrepresented,
and it has been stated that I attribute the modification of species
exclusively to natural selection, I may be permitted to remark
that in the first edition of this work, and subsequently, I placed
in a most conspicuous position—namely, at the close of the
Introduction—the following words: 'I am convinced that natural
selection has been the main but not the exclusive means of
modification.'"
But apart from the invention of this reasonable hypothesis, which may well,
as Huxley estimated, "be the guide of biological and psychological
speculation for the next three or four generations," Darwin made a more
significant and imperishable contribution. Not for a few generations, but
through all ages he should be remembered as the first who showed clearly
that the problems of Heredity and Variation are soluble by observation, and
laid down the course by which we must proceed to their solution.[61] The
moment of inspiration did not come with the reading of Malthus, but with
the opening of the "first note-book on Transmutation of Species."[62]
Evolution is a process of Variation and Heredity. The older writers, though
they had some vague idea that it must be so, did not study Variation and
Heredity. Darwin did, and so begat not a theory, but a science.
The extent to which this is true, the scientific world is only beginning to
realise. So little was the fact appreciated in Darwin's own time that the
success of his writings was followed by an almost total cessation of work in
that special field. Of the causes which led to these remarkable consequences
I have spoken elsewhere. They proceeded from circumstances peculiar to
the time; but whatever the causes there is no doubt that this statement of the
result is historically exact, and those who make it their business to collect

facts elucidating the physiology of Heredity and Variation are well aware
that they will find little to reward their quest in the leading scientific
Journals of the Darwinian epoch.
In those thirty years the original stock of evidence current and in circulation
even underwent a process of attrition. As in the story of the Eastern sage
who first wrote the collected learning of the universe for his sons in a
thousand volumes and by successive compression and burning reduced
them to one and from this by further burning distilled the single ejaculation
of the Faith "There is no god but God and Mohammed is the Prophet of
God," which was all his maturer wisdom deemed essential:—so in the
books of that period do we find the corpus of genetic knowledge dwindle to
a few prerogative instances and these at last to the brief formula of an
unquestioned creed.
And yet in all else that concerns biological science this period was, in very
truth, our Golden Age, when the natural history of the earth was explored as
never before; morphology and embryology were exhaustively ransacked;
the physiology of plants and animals began to rival chemistry and physics
in precision of method and in the rapidity of its advances; and the
foundations of pathology were laid.
In contrast with this immense activity elsewhere the neglect which befel the
special physiology of Descent, or Genetics as we now call it, is astonishing.
This may of course be interpreted as meaning that the favoured studies
seemed to promise a quicker return for effort, but it would be more true to
say that those who chose these other pursuits did so without making any
such comparison; for the idea that the physiology of Heredity and Variation
was a coherent science, offering possibilities of extraordinary discovery,
was not present to their minds at all. In a word, the existence of such a
science was well nigh forgotten. It is true that in ancillary periodicals, as for
example those that treat of entomology or horticulture, or in the writings of
the already isolated systematists,[63] observations with this special bearing
were from time to time related, but the class of fact on which Darwin built
his conceptions of Heredity and Variation was not seen in the highways of
biology. It formed no part of the official curriculum of biological students,
and found no place among the subjects which their teachers were
investigating.

During this period nevertheless one distinct advance was made, that with
which Weismann's name is prominently connected. In Darwin's genetic
scheme the hereditary transmission of parental experience and its
consequences played a considerable role. Exactly how great that role was
supposed to be, he with his habitual caution refrained from specifying, for
the sufficient reason that he did not know. Nevertheless much of the process
of Evolution, especially that by which organs have become degenerate and
rudimentary, was certainly attributed by Darwin to such inheritance, though
since belief in the inheritance of acquired characters fell into dispute, the
fact has been a good deal overlooked. The Origin without "use and disuse"
would be a materially different book. A certain vacillation is discernible in
Darwin's utterances on this question, and the fact gave to the astute Butler
an opportunity for his most telling attack. The discussion which best
illustrates the genetic views of the period arose in regard to the production
of the rudimentary condition of the wings of many beetles in the Madeira
group of islands, and by comparing passages from the Origin[64] Butler
convicts Darwin of saying first that this condition was in the main the result
of Selection, with disuse aiding, and in another place that the main cause of
degeneration was disuse, but that Selection had aided. To Darwin however I
think the point would have seemed one of dialetics merely. To him the one
paramount purpose was to show that somehow an Evolution by means of
Variation and Heredity might have brought about the facts observed, and
whether they had come to pass in the one way or the other was a matter of
subordinate concern.
To us moderns the question at issue has a diminished significance. For over
all such debates a change has been brought by Weismann's challenge for
evidence that use and disuse have any transmitted effects at all. Hitherto the
transmission of many acquired characteristics had seemed to most
naturalists so obvious as not to call for demonstration.[65] Weismann's
demand for facts in support of the main proposition revealed at once that
none having real cogency could be produced. The time-honoured examples
were easily shown to be capable of different explanations. A few certainly
remain which cannot be so summarily dismissed, but—though it is
manifestly impossible here to do justice to such a subject—I think no one
will dispute that these residual and doubtful phenomena, whatever be their
true nature, are not of a kind to help us much in the interpretation of any of

those complex cases of adaptation which on the hypothesis of unguided
Natural Selection are especially difficult to understand. Use and disuse were
invoked expressly to help us over these hard places; but whatever changes
can be induced in offspring by direct treatment of the parents, they are not
of a kind to encourage hope of real assistance from that quarter. It is not to
be denied that through the collapse of this second line of argument the
Selection hypothesis has had to take an increased and perilous burden.
Various ways of meeting the difficulty have been proposed, but these
mostly resolve themselves into improbable attempts to expand or magnify
the powers of Natural Selection.
Weismann's interpellation, though negative in purpose, has had a lasting
and beneficial effect, for through his thorough demolition of the old loose
and distracting notions of inherited experience, the ground has been cleared
for the construction of a true knowledge of heredity based on experimental
fact.
In another way he made a contribution of a more positive character, for his
elaborate speculations as to the genetic meaning of cytological appearances
have led to a minute investigation of the visible phenomena occurring in
those cell-divisions by which germ-cells arise. Though the particular views
he advocated have very largely proved incompatible with the observed facts
of heredity, yet we must acknowledge that it was chiefly through the
stimulus of Weismann's ideas that those advances in cytology were made;
and though the doctrine of the continuity of germ-plasm cannot be
maintained in the form originally propounded, it is in the main true and
illuminating.[66] Nevertheless in the present state of knowledge we are still
as a rule quite unable to connect cytological appearances with any genetic
consequences and save in one respect (obviously of extreme importance—
to be spoken of later) the two sets of phenomena might, for all we can see,
be entirely distinct.
I cannot avoid attaching importance to this want of connection between the
nuclear phenomena and the features of bodily organisation. All attempts to
investigate Heredity by cytological means lie under the disadvantage that it
is the nuclear changes which can alone be effectively observed. Important
as they must surely be, I have never been persuaded that the rest of the cell
counts for nothing. What we know of the behaviour and variability of

chromosomes seems in my opinion quite incompatible with the belief that
they alone govern form, and are the sole agents responsible in heredity.[67]
If, then, progress was to be made in Genetics, work of a different kind was
required. To learn the laws of Heredity and Variation there is no other way
than that which Darwin himself followed, the direct examination of the
phenomena. A beginning could be made by collecting fortuitous
observations of this class, which have often thrown a suggestive light, but
such evidence can be at best but superficial and some more penetrating
instrument of research is required. This can only be provided by actual
experiments in breeding.
The truth of these general considerations was becoming gradually clear to
many of us when in 1900 Mendel's work was rediscovered. Segregation, a
phenomenon of the utmost novelty, was thus revealed. From that moment
not only in the problem of the origin of species, but in all the great
problems of biology a new era began. So unexpected was the discovery that
many naturalists were convinced it was untrue, and at once proclaimed
Mendel's conclusions as either altogether mistaken, or if true, of very
limited application. Many fantastic notions about the workings of Heredity
had been asserted as general principles before: this was probably only
another fancy of the same class.
Nevertheless those who had a preliminary acquaintance with the facts of
Variation were not wholly unprepared for some such revelation. The
essential deduction from the discovery of segregation was that the
characters of living things are dependent on the presence of definite
elements or factors, which are treated as units in the processes of Heredity.
These factors can thus be recombined in various ways. They act sometimes
separately, and sometimes they interact in conduction with each other,
producing their various effects. All this indicates a definiteness and specific
order in heredity, and therefore in variation. This order cannot by the nature
of the case be dependent on Natural Selection for its existence, but must be
a consequence of the fundamental chemical and physical nature of living
things. The study of Variation had from the first shown that an orderliness
of this kind was present. The bodies and the properties of livings things are
cosmic, not chaotic. No matter how low in the scale we go, never do we
find the slightest hint of a diminution in that all-pervading orderliness, nor

can we conceive an organism existing for a moment in any other state.
Moreover not only does this order prevail in normal forms, but again and
again it is to be seen in newly-sprung varieties, which by general consent
cannot have been subjected to a prolonged Selection. The discovery of
Mendelian elements admirably coincided with and at once gave a rationale
of these facts. Genetic Variation is then primarily the consequence of
additions to, or omissions from, the stock of elements which the species
contains. The further investigation of the species-problem must thus
proceed by the analytical method which breeding experiments provide.
In the nine years which have elapsed since Mendel's clue became generally
known, progress has been rapid. We now understand the process by which a
polymorphic race maintains its polymorphism. When a family consists of
dissimilar members, given the numerical proportions in which these
members are occurring, we can represent their composition symbolically
and state what types can be transmitted by the various members. The
difficulty of the "swamping effects of inter-crossing" is practically at an
end. Even the famous puzzle of sex-limited inheritance is solved, at all
events in its more regular manifestations, and we know now how it is
brought about that the normal sisters of a colour-blind man can transmit the
colour-blindness while his normal brothers cannot transmit it.
We are still only on the fringe of the inquiry. It can be seen extending and
ramifying in many directions. To enumerate these here would be
impossible. A whole new range of possibilities is being brought into view
by study of the inter-relations between the simple factors. By following up
the evidence as to segregation, indications have been obtained which can
only be interpreted as meaning that when many factors are being
simultaneously redistributed among the germ-cells, certain of them exert
what must be described as a repulsion upon other factors. We cannot
surmise whither this discovery may lead.
In the new light all the old problems wear a fresh aspect. Upon the question
of the nature of Sex, for example, the bearing of Mendelian evidence is
close. Elsewhere I have shown that from several sets of parallel
experiments the conclusion is almost forced upon us that, in the types
investigated, of the two sexes the female is to be regarded as heterozygous
in sex, containing one unpaired dominant element, while the male is

similarly homozygous in the absence of that element.[68] It is not a little
remarkable that on this point—which is the only one where observations of
the nuclear processes of gameto-genesis have yet been brought into relation
with the visible characteristics of the organisms themselves—there should
be diametrical opposition between the results of breeding experiments and
those derived from cytology.
Those who have followed the researches of the American school will be
aware that, after it had been found in certain insects that the spermatozoa
were of two kinds according as they contained or did not contain the
accessory chromosome, E. B. Wilson succeeded in proving that the sperms
possessing this accessory body were destined to form females on
fertilisation, while sperms without it form males, the eggs being apparently
indifferent. Perhaps the most striking of all this series of observations is that
lately made by T. H. Morgan,[69] since confirmed by von Baehr, that in a
Phylloxeran two kinds of spermatids are formed, respectively with and
without an accessory (in this case, double) chromosome. Of these, only
those possessing the accessory body become functional spermatozoa, the
others degenerating. We have thus an elucidation of the puzzling fact that in
these forms fertilisation results in the formation of females only. How the
males are formed—for of course males are eventually produced by the
parthenogenetic females—we do not know.
If the accessory body is really to be regarded as bearing the factor for
femaleness, then in Mendelian terms female is DD and male is DR. The
eggs are indifferent and the spermatozoa are each male, or female. But
according to the evidence derived from a study of the sex-limited descent of
certain features in other animals the conclusion seems equally clear that in
them female must be regarded as DR and male as RR. The eggs are thus
each either male or female and the spermatozoa are indifferent. How this
contradictory evidence is to be reconciled we do not yet know. The
breeding work concerns fowls, canaries, and the Currant moth (Abraxas
grossulariata). The accessory chromosome has been now observed in most
of the great divisions of insects,[70] except, as it happens, Lepidoptera. At
first sight it seems difficult to suppose that a feature apparently so
fundamental as sex should be differently constituted in different animals,
but that seems at present the least improbable inference. I mention these
two groups of facts as illustrating the nature and methods of modern genetic

work. We must proceed by minute and specific analytical investigation.
Wherever we look we find traces of the operation of precise and specific
rules.
In the light of present knowledge it is evident that before we can attack the
Species-problem with any hope of success there are vast arrears to be made
up. He would be a bold man who would now assert that there was no sense
in which the term Species might not have a strict and concrete meaning in
contradistinction to the term Variety. We have been taught to regard the
difference between species and variety as one of degree. I think it unlikely
that this conclusion will bear the test of further research. To Darwin the
question, What is a variation? presented no difficulties. Any difference
between parent and offspring was a variation. Now we have to be more
precise. First we must, as de Vries has shown, distinguish real, genetic,
variation from fluctuational variations, due to environmental and other
accidents, which cannot be transmitted. Having excluded these sources of
error the variations observed must be expressed in terms of the factors to
which they are due before their significance can be understood. For
example, numbers of the variations seen under domestication, and not a few
witnessed in nature, are simply the consequence of some ingredient being in
an unknown way omitted from the composition of the varying individual.
The variation may on the contrary be due to the addition of some new
element, but to prove that it is so is by no means an easy matter. Casual
observation is useless, for though these latter variations will always be
dominants, yet many dominant characteristics may arise from another
cause, namely the meeting of complementary factors, and special study of
each case in two generations at least is needed before these two phenomena
can be distinguished.
When such considerations are fully appreciated it will be realised that
medleys of most dissimilar occurrences are all confused together under the
term Variation. One of the first objects of genetic analysis is to disentangle
this mass of confusion.
To those who have made no study of heredity it sometimes appears that the
question of the effect of conditions in causing variation is one which we
should immediately investigate, but a little thought will show that before
any critical inquiry into such possibilities can be attempted, a knowledge of

the working of heredity under conditions as far as possible uniform must be
obtained. At the time when Darwin was writing, if a plant brought into
cultivation gave off an albino variety, such an event was without hesitation
ascribed to the change of life. Now we see that albino gametes, germs, that
is to say, which are destitute of the pigment-forming factor, may have been
originally produced by individuals standing an indefinite number of
generations back in the ancestry of the actual albino, and it is indeed almost
certain that the variation to which the appearance of the albino is due cannot
have taken place in a generation later than that of the grandparents. It is true
that when a new dominant appears we should feel greater confidence that
we were witnessing the original variation, but such events are of extreme
rarity, and no such case has come under the notice of an experimenter in
modern times, as far as I am aware. That they must have appeared is clear
enough. Nothing corresponding to the Brown-breasted Game fowl is known
wild, yet that colour is a most definite dominant, and at some moment since
Gallus bankiva was domesticated, the element on which that special colour
depends must have at least once been formed in the germ-cell of a fowl; but
we need harder evidence than any which has yet been produced before we
can declare that this novelty came through over-feeding, or change of
climate, or any other disturbance consequent on domestication. When we
reflect on the intricacies of genetic problems as we must now conceive
them there come moments when we feel almost thankful that the Mendelian
principles were unknown to Darwin. The time called for a bold
pronouncement, and he made it, to our lasting profit and delight. With fuller
knowledge we pass once more into a period of cautious expectation and
reserve.
In every arduous enterprise it is pleasanter to look back at difficulties
overcome than forward to those which still seem insurmountable, but in the
next stage there is nothing to be stained by disguising the fact that the
attributes of living things are not what we used to suppose. If they are more
complex in the sense that the properties they display are throughout so
regular[71] that the Selection of minute random variations is an unacceptable
account of the origin of their diversity, yet by virtue of that very regularity
the problem is limited in scope and thus simplified.
To begin with, we must relegate Selection to its proper place. Selection
permits the viable to continue and decides that the non-viable shall perish;

just as the temperature of our atmosphere decides that no liquid carbon shall
be found on the face of the earth: but we do not suppose that the form of the
diamond has been gradually achieved by a process of Selection. So again,
as the course of descent branches in the successive generations, Selection
determines along which branch Evolution shall proceed, but it does not
decide what novelties that branch shall bring forth. "La Nature contient le
fonds de toutes ces variétés, mais le hazard ou l'art les mettent en œuvre,"
as Maupertuis most truly said.
Not till knowledge of the genetic properties of organisms has attained to far
greater completeness can evolutionary speculations have more than a
suggestive value. By genetic experiment, cytology and physiological
chemistry aiding, we may hope to acquire such knowledge. In 1872
Nathusius wrote:[72] "Das Gesetz der Vererbung ist noch nicht erkannt; der
Apfel ist noch nicht vom Baum der Erkenntniss gefallen, welcher, der Sage
nach, Newton auf den rechten Weg zur Ergründung der Gravitationsgesetze
führte." We cannot pretend that the words are not still true, but in
Mendelian analysis the seeds of that apple-tree at last are sown.
If we were asked what discovery would do most to forward our inquiry,
what one bit of knowledge would more than any other illuminate the
problem, I think we may give the answer without hesitation. The greatest
advance that we can foresee will be made when it is found possible to
connect the geometrical phenomena of development with the chemical. The
geometrical symmetry of living things is the key to a knowledge of their
regularity, and the forces which cause it. In the symmetry of the dividing
cell the basis of that resemblance we call Heredity is contained. To imitate
the morphological phenomena of life we have to devise a system which can
divide. It must be able to divide, and to segment as—grossly—a vibrating
plate or rod does, or as an icicle can do as it becomes ribbed in a continuous
stream of water; but with this distinction, that the distribution of chemical
differences and properties must simultaneously be decided and disposed in
orderly relation to the pattern of the segmentation. Even if a model which
would do this could be constructed it might prove to be a useful beginning.
This may be looking too far ahead. If we had to choose some one piece of
more proximate knowledge which we would more especially like to
acquire, I suppose we should ask for the secret of interracial sterility.

Nothing has yet been discovered to remove the grave difficulty, by which
Huxley in particular was so much oppressed, that among the many varieties
produced under domestication—which we all regard as analogous to the
species seen in nature—no clear case of interracial sterility has been
demonstrated. The phenomenon is probably the only one to which the
domesticated products seem to afford no parallel. No solution of the
difficulty can be offered which has positive value, but it is perhaps worth
considering the facts in the light of modern ideas. It should be observed that
we are not discussing incompatibility of two species to produce offspring (a
totally distinct phenomenon), but the sterility of the offspring which many
of them do produce.
When two species, both perfectly fertile severally, produce on crossing a
sterile progeny, there is a presumption that the sterility is due to the
development in the hybrid of some substance which can only be formed by
the meeting of two complementary factors. That some such account is
correct in essence may be inferred from the well-known observation that if
the hybrid is not totally sterile but only partially so, and thus is able to form
some good germ-cells which develop into new individuals, the sterility of
these daughter-individuals is sensibly reduced or may be entirely absent.
The fertility once re-established, the sterility does not return in the later
progeny, a fact strongly suggestive of segregation. Now if the sterility of the
cross-bred be really the consequence of the meeting of two complementary
factors, we see that the phenomenon could only be produced among the
divergent offspring of one species by the acquisition of at least two new
factors; for if the acquisition of a single factor caused sterility the line
would then end. Moreover each factor must be separately acquired by
distinct individuals, for if both were present together, the possessors would
by hypothesis be sterile. And in order to imitate the case of species each of
these factors must be acquired by distinct breeds. The factors need not, and
probably would not, produce any other perceptible effects; they might, like
the colour-factors present in white flowers, make no difference in the form
or other characters. Not till the cross was actually made between the two
complementary individuals would either factor come into play, and the
effects even then might be unobserved until an attempt was made to breed
from the cross-bred.

Next, if the factors responsible for sterility were acquired, they would in all
probability be peculiar to certain individuals and would not readily be
distributed to the whole breed. Any member of the breed also into which
both the factors were introduced would drop out of the pedigree by virtue of
its sterility. Hence the evidence that the various domesticated breeds say of
dogs or fowls can when mated together produce fertile offspring, is beside
the mark. The real question is, Do they ever produce sterile offspring? I
think the evidence is clearly that sometimes they do, oftener perhaps than is
commonly supposed. These suggestions are quite amenable to experimental
tests. The most obvious way to begin is to get a pair of parents which are
known to have had any sterile offspring, and to find the proportions in
which these steriles were produced. If, as I anticipate, these proportions are
found to be definite, the rest is simple.
In passing, certain other considerations may be referred to. First, that there
are observations favouring the view that the production of totally sterile
cross-breds is seldom a universal property of two species, and that it may be
a matter of individuals, which is just what on the view here proposed would
be expected. Moreover, as we all know now, though incompatibility may be
dependent to some extent on the degree to which the species are dissimilar,
no such principle can be demonstrated to determine sterility or fertility in
general. For example, though all our Finches can breed together, the hybrids
are all sterile. Of Ducks some species can breed together without producing
the slightest sterility; others have totally sterile offspring, and so on. The
hybrids between several genera of Orchids are perfectly fertile on the
female side, and some on the male side also, but the hybrids produced
between the Turnip (Brassica napus) and the Swede (Brassica campestris),
which, according to our estimates of affinity, should be nearly allied forms,
are totally sterile.[73] Lastly, it may be recalled that in sterility we are almost
certainly considering a meristic phenomenon. Failure to divide is, we may
feel fairly sure, the immediate "cause" of the sterility. Now, though we
know very little about the heredity of meristic differences, all that we do
know points to the conclusion that the less-divided is dominant to the more-
divided, and we are thus justified in supposing that there are factors which
can arrest or prevent cell-division. My conjecture therefore is that in the
case of sterility of cross-breds we see the effect produced by a

complementary pair of such factors. This and many similar problems are
now open to our analysis.
The question is sometimes asked, Do the new lights on Variation and
Heredity make the process of Evolution easier to understand? On the whole
the answer may be given that they do. There is some appearance of loss of
simplicity, but the gain is real. As was said above, the time is not ripe for
the discussion of the origin of species. With faith in Evolution unshaken—if
indeed the word faith can be used in application to that which is certain—
we look on the manner and causation of adapted differentiation as still
wholly mysterious. As Samuel Butler so truly said: "To me it seems that the
'Origin of Variation,' whatever it is, is the only true 'Origin of Species,'"[74]
and of that Origin not one of us knows anything. But given Variation—and
it is given: assuming further that the variations are not guided into paths of
adaptation—and both to the Darwinian and to the modern school this
hypothesis appears to be sound if unproven—an evolution of species
proceeding by definite steps is more, rather than less, easy to imagine than
an evolution proceeding by the accumulation of indefinite and insensible
steps. Those who have lost themselves in contemplating the miracles of
Adaptation (whether real or spurious) have not unnaturally fixed their hopes
rather on the indefinite than on the definite changes. The reasons are
obvious. By suggesting that the steps through which an adaptative
mechanism arose were indefinite and insensible, all further trouble is
spared. While it could be said that species arise by an insensible and
imperceptible process of variation, there was clearly no use in tiring
ourselves by trying to perceive that process. This labour-saving counsel
found great favour. All that had to be done to develop evolution-theory was
to discover the good in everything, a task which, in the complete absence of
any control or test whereby to check the truth of the discovery, is not very
onerous. The doctrine "que tout est au mieux" was therefore preached with
fresh vigour, and examples of that illuminating principle were discovered
with a facility that Pangloss himself might have envied, till at last even the
spectators wearied of such dazzling performances.
But in all seriousness, why should indefinite and unlimited variation have
been regarded as a more probable account of the origin of Adaptation?
Only, I think, because the obstacle was shifted one plane back, and so
looked rather less prominent. The abundance of Adaptation, we all grant, is

an immense, almost an unsurpassable difficulty in all non-Lamarckian
views of Evolution; but if the steps by which that adaptation arose were
fortuitious, to imagine them insensible is assuredly no help. In one most
important respect indeed, as has often been observed, it is a multiplication
of troubles. For the smaller the steps, the less could Natural Selection act
upon them. Definite variations—and of the occurrence of definite variations
in abundance we have now the most convincing proof—have at least the
obvious merit that they can make and often do make a real difference in the
chances of life.
There is another aspect of the Adaptation problem to which I can allude
very briefly. May not our present ideas of the universality and precision of
Adaptation be greatly exaggerated? The fit of organism to its environment
is not after all so very close—a proposition unwelcome perhaps, but one
which could be illustrated by very copious evidence. Natural Selection is
stern, but she has her tolerant moods.
We have now most certain and irrefragable proof that much definiteness
exists in living things apart from Selection, and also much that may very
well have been preserved and so in a sense constituted by Selection. Here
the matter is likely to rest. There is a passage in the sixth edition of the
Origin which has I think been overlooked. On page 70 Darwin says, "The
tuft of hair on the breast of the wild turkey-cock cannot be of any use, and it
is doubtful whether it can be ornamental in the eyes of the female bird."
This tuft of hair is a most definite and unusual structure, and I am afraid
that the remark that it "cannot be of any use" may have been made
inadvertently; but it may have been intended, for in the first edition the
usual qualification was given and must therefore have been deliberately
excised. Anyhow I should like to think that Darwin did throw over that tuft
of hair, and that he felt relief when he had done so. Whether however we
have his great authority for such a course or not, I feel quite sure that we
shall be rightly interpreting the facts of nature if we cease to expect to find
purposefulness wherever we meet with definite structures or patterns. Such
things are, as often as not, I suspect rather of the nature of tool-marks, mere
incidents of manufacture, benefiting their possessor not more than the wire-
marks in a sheet of paper, or the ribbing on the bottom of an oriental plate
renders those objects more attractive in our eyes.

If Variation may be in any way definite, the question once more arises, may
it not be definite in direction? The belief that it is has had many supporters,
from Lamarck onwards, who held that it was guided by need, and others
who, like Nägeli, while laying no emphasis on need, yet were convinced
that there was guidance of some kind. The latter view under the name of
"Orthogenesis," devised I believe by Eimer, at the present day commends
itself to some naturalists. The objection to such a suggestion is of course
that no fragment of real evidence can be produced in its support. On the
other hand, with the experimental proof that variation consists largely in the
unpacking and repacking of an original complexity, it is not so certain as we
might like to think that the order of these events is not predetermined.
For instance the original "pack" may have been made in such a way that at
the nth division of the germ-cells of a Sweet Pea a colour-factor might be
dropped, and that at the n+nth division the hooded variety be given off, and
so on. I see no ground whatever for holding such a view, but in fairness the
possibility should not be forgotten, and in the light of modern research it
scarcely looks so absurdly improbable as before.
No one can survey the work of recent years without perceiving that
evolutionary orthodoxy developed too fast, and that a great deal has got to
come down; but this satisfaction at least remains, that in the experimental
methods which Mendel inaugurated, we have means of reaching certainty in
regard to the physiology of Heredity and Variation upon which a more
lasting structure may be built.

FOOTNOTES:
[56] Vénus Physique, contenant deux Dissertations, l'une sur l'origine des
Hommes et des Animaux; Et l'autre sur l'origine des Noirs, La Haye, 1746, pp.
124 and 129. For an introduction to the writings of Maupertuis I am indebted to
an article by Professor Lovejoy in Popular Sci. Monthly, 1902.
[57] For the fullest account of the views of these pioneers of Evolution, see the
works of Samuel Butler, especially Evolution, Old and New (2nd edit.) 1882.
Butler's claims on behalf of Buffon have met with some acceptance; but after
reading what Butler has said, and a considerable part of Buffon's own works, the
word "hinted" seems to me a sufficiently correct description of the part he
played. It is interesting to note that in the chapter on the Ass, which contains
some of his evolutionary passages, there is a reference to "plusieurs idées très-
élevées sur la génération" contained in the Letters of Maupertuis.

[58] See especially W. Lawrence, Lectures on Physiology, London, 1823, pp.
213 f.
[59] See the chapter contributed to the Life and Letters of Charles Darwin, ii. p.
195. I do not clearly understand the sense in which Darwin wrote
(Autobiography, ibid. i. p. 87): "It has sometimes been said that the success of
the Origin proved 'that the subject was in the air,' or 'that men's minds were
prepared for it.' I do not think that this is strictly true, for I occasionally sounded
not a few naturalists, and never happened to come across a single one who
seemed to doubt about the permanence of species." This experience may perhaps
have been an accident due to Darwin's isolation. The literature of the period
abounds with indications of "critical expectancy." A most interesting expression
of that feeling is given in the charming account of the "Early Days of
Darwinism" by Alfred Newton, Macmillan's Magazine, lvii. 1888, p. 241. He
tells how in 1858 when spending a dreary summer in Iceland, he and his friend,
the ornithologist John Wolley, in default of active occupation, spent their days in
discussion. "Both of us taking a keen interest in Natural History, it was but
reasonable that a question, which in those days was always coming up wherever
two or more naturalists were gathered together, should be continually recurring.
That question was, 'What is a species?' and connected therewith was the other
question, 'How did a species begin?'... Now we were of course fairly well
acquainted with what had been published on these subjects." He then enumerates
some of these publications, mentioning among others T. Vernon Wollaston's
Variation of Species—a work which has in my opinion never been adequately
appreciated. He proceeds: "Of course we never arrived at anything like a
solution of these problems, general or special, but we felt very strongly that a
solution ought to be found, and that quickly, if the study of Botany and Zoology
was to make any great advance." He then describes how on his return home he
received the famous number of the Linnean Journal on a certain evening. "I sat
up late that night to read it; and never shall I forget the impression it made upon
me. Herein was contained a perfectly simple solution of all the difficulties which
had been troubling me for months past.... I went to bed satisfied that a solution
had been found."
[60] Origin, 6th edit. (1882), p. 421.
[61] Whatever be our estimate of the importance of Natural Selection, in this we
all agree. Samuel Butler, the most brilliant, and by far the most interesting of
Darwin's opponents—whose works are at length emerging from oblivion—in his
Preface (1882) to the 2nd edition of Evolution, Old and New, repeats his earlier
expression of homage to one whom he had come to regard as an enemy: "To the
end of time, if the question be asked, 'Who taught people to believe in
Evolution?' the answer must be that it was Mr. Darwin. This is true, and it is hard
to see what palm of higher praise can be awarded to any philosopher."
[62] Life and Letters, i. pp. 276 and 83.
[63] This isolation of the systematists is the one most melancholy sequela of
Darwinism. It seems an irony that we should read in the peroration to the Origin
that when the Darwinian view is accepted "Systematists will be able to pursue
their labours as at present; but they will not be incessantly haunted by the
shadowy doubt whether this or that form be a true species. This, I feel sure, and I

speak after experience, will be no slight relief. The endless disputes whether or
not some fifty species of British brambles are good species will cease." Origin,
6th edit. (1882), p. 425. True they have ceased to attract the attention of those
who lead opinion, but anyone who will turn to the literature of systematics will
find that they have not ceased in any other sense. Should there not be something
disquieting in the fact that among the workers who come most into contact with
specific differences, are to be found the only men who have failed to be
persuaded of the unreality of those differences?
[64] 6th edit. pp. 109 and 401. See Butler, Essays on Life, Art, and Science, p.
265, reprinted 1908, and Evolution, Old and New, chap. xxii. (2nd edit.), 1882.
[65] W. Lawrence was one of the few who consistently maintained the contrary
opinion. Prichard, who previously had expressed himself in the same sense, does
not, I believe, repeat these views in his later writings, and there are signs that he
came to believe in the transmission of acquired habits. See Lawrence, Lect.
Physiol. 1823, pp. 436-437, 447. Prichard, Edin. Inaug. Disp. 1808 [not seen by
me], quoted ibid. and Nat. Hist. Man, 1843, pp. 34 f.
[66] It is interesting to see how nearly Butler was led by natural penetration, and
from absolutely opposite conclusions, back to this underlying truth: "So that
each ovum when impregnate should be considered not as descended from its
ancestors, but as being a continuation of the personality of every ovum in the
chain of its ancestry, which every ovum it actually is quite as truly as the
octogenarian is the same identity with the ovum from which he has been
developed. This process cannot stop short of the primordial cell, which again
will probably turn out to be but a brief resting-place. We therefore prove each
one of us to be actually the primordial cell which never died nor dies, but has
differentiated itself into the life of the world, all living beings whatever, being
one with it and members one of another," Life and Habit, 1878, p. 86.
[67] This view is no doubt contrary to the received opinion. I am however
interested to see it lately maintained by Driesch (Science and Philosophy of the
Organism, London, 1907, p. 233), and from the recent observations of
Godlewski it has received distinct experimental support.
[68] In other words, the ova are each either female, or male (i.e. non-female),
but the sperms are all non-female.
[69] Morgan, Proc. Soc. Exp. Biol. Med. v. 1908, and von Baehr, Zool. Anz.
xxxii. p. 507, 1908.

[70] As Wilson has proved, the unpaired body is not a universal feature even in
those orders in which it has been observed. Nearly allied types may differ. In
some it is altogether unpaired. In others it is paired with a body of much smaller
size, and by selection of various types all gradations can be demonstrated
ranging to the condition in which the members of the pair are indistinguishable
from each other.
[71] I have in view, for example, the marvellous and specific phenomena of
regeneration, and those discovered by the students of "Entwicklungsmechanik."
The circumstances of its occurrence here preclude any suggestion that this
regularity has been brought about by the workings of Selection. The attempts

thus to represent the phenomena have resulted in mere parodies of scientific
reasoning.
[72] Vorträge über Viehzucht und Rassenerkenntniss, p. 120, Berlin, 1872.
[73] See Sutton, A. W., Journ. Linn. Soc. xxxviii. p. 341, 1908.
[74] Life and Habit, London, p. 263, 1878

IV

"THE DESCENT OF MAN"
By G. Schwalbe

Professor of Anatomy in the University of Strassburg

The problem of the origin of the human race, of the descent of man, is
ranked by Huxley in his epoch-making book Man's Place in Nature, as the
deepest with which biology has to concern itself, "the question of
questions,"—the problem which underlies all others. In the same brilliant
and lucid exposition, which appeared in 1863, soon after the publication of
Darwin's Origin of Species, Huxley stated his own views in regard to this
great problem. He tells us how the idea of a natural descent of man
gradually grew up in his mind. It was especially the assertions of Owen in
regard to the total difference between the human and the simian brain that
called forth strong dissent from the great anatomist Huxley, and he easily
succeeded in showing that Owen's supposed differences had no real
existence; he even established, on the basis of his own anatomical
investigations, the proposition that the anatomical differences between the
Marmoset and the Chimpanzee are much greater than those between the
Chimpanzee and Man.
But why do we thus introduce the study of Darwin's Descent of Man, which
is to occupy us here, by insisting on the fact that Huxley had taken the field
in defence of the descent of man in 1863, while Darwin's book on the
subject did not appear till 1871? It is in order that we may clearly
understand how it happened that from this time onwards Darwin and
Huxley followed the same great aim in the most intimate association.
Huxley and Darwin working at the same Problema maximum! Huxley fiery,
impetuous, eager for battle, contemptuous of the resistance of a dull world,
or energetically triumphing over it. Darwin calm, weighing every problem
slowly, letting it mature thoroughly,—not a fighter, yet having the greater
and more lasting influence by virtue of his immense mass of critically sifted
proofs. Darwin's friend, Huxley, was the first to do him justice, to

understand his nature, and to find in it the reason why the detailed and
carefully considered book on the descent of man made its appearance so
late. Huxley, always generous, never thought of claiming priority for
himself. In enthusiastic language he tells how Darwin's immortal work, The
Origin of Species, first shed light for him on the problem of the descent of
man; the recognition of a vera causa in the transformation of species
illuminated his thoughts as with a flash. He was now content to leave what
perplexed him, what he could not yet solve, as he says himself, "in the
mighty hands of Darwin." Happy in the bustle of strife against old and
deep-rooted prejudices, against intolerance and superstition, he wielded his
sharp weapons on Darwin's behalf; wearing Darwin's armour he joyously
overthrew adversary after adversary. Darwin spoke of Huxley as his
"general agent."[75] Huxley says of himself "I am Darwin's bulldog."[76]
Thus Huxley openly acknowledged that it was Darwin's Origin of Species
that first set the problem of the descent of man in its true light, that made
the question of the origin of the human race a pressing one. That this was
the logical consequence of his book Darwin himself had long felt. He had
been reproached with intentionally shirking the application of his theory to
Man. Let us hear what he says on this point in his autobiography: "As soon
as I had become, in the year 1837 or 1838, convinced that species were
mutable productions, I could not avoid the belief that man must come under
the same law. Accordingly I collected notes on the subject for my own
satisfaction, and not for a long time with any intention of publishing.
Although in the 'Origin of Species' the derivation of any particular species
is never discussed, yet I thought it best, in order that no honourable man
should accuse me of concealing my views,[77] to add that by the work 'light
would be thrown on the origin of man and his history.' It would have been
useless and injurious to the success of the book to have paraded, without
giving any evidence, my conviction with respect to his origin."[78]
In a letter written in January, 1860, to the Rev. L. Blomefield, Darwin
expresses himself in similar terms. "With respect to man, I am very far from
wishing to obtrude my belief; but I thought it dishonest to quite conceal my
opinion."[79]
The brief allusion in the Origin of Species is so far from prominent and so
incidental that it was excusable to assume that Darwin had not touched

upon the descent of man in this work. It was solely the desire to have his
mass of evidence sufficiently complete, solely Darwin's great characteristic
of never publishing till he had carefully weighed all aspects of his subject
for years, solely, in short, his most fastidious scientific conscience that
restrained him from challenging the world in 1859 with a book in which the
theory of the descent of man was fully set forth. Three years, frequently
interrupted by ill-health, were needed for the actual writing of the book:[80]
the first edition, which appeared in 1871, was followed in 1874 by a much
improved second edition, the preparation of which he very reluctantly
undertook.[81]
This, briefly, is the history of the work, which, with the Origin of Species,
marks an epoch in the history of biological sciences—the work with which
the cautious, peace-loving investigator ventured forth from his
contemplative life into the arena of strife and unrest, and laid himself open
to all the annoyances that deep-rooted belief and prejudice, and the
prevailing tendency of scientific thought at the time could devise.
Darwin did not take this step lightly. Of great interest in this connection is a
letter written to Wallace on Dec. 22, 1857,[82] in which he says, "You ask
me whether I shall discuss 'man.' I think I shall avoid the whole subject, as
so surrounded with prejudices; though I fully admit that it is the highest and
most interesting problem for the naturalist." But his conscientiousness
compelled him to state briefly his opinion on the subject in the Origin of
Species in 1859. Nevertheless he did not escape reproaches for having been
so reticent. This is unmistakably apparent from a letter to Fritz Müller dated
Feb. 22 [1869?], in which he says: "I am thinking of writing a little essay on
the Origin of Mankind, as I have been taunted with concealing my
opinions."[83]
It might be thought that Darwin behaved thus hesitatingly, and was so slow
in deciding on the full publication of his collected material in regard to the
descent of man, because he had religious difficulties to overcome.
But this was not the case, as we can see from his admirable confession of
faith, the publication of which we owe to his son Francis.[84] Whoever
wishes really to understand the lofty character of this great man should read
these immortal lines in which he unfolds to us in simple and straightforward

words the development of his conception of the universe. He describes how,
though he was still quite orthodox during his voyage round the world on
board the Beagle, he came gradually to see, shortly afterwards (1836-1839)
that the Old Testament was no more to be trusted than the Sacred Books of
the Hindoos; the miracles by which Christianity is supported, the
discrepancies between the accounts in the different Gospels, gradually led
him to disbelieve in Christianity as a divine revelation. "Thus," he writes,
[85] "disbelief crept over me at a very slow rate, but was at last complete.
The rate was so slow that I felt no distress." But Darwin was too modest to
presume to go beyond the limits laid down by science. He wanted nothing
more than to be able to go, freely and unhampered by belief in authority or
in the Bible, as far as human knowledge could lead him. We learn this from
the concluding words of his chapter on religion "The mystery of the
beginning of all things is insoluble by us; and I for one must be content to
remain an Agnostic."[86]
Darwin was always very unwilling to give publicity to his views in regard
to religion. In a letter to Asa Gray on May 22, 1860,[87] he declares that it is
always painful to him to have to enter into discussion of religious problems.
He had, he said, no intention of writing atheistically.
Finally, let us cite one characteristic sentence from a letter from Darwin to
C. Ridley[88] (Nov. 28, 1878). A clergyman, Dr. Pusey, had asserted that
Darwin had written the Origin of Species with some relation to theology.
Darwin writes emphatically, "Many years ago when I was collecting facts
for the 'Origin,' my belief in what is called a personal God was as firm as
that of Dr. Pusey himself, and as to the eternity of matter I never troubled
myself about such insoluble questions." The expression "many years ago"
refers to the time of his voyage round the world, as has already been
pointed out. Darwin means by this utterance that the views which had
gradually developed in his mind in regard to the origin of species were quite
compatible with the faith of the Church.
If we consider all these utterances of Darwin in regard to religion and to his
outlook on life (Weltanschauung), we shall see at least so much, that
religious reflection could in no way have influenced him in regard to the
writing and publishing of his book on The Descent of Man. Darwin had
early won for himself freedom of thought, and to this freedom he remained

true to the end of his life, uninfluenced by the customs and opinions of the
world around him.
Darwin was thus inwardly fortified and armed against the host of
calumnies, accusations, and attacks called forth by the publication of the
Origin of Species, and to an even greater extent by the appearance of the
Descent of Man. But in his defence he could rely on the aid of a band of
distinguished auxiliaries of the rarest ability. His faithful confederate,
Huxley, was joined by the botanist Hooker, and, after longer resistance, by
the famous geologist Lyell, whose "conversion" afforded Darwin peculiar
satisfaction. All three took the field with enthusiasm in defence of the
natural descent of man. From Wallace, on the other hand, though he shared
with him the idea of natural selection, Darwin got no support in this matter.
Wallace expressed himself in a strange manner. He admitted everything in
regard to the morphological descent of man, but maintained, in a mystic
way, that something else, something of a spiritual nature must have been
added to what man inherited from his animal ancestors. Darwin, whose
esteem for Wallace was extraordinarily high, could not understand how he
could give utterance to such a mystical view in regard to man; the idea
seemed to him so "incredibly strange" that he thought some one else must
have added these sentences to Wallace's paper.
Even now there are thinkers who, like Wallace, shrink from applying to
man the ultimate consequences of the theory of descent. The idea that man
is derived from ape-like forms is to them unpleasant and humiliating.
So far I have been depicting the development of Darwin's work on the
descent of man. In what follows I shall endeavour to give a condensed
survey of the contents of the book.
It must at once be said that the contents of Darwin's work fall into two
parts, dealing with entirely different subjects. The Descent of Man includes
a very detailed investigation in regard to secondary sexual characters in the
animal series, and on this investigation Darwin founded a new theory, that
of sexual selection. With astonishing patience he gathered together an
immense mass of material, and showed, in regard to Arthropods and
Vertebrates, the wide distribution of secondary characters, which develop
almost exclusively in the male, and which enable him, on the one hand, to

get the better of his rivals in the struggle for the female by the greater
perfection of his weapons, and, on the other hand, to offer greater
allurements to the female through the higher development of decorative
characters, of song, or of scent-producing glands. The best equipped males
will thus crowd out the less well-equipped in the matter of reproduction,
and thus the relevant characters will be increased and perfected through
sexual selection. It is, of course, a necessary assumption that these
secondary sexual characters may be transmitted to the female, although
perhaps in rudimentary form.
As we have said, this story of sexual selection takes up a great deal of space
in Darwin's book, and it need only be considered here in so far as Darwin
applied it to the descent of man. To this latter problem the whole of Part I is
devoted, while Part III contains a discussion of sexual selection in relation
to man, and a general summary. Part II treats of sexual selection in general,
and may be disregarded in our present study. Moreover, many interesting
details must necessarily be passed over in what follows, for want of space.
The first part of the Descent of Man begins with an enumeration of the
proofs of the animal descent of man taken from the structure of the human
body. Darwin chiefly emphasises the fact that the human body consists of
the same organs and of the same tissues as those of the other mammals; he
shows also that man is subject to the same diseases and tormented by the
same parasites as the apes. He further dwells on the general agreement
exhibited by young embryonic forms, and he illustrates this by two figures
placed one above the other, one representing a human embryo, after Ecker,
the other a dog embryo, after Bischoff.[89]
Darwin finds further proofs of the animal origin of man in the reduced
structures, in themselves extremely variable, which are either absolutely
useless to their possessors, or of so little use that they could never have
developed under existing conditions. Of such vestiges he enumerates: the
defective development of the panniculus carnosus (muscle of the skin) so
widely distributed among mammals, the ear-muscles, the occasional
persistence of the animal ear-point in man, the rudimentary nictitating
membrane (plica semilunaris) in the human eye, the slight development of
the organ of smell, the general hairiness of the human body, the frequently
defective development or entire absence of the third molar (the wisdom

tooth), the vermiform appendix, the occasional reappearance of a bony
canal (foramen supracondyloideum) at the lower end of the humerus, the
rudimentary tail of man (the so-called taillessness), and so on. Of these
rudimentary structures the occasional occurrence of the animal ear-point in
man is most fully discussed. Darwin's attention was called to this interesting
structure by the sculptor Woolner. He figures such a case observed in man,
and also the head of an alleged orang-foetus, the photograph of which he
received from Nitsche.
Darwin's interpretation of Woolner's case as having arisen through a folding
over of the free edge of a pointed ear has been fully borne out by my
investigations on the external ear.[90] In particular, it was established by
these investigations that the human foetus, about the middle of its
embryonic life, possesses a pointed ear somewhat similar to that of the
monkey genus Macacus. One of Darwin's statements in regard to the head
of the orang-foetus must be corrected. A large ear with a point is shown in
the photograph,[91] but it can easily be demonstrated—and Deniker has
already pointed this out—that the figure is not that of an orang foetus at all,
for that form has much smaller ears with no point; nor can it be a gibbon-
foetus, as Deniker supposes, for the gibbon ear is also without a point. I
myself regard it as that of a Macacus-embryo. But this mistake, which is
due to Nitsche, in no way affects the fact recognised by Darwin, that ear-
forms showing the point characteristic of the animal ear occur in man with
extraordinary frequency.
Finally, there is a discussion of those rudimentary structures which occur
only in one sex, such as the rudimentary mammary glands in the male, the
vesicula prostatica, which corresponds to the uterus of the female, and
others. All these facts tell in favour of the common descent of man and all
other vertebrates. The conclusion of this section is characteristic: "It is only
our natural prejudice, and that arrogance which made our forefathers
declare that they were descended from demi-gods, which leads us to demur
to this conclusion. But the time will before long come, when it will be
thought wonderful that naturalists, who were well acquainted with the
comparative structure and development of man, and other mammals, should
have believed that each was the work of a separate act of creation."[92]

In the second chapter there is a more detailed discussion, again based upon
an extraordinary wealth of facts, of the problem as to the manner in which,
and the causes through which, man evolved from a lower form. Precisely
the same causes are here suggested for the origin of man, as for the origin
of species in general. Variability, which is a necessary assumption in regard
to all transformations, occurs in man to a high degree. Moreover, the rapid
multiplication of the human race creates conditions which necessitate an
energetic struggle for existence, and thus afford scope for the intervention
of natural selection. Of the exercise of artificial selection in the human race,
there is nothing to be said, unless we cite such cases as the grenadiers of
Frederick William I, or the population of ancient Sparta. In the passages
already referred to and in those which follow, the transmission of acquired
characters, upon which Darwin does not dwell, is taken for granted. In man,
direct effects of changed conditions can be demonstrated (for instance in
regard to bodily size), and there are also proofs of the influence exerted on
his physical constitution by increased use or disuse. Reference is here made
to the fact, established by Forbes, that the Quechua Indians of the high
plateaus of Peru show a striking development of lungs and thorax, as a
result of living constantly at high altitudes.
Such special forms of variation as arrests of development (microcephalism)
and reversion to lower forms are next discussed. Darwin himself felt[93] that
these subjects are so nearly related to the cases mentioned in the first
chapter, that many of them might as well have been dealt with there. It
seems to me that it would have been better so, for the citation of additional
instances of reversion at this place rather disturbs the logical sequence of
his ideas as to the conditions which have brought about the evolution of
man from lower forms. The instances of reversion here discussed are
microcephalism, which Darwin wrongly interpreted as atavistic,
supernumerary mammae, supernumerary digits, bicornuate uterus, the
development of abnormal muscles, and so on. Brief mention is also made of
correlative variations observed in man.
Darwin next discusses the question as to the manner in which man attained
to the erect position from the state of a climbing quadruped. Here again he
puts the influence of Natural Selection in the first rank. The immediate
progenitors of man had to maintain a struggle for existence in which
success was to the more intelligent, and to those with social instincts. The

hand of these climbing ancestors, which had little skill and served mainly
for locomotion, could only undergo further development when some early
member of the Primate series came to live more on the ground and less
among trees.
A bipedal existence thus became possible, and with it the liberation of the
hand from locomotion, and the one-sided development of the human foot.
The upright position brought about correlated variations in the bodily
structure; with the free use of the hand it became possible to manufacture
weapons and to use them; and this again resulted in a degeneration of the
powerful canine teeth and the jaws, which were then no longer necessary
for defence. Above all, however, the intelligence immediately increased,
and with it skull and brain. The nakedness of man, and the absence of a tail
(rudimentariness of the tail vertebrae) are next discussed. Darwin is inclined
to attribute the nakedness of man, not to the action of natural selection on
ancestors who originally inhabited a tropical land, but to sexual selection,
which, for aesthetic reasons, brought about the loss of the hairy covering in
man, or primarily in woman. An interesting discussion of the loss of the tail,
which, however, man shares with the anthropoid apes, some other monkeys
and lemurs, forms the conclusion of the almost superabundant material
which Darwin worked up in the second chapter. His object was to show that
some of the most distinctive human characters are in all probability directly
or indirectly due to natural selection. With characteristic modesty he adds:
[94] "Hence, if I have erred in giving to natural selection great power, which
I am very far from admitting, or in having exaggerated its power, which is
in itself probable, I have at least, as I hope, done good service in aiding to
overthrow the dogma of separate creations." At the end of the chapter he
touches upon the objection as to man's helpless and defenceless condition.
Against this he urges his intelligence and social instincts.
The two following chapters contain a detailed discussion of the objections
drawn from the supposed great differences between the mental powers of
men and animals. Darwin at once admits that the differences are enormous,
but not that any fundamental difference between the two can be found. Very
characteristic of him is the following passage: "In what manner the mental
powers were first developed in the lowest organisms, is as hopeless an
enquiry as how life itself first originated. These are problems for the distant
future, if they are ever to be solved by man."[95]

After some brief observations on instinct and intelligence, Darwin brings
forward evidence to show that the greater number of the emotional states,
such as pleasure and pain, happiness and misery, love and hate are common
to man and the higher animals. He goes on to give various examples
showing that wonder and curiosity, imitation, attention, memory and
imagination (dreams of animals), can also be observed in the higher
mammals, especially in apes. In regard even to reason there are no sharply
defined limits. A certain faculty of deliberation is characteristic of some
animals, and the more thoroughly we know an animal the more intelligence
we are inclined to credit it with. Examples are brought forward of the
intelligent and deliberate actions of apes, dogs and elephants. But although
no sharply defined differences exist between man and animals, there is,
nevertheless, a series of other mental powers which are characteristics
usually regarded as absolutely peculiar to man. Some of these
characteristics are examined in detail, and it is shown that the arguments
drawn from them are not conclusive. Man alone is said to be capable of
progressive improvement; but against this must be placed as something
analogous in animals, the fact that they learn cunning and caution through
long continued persecution. Even the use of tools is not in itself peculiar to
man (monkeys use sticks, stones and twigs), but man alone fashions and
uses implements designed for a special purpose. In this connection the
remarks taken from Lubbock in regard to the origin and gradual
development of the earliest flint implements will be read with interest; these
are similar to the observations on modern eoliths, and their bearing on the
development of the stone industry. It is interesting to learn from a letter to
Hooker,[96] that Darwin himself at first doubted whether the stone
implements discovered by Boucher de Perthes were really of the nature of
tools. With the relentless candour as to himself which characterised him, he
writes four years later in a letter to Lyell in regard to this view of Boucher
de Perthes' discoveries: "I know something about his errors, and looked at
his book many years ago, and am ashamed to think that I concluded the
whole was rubbish! Yet he has done for man something like what Agassiz
did for glaciers."[97]
To return to Darwin's further comparisons between the higher mental
powers of man and animals; He takes much of the force from the argument
that man alone is capable of abstraction and self-consciousness by his own

observations on dogs. One of the main differences between man and
animals, speech, receives detailed treatment. He points out that various
animals (birds, monkeys, dogs) have a large number of different sounds for
different emotions, that, further, man produces in common with animals a
whole series of inarticulate cries combined with gestures, and that dogs
learn to understand whole sentences of human speech. In regard to human
language, Darwin expresses a view contrary to that held by Max Müller:[98]
"I cannot doubt that language owes its origin to the imitation and
modification of various natural sounds, the voices of other animals, and
man's own instinctive cries, aided by signs and gestures." The development
of actual language presupposes a higher degree of intelligence than is found
in any kind of ape. Darwin remarks on this point:[99] "The fact of the higher
apes not using their vocal organs for speech no doubt depends on their
intelligence not having been sufficiently advanced."
The sense of beauty, too, has been alleged to be peculiar to man. In
refutation of this assertion Darwin points to the decorative colours of birds,
which are used for display. And to the last objection, that man alone has
religion, that he alone has a belief in God, it is answered "that numerous
races have existed, and still exist, who have no idea of one or more gods,
and who have no words in their languages to express such an idea."[100]
The result of the investigations recorded in this chapter is to show that,
great as the difference in mental powers between man and the higher
animals may be, it is undoubtedly only a difference "of degree and not of
kind."[101]
In the fourth chapter Darwin deals with the moral sense or conscience,
which is the most important of all differences between man and animals. It
is a result of social instincts, which lead to sympathy for other members of
the same society, to non-egoistic actions for the good of others. Darwin
shows that social tendencies are found among many animals, and that
among these love and kin-sympathy exist, and he gives examples of
animals (especially dogs) which may exhibit characters that we should call
moral in man (e.g. disinterested self-sacrifice for the sake of others). The
early ape-like progenitors of the human race were undoubtedly social. With
the increase of intelligence the moral sense develops farther; with the
acquisition of speech public opinion arises, and finally, moral sense

becomes habit. The rest of Darwin's detailed discussions on moral
philosophy may be passed over.
The fifth chapter may be very briefly summarised. In it Darwin shows that
the intellectual and moral faculties are perfected through natural selection.
He inquires how it can come about that a tribe at a low level of evolution
attains to a higher, although the best and bravest among them often pay for
their fidelity and courage with their lives without leaving any descendants.
In this case it is the sentiment of glory, praise and blame, the admiration of
others, which bring about the increase of the better members of the tribe.
Property, fixed dwellings, and the association of families into a community
are also indispensable requirements for civilisation. In the longer second
section of the fifth chapter Darwin acts mainly as recorder. On the basis of
numerous investigations, especially those of Greg, Wallace, and Galton, he
inquires how far the influence of natural selection can be demonstrated in
regard to civilised nations. In the final section, which deals with the proofs
that all civilised nations were once barbarians, Darwin again uses the results
gained by other investigators, such as Lubbock and Tylor. There are two
sets of facts which prove the proposition in question. In the first place, we
find traces of a former lower state in the customs and beliefs of all civilised
nations, and in the second place, there are proofs to show that savage races
are independently able to raise themselves a few steps in the scale of
civilisation, and that they have thus raised themselves.
In the sixth chapter of the work, Morphology comes into the foreground
once more. Darwin first goes back, however, to the argument based on the
great difference between the mental powers of the highest animals and
those of man. That this is only quantitative, not qualitative, he has already
shown. Very instructive in this connection is the reference to the enormous
difference in mental powers in another class. No one would draw from the
fact that the cochineal insect (Coccus) and the ant exhibit enormous
differences in their mental powers, the conclusion that the ant should
therefore be regarded as something quite distinct, and withdrawn from the
class of insects altogether.
Darwin next attempts to establish the specific genealogical tree of man, and
carefully weighs the differences and resemblances between the different
families of the Primates. The erect position of man is an adaptive character,

just as are the various characters referable to aquatic life in the seals, which,
notwithstanding these, are ranked as a mere family of the carnivores. The
following utterance is very characteristic of Darwin:[102] "If man had not
been his own classifier, he would never have thought of founding a separate
order for his own reception." In numerous characters not mentioned in
systematic works, in the features of the face, in the form of the nose, in the
structure of the external ear, man resembles the apes. The arrangement of
the hair in man has also much in common with the apes; as also the
occurrence of hair on the forehead of the human embryo, the beard, the
convergence of the hair of the upper and under arm towards the elbow,
which occurs not only in the anthropoid apes, but also in some American
monkeys. Darwin here adopts Wallace's explanation of the origin of the
ascending direction of the hair in the forearm of the orang,—that it has
arisen through the habit of holding the hands over the head in rain. But this
explanation cannot be maintained when we consider that this disposition of
the hair is widely distributed among the most different mammals, being
found in the dog, in the sloth, and in many of the lower monkeys.
After further careful analysis of the anatomical characters Darwin reaches
the conclusion that the New World monkeys (Platyrrhine) may be excluded
from the genealogical tree altogether, but that man is an offshoot from the
Old World monkeys (Catarrhine) whose progenitors existed as far back as
the Miocene period. Among these Old World monkeys the forms to which
man shows the greatest resemblance are the anthropoid apes, which, like
him, possess neither tail nor ischial callosities. The platyrrhine and
catarrhine monkeys have their primitive ancestor among extinct forms of
the Lemuridae. Darwin also touches on the question of the original home of
the human race and supposes that it may have been in Africa, because it is
there that man's nearest relatives, the gorilla and the chimpanzee, are found.
But he regards speculation on this point as useless. It is remarkable that, in
this connection, Darwin regards the loss of the hair-covering in man as
having some relation to a warm climate, while elsewhere he is inclined to
make sexual selection responsible for it. Darwin recognises the great gap
between man and his nearest relatives, but similar gaps exist at other parts
of the mammalian genealogical tree: the allied forms have become extinct.
After the extermination of the lower races of mankind, on the one hand, and
of the anthropoid apes on the other, which will undoubtedly take place, the

gulf will be greater than ever, since the baboons will then bound it on the
one side, and the white races on the other. Little weight need be attached to
the lack of fossil remains to fill up this gap, since the discovery of these
depends upon chance. The last part of the chapter is devoted to a discussion
of the earlier stages in the genealogy of man. Here Darwin accepts in the
main the genealogical tree, which had meantime been published by
Haeckel, who traces the pedigree back through Monotrems, Reptiles,
Amphibians, and Fishes, to Amphioxus.
Then follows an attempt to reconstruct, from the atavistic characters, a
picture of our primitive ancestor who was undoubtedly an arboreal animal.
The occurrence of rudiments of parts in one sex which only come to full
development in the other is next discussed. This state of things Darwin
regards as derived from an original hermaphroditism. In regard to the
mammary glands of the male he does not accept the theory that they are
vestigial, but considers them rather as not fully developed.
The last chapter of Part I deals with the question whether the different races
of man are to be regarded as different species, or as sub-species of a race of
monophyletic origin. The striking differences between the races are first
emphasised, and the question of the fertility or infertility of hybrids is
discussed. That fertility is the more usual is shown by the excessive fertility
of the hybrid population of Brazil. This, and the great variability of the
distinguishing characters of the different races, as well as the fact that all
grades of transition stages are found between these, while considerable
general agreement exists, tell in favour of the unity of the races and lead to
the conclusion that they all had a common primitive ancestor.
Darwin therefore classifies all the different races as sub-species of one and
the same species. Then follows an interesting inquiry into the reasons for
the extinction of human races. He recognises as the ultimate reason the
injurious effects of a change of the conditions of life, which may bring
about an increase in infantile mortality, and a diminished fertility. It is
precisely the reproductive system, among animals also, which is most
susceptible to changes in the environment.
The final section of this chapter deals with the formation of the races of
mankind. Darwin discusses the question how far the direct effect of

different conditions of life, or the inherited effects of increased use or
disuse may have brought about the characteristic differences between the
different races. Even in regard to the origin of the colour of the skin he
rejects the transmitted effects of an original difference of climate as an
explanation. In so doing he is following his tendency to exclude
Lamarckian explanations as far as possible. But here he makes gratuitous
difficulties from which, since natural selection fails, there is no escape
except by bringing in the principle of sexual selection, to which, he
regarded it as possible, skin-colouring, arrangement of hair, and form of
features might be traced. But with his characteristic conscientiousness he
guards himself thus: "I do not intend to assert that sexual selection will
account for all the differences between the races."[103]
I may be permitted a remark as to Darwin's attitude towards Lamarck.
While, at an earlier stage, when he was engaged in the preliminary labours
for his immortal work, The Origin of Species, Darwin expresses himself
very forcibly against the views of Lamarck, speaking of Lamarckian
"nonsense,"[104] and of Lamarck's "absurd, though clever work"[105] and
expressly declaring, "I attribute very little to the direct action of climate,
etc."[106] yet in later life he became more and more convinced of the
influence of external conditions. In 1876, that is, two years after the
appearance of the second edition of The Descent of Man, he writes with his
usual candid honesty: "In my opinion the greatest error which I have
committed, has been not allowing sufficient weight to the direct action of
the environment, i.e. food, climate, etc. independently of a natural
selection."[107] It is certain from this change of opinion that, if he had been
able to make up his mind to issue a third edition of The Descent of Man, he
would have ascribed a much greater influence to the effect of external
conditions in explaining the different characters of the races of man than he
did in the second edition. He would also undoubtedly have attributed less
influence to sexual selection as a factor in the origin of the different bodily
characteristics, if indeed he would not have excluded it altogether.
In Part III of the Descent two additional chapters are devoted to the
discussion of sexual selection in relation to man. These may be very briefly
referred to. Darwin here seeks to show that sexual selection has been
operative on man and his primitive progenitor. Space fails me to follow out
his interesting arguments. I can only mention that he is inclined to trace

back hairlessness, the development of the beard in man, and the
characteristic colour of the different human races to sexual selection. Since
bareness of the skin could be no advantage, but rather a disadvantage, this
character cannot have been brought about by natural selection. Darwin also
rejected a direct influence of climate as a cause of the origin of the skin-
colour. I have already expressed the opinion, based on the development of
his views as shown in his letters, that in a third edition Darwin would
probably have laid more stress on the influence of external environment. He
himself feels that there are gaps in his proofs here, and says in self-
criticism: "The views here advanced, on the part which sexual selection has
played in the history of man, want scientific precision."[108] I need here only
point out that it is impossible to explain the graduated stages of skin-colour
by sexual selection, since it would have produced races sharply defined by
their colour and not united to other races by transition stages, and this, it is
well known, is not the case. Moreover, the fact established by me,[109] that
in all races the ventral side of the trunk is paler than the dorsal side, and the
inner surface of the extremities paler than the outer side, cannot be
explained by sexual selection in the Darwinian sense.
With this I conclude my brief survey of the rich contents of Darwin's book.
I may be permitted to conclude by quoting the magnificent final words of
The Descent of Man: "We must, however, acknowledge, as it seems to me,
that man, with all his noble qualities, with sympathy which feels for the
most debased, with benevolence which extends not only to other men but to
the humblest living creature, with his god-like intellect which has
penetrated into the movements and constitution of the solar system—with
all these exalted powers—Man still bears in his bodily frame the indelible
stamp of his lowly origin."[110]
What has been the fate of Darwin's doctrines since his great achievement?
How have they been received and followed up by the scientific and lay
world? And what do the successors of the mighty hero and genius think
now in regard to the origin of the human race?
At the present time we are incomparably more favourably placed than
Darwin was for answering this question of all questions. We have at our
command an incomparably greater wealth of material than he had at his
disposal. And we are more fortunate than he in this respect, that we now

know transition-forms which help to fill up the gap, still great, between the
lowest human races and the highest apes. Let us consider for a little the
more essential additions to our knowledge since the publication of The
Descent of Man.
Since that time our knowledge of animal embryos has increased
enormously. While Darwin was obliged to content himself with comparing
a human embryo with that of a dog, there are now available the youngest
embryos of monkeys of all possible groups (Orang, Gibbon,
Semnopithecus, Macacus), thanks to Selenka's most successful tour in the
East Indies in search of such material. We can now compare corresponding
stages of the lower monkeys and of the Anthropoid apes with human
embryos, and convince ourselves of their great resemblance to one another,
thus strengthening enormously the armour prepared by Darwin in defence
of his view on man's nearest relatives. It may be said that Selenka's material
fills up the blanks in Darwin's array of proofs in the most satisfactory
manner.
The deepening of our knowledge of comparative anatomy also gives us
much surer foundations than those on which Darwin was obliged to build.
Just of late there have been many workers in the domain of the anatomy of
apes and lemurs, and their investigations extend to the most different
organs. Our knowledge of fossil apes and lemurs has also become much
wider and more exact since Darwin's time: the fossil lemurs have been
especially worked up by Cope, Forsyth Major, Ameghino, and others.
Darwin knew very little about fossil monkeys. He mentions two or three
anthropoid apes as occurring in the Miocene of Europe,[111] but only names
Dryopithecus, the largest form from the Miocene of France. It was
erroneously supposed that this form was related to Hylobates. We now
know not only a form that actually stands near to the gibbon (Pliopithecus),
and remains of other anthropoids (Pliohylobates and the fossil chimpanzee,
Palaeopithecus), but also several lower catarrhine monkeys, of which
Mesopithecus, a form nearly related to the modern Sacred Monkeys (a
species of Semnopithecus) and found in strata of the Miocene period in
Greece, is the most important. Quite recently, too, Ameghino's
investigations have made us acquainted with fossil monkeys from South
America (Anthropops, Homunculus), which, according to their discoverer,
are to be regarded as in the line of human descent.

What Darwin missed most of all—intermediate forms between apes and
man—has been recently furnished. E. Dubois, as is well known, discovered
in 1893, near Trinil in Java, in the alluvial deposits of the river Bengawan,
an important form represented by a skull-cap, some molars, and a femur.
His opinion—much disputed as it has been—that in this form, which he
named Pithecanthropus, he has found a long-desired transition-form is
shared by the present writer. And although the geological age of these
fossils, which, according to Dubois, belong to the uppermost Tertiary series,
the Pliocene has recently been fixed at a later date (the older Diluvium), the
morphological value of these interesting remains, that is, the intermediate
position of Pithecanthropus, still holds good. Volz says with justice,[112]
that even if Pithecanthropus is not the missing link, it is undoubtedly a
missing link.
As on the one hand there has been found in Pithecanthropus a form which,
though intermediate between apes and man, is nevertheless more closely
allied to the apes, so on the other hand, much progress has been made since
Darwin's day in the discovery and description of the oldest human remains.
Since the famous roof of a skull and the bones of the extremities belonging
to it were found in 1856 in the Neandertal near Düsseldorf, the most varied
judgments have been expressed in regard to the significance of the remains
and of the skull in particular. In Darwin's Descent of Man there is only a
passing allusion to them[113] in connection with the discussion of the skull-
capacity, although the investigations of Schaaffhausen, King, and Huxley
were then known. I believe I have shown, in a series of papers, that the
skull in question belongs to a form different from any of the races of man
now living, and, with King and Cope, I regard it as at least a different
species from living man, and have therefore designated it Homo
primigenius. The form unquestionably belongs to the older Diluvium, and
in the later Diluvium human forms already appear, which agree in all
essential points with existing human races.
As far back as 1886 the value of the Neandertal skull was greatly enhanced
by Fraipont's discovery of two skulls and skeletons from Spy in Belgium.
These are excellently described by their discoverer,[114] and are regarded as
belonging to the same group of forms as the Neandertal remains. In 1899
and the following years came the discovery by Gorjanovič-Kramberger of
different skeletal parts of at least ten individuals in a cave near Krapina in

Croatia.[115] It is in particular the form of the lower jaw which is different
from that of all recent races of man, and which clearly indicates the lowly
position of Homo primigenius, while, on the other hand, the long-known
skull from Gibraltar, which I[116] have referred to Homo primigenius, and
which has lately been examined in detail by Sollas,[117] has made us
acquainted with the surprising shape of the eye-orbit, of the nose, and of the
whole upper part of the face. Isolated lower jaws found at La Naulette in
Belgium, and at Malarnaud in France, increase our material which is now as
abundant as could be desired. The most recent discovery of all is that of a
skull dug up in August of this year [1908] by Klaatsch and Hauser in the
lower grotto of the Le Moustier in Southern France, but this skull has not
yet been fully described. Thus Homo primigenius must also be regarded as
occupying a position in the gap existing between the highest apes and the
lowest human races, Pithecanthropus, standing in the lower part of it, and
Homo primigenius in the higher, near man. In order to prevent
misunderstanding, I should like here to emphasise that in arranging this
structural series—anthropoid apes, Pithecanthropus, Homo primigenius,
Homo sapiens—I have no intention of establishing it as a direct
genealogical series. I shall have something to say in regard to the genetic
relations of these forms, one to another, when discussing the different
theories of descent current at the present day.[118]
In quite a different domain from that of morphological relationship, namely
in the physiological study of the blood, results have recently been gained
which are of the highest importance to the doctrine of descent. Uhlenhuth,
Nuttall, and others have established the fact that the blood-serum of a rabbit
which has previously had human blood injected into it, forms a precipitate
with human blood. This biological reaction was tried with a great variety of
mammalian species, and it was found that those far removed from man gave
no precipitate under these conditions. But as in other cases among
mammals all nearly related forms yield an almost equally marked
precipitate, so the serum of a rabbit treated with human blood and then
added to the blood of an anthropoid ape gives almost as marked a
precipitate as in human blood; the reaction to the blood of the lower Eastern
monkeys is weaker, that to the Western monkeys weaker still; indeed in this
last case there is only a slight clouding after a considerable time and no
actual precipitate. The blood of the Lemuridae (Nuttall) gives no reaction or

an extremely weak one, that of the other mammals none whatever. We have
in this not only a proof of the literal blood relationship between man and
apes, but the degree of relationship with the different main groups of apes
can be determined beyond possibility of mistake.
Finally, it must be briefly mentioned that in regard to remains of human
handicraft also, the material at our disposal has greatly increased of late
years, that, as a result of this, the opinions of archaeologists have undergone
many changes, and that, in particular, their views in regard to the age of the
human race have been greatly influenced. There is a tendency at the present
time to refer the origin of man back to Tertiary times. It is true that no
remains of Tertiary man have been found, but flints have been discovered
which, according to the opinion of most investigators, bear traces either of
use, or of very primitive workmanship. Since Rutot's time, following
Mortillet's example, investigators have called these "eoliths," and they have
been traced back by Verworn to the Miocene of the Auvergne, and by Rutot
even to the upper Oligocene. Although these eoliths are even nowadays the
subject of many different views, the preoccupation with them has kept the
problem of the age of the human race continually before us.
Geology, too, has made great progress since the days of Darwin and Lyell,
and has endeavoured with satisfactory results to arrange the human remains
of the Diluvial period in chronological order (Penck). I do not intend to
enter upon the question of the primitive home of the human race; since the
space at my disposal will not allow of my touching even very briefly upon
all the departments of science which are concerned in the problem of the
descent of man. How Darwin would have rejoiced over each of the
discoveries here briefly outlined! What use he would have made of the new
and precious material, which would have prevented the discouragement
from which he suffered when preparing the second edition of The Descent
of Man! But it was not granted to him to see this progress towards filling up
the gaps in his edifice of which he was so painfully conscious.
He did, however, have the satisfaction of seeing his ideas steadily gaining
ground, notwithstanding much hostility and deep-rooted prejudice. Even in
the years between the appearance of The Origin of Species and of the first
edition of the Descent, the idea of a natural descent of man, which was only
briefly indicated in the work of 1859, had been eagerly welcomed in some

quarters. It has been already pointed out how brilliantly Huxley contributed
to the defence and diffusion of Darwin's doctrines, and how in Man's Place
in Nature he has given us a classic work as a foundation for the doctrine of
the descent of man. As Huxley was Darwin's champion in England, so in
Germany Carl Vogt, in particular, made himself master of the Darwinian
ideas. But above all it was Haeckel who, in energy, eagerness for battle, and
knowledge may be placed side by side with Huxley, who took over the
leadership in the controversy over the new conception of the universe. As
far back as 1866, in his Generelle Morphologie, he had inquired minutely
into the question of the descent of man, and not content with urging merely
the general theory of descent from lower animal forms, he drew up for the
first time genealogical trees showing the close structural relationships of the
different animal groups; the last of these illustrated the relationships of
Mammals, and among them of all groups of the Primates, including man. It
was Haeckel's genealogical trees that formed the basis of the special
discussion of the relationships of man, in the sixth chapter of Darwin's
Descent of Man.
In the last section of this essay I shall return to Haeckel's conception of the
special descent of man, the main features of which he still upholds, and
rightly so. Haeckel has contributed more than any one else to the spread of
the Darwinian doctrine.
I can only allow myself a few words as to the spread of the theory of the
natural descent of man in other countries. The Parisian anthropological
school, founded and guided by the genius of Broca, took up the idea of the
descent of man, and made many notable contributions to it (Broca,
Manouvrier, Mahoudeau, Deniker and others). In England itself Darwin's
work did not die. Huxley took care of that, for he, with his lofty and
unprejudiced mind, dominated and inspired English biology until his death
on June 29, 1895. He had the satisfaction shortly before his death of
learning of Dubois' discovery, which he illustrated by a humourous sketch.
[119] But there are still many followers in Darwin's footsteps in England.
Keane has worked at the special genealogical tree of the Primates; Keith
has inquired which of the anthropoid apes has the greatest number of
characters in common with man; Morris concerns himself with the
evolution of man in general, especially with his acquisition of the erect
position. The recent discoveries of Pithecanthropus and Homo primigenius

are being vigorously discussed; but the present writer is not in a position to
form an opinion of the extent to which the idea of descent has penetrated
throughout England generally.
In Italy independent work in the domain of the descent of man is being
produced, especially by Morselli; with him are associated, in the
investigation of related problems, Sergi and Giuffrida-Ruggeri. From the
ranks of American investigators we may single out in particular the eminent
geologist Cope, who championed with much decision the idea of the
specific difference of Homo neandertalensis (primigenius) and maintained a
more direct descent of man from the fossil Lemuridae. In South America
too, in Argentina, new life is stirring in this department of science.
Ameghino in Buenos Ayres has awakened the fossil primates of the Pampas
formation to new life; he even believes that in his Tetraprothomo,
represented by a femur, he has discovered a direct ancestor of man.
Lehmann-Nitsche is working at the other side of the gulf between apes and
man, and he describes a remarkable first cervical vertebra (atlas) from
Monte Hermoso as belonging to a form which may bear the same relation to
Homo sapiens in South America as Homo primigenius does in the Old
World. After a minute investigation he establishes a human species Homo
neogaeus, while Ameghino ascribes this atlas vertebra to his
Tetraprothomo.
Thus throughout the whole scientific world there is arising a new life, an
eager endeavour to get nearer to Huxley's problema maximum, to penetrate
more deeply into the origin of the human race. There are to-day very few
experts in anatomy and zoology who deny the animal descent of man in
general. Religious considerations, old prejudices, the reluctance to accept
man, who so far surpasses mentally all other creatures, as descended from
"soulless" animals, prevent a few investigators from giving full adherence
to the doctrine. But there are very few of these who still postulate a special
act of creation for man. Although the majority of experts in anatomy and
zoology accept unconditionally the descent of man from lower forms, there
is much diversity of opinion among them in regard to the special line of
descent.
In trying to establish any special hypothesis of descent, whether by the
graphic method of drawing up genealogical trees or otherwise, let us always

bear in mind Darwin's words[120] and use them as a critical guiding line:
"As we have no record of the lines of descent, the pedigree can be
discovered only by observing the degrees of resemblance between the
beings which are to be classed." Darwin carries this further by stating "that
resemblances in several unimportant structures, in useless and rudimentary
organs, or not now functionally active, or in an embryological condition, are
by far the most serviceable for classification."[121] It has also to be
remembered that numerous separate points of agreement are of much
greater importance than the amount of similarity or dissimilarity in a few
points.
The hypotheses as to descent current at the present day may be divided into
two main groups. The first group seeks for the roots of the human race not
among any of the families of the apes—the anatomically nearest forms—
nor among their very similar but less specialised ancestral forms, the fossil
representatives of which we can know only in part, but, setting the monkeys
on one side, it seeks for them lower down among the fossil Eocene Pseudo-
lemuridae or Lemuridae (Cope), or even among the primitive
pentadactylous Eocene forms, which may either have led directly to the
evolution of man (Adloff), or have given rise to an ancestral form common
to apes and men (Klaatsch,[122] Giuffrida-Ruggeri). The common ancestral
form, from which man and apes are thus supposed to have arisen
independently, may explain the numerous resemblances which actually
exist between them. That is to say, all the characters upon which the great
structural resemblance between apes and man depends must have been
present in their common ancestor. Let us take an example of such a
common character. The bony external ear-passage is in general as highly
developed in the lower Eastern monkeys and the anthropoid apes as in man.
This character must, therefore, have already been present in the common
primitive form. In that case it is not easy to understand why the Western
monkeys have not also inherited the character, instead of possessing only a
tympanic ring. But it becomes more intelligible if we assume that forms
with a primitive tympanic ring were the original type, and that from these
were evolved, on the one hand, the existing New World monkeys with
persistent tympanic ring, and on the other an ancestral form common to the
lower Old World monkeys, the anthropoid apes and man. For man shares
with these the character in question, and it is also one of the "unimportant"

characters required by Darwin. Thus we have two divergent lines arising
from the ancestral form, the Western monkeys (Platyrrhine) on the one
hand, and an ancestral form common to the lower Eastern monkeys, the
anthropoid apes, and man, on the other. But considerations similar to those
which showed it to be impossible that man should have developed from an
ancestor common to him and the monkeys, yet outside of and parallel with
these, may be urged also against the likelihood of a parallel evolution of the
lower Eastern monkeys, the anthropoid apes, and man. The anthropoid apes
have in common with man many characters which are not present in the
lower Old World monkeys. These characters must therefore have been
present in the ancestral form common to the three groups. But here, again, it
is difficult to understand why the lower Eastern monkeys should not also
have inherited these characters. As this is not the case, there remains no
alternative but to assume divergent evolution from an indifferent form. The
lower Eastern monkeys are carrying on the evolution in one direction—I
might almost say towards a blind alley—while anthropoids and men have
struck out a progressive path, at first in common, which explains the many
points of resemblance between them, without regarding man as derived
directly from the anthropoids. Their many striking points of agreement
indicate a common descent, and cannot be explained as phenomena of
convergence.
I believe I have shown in the above sketch that a theory which derives man
directly from lower forms without regarding apes as transition-types leads
ad absurdum. The close structural relationship between man and monkeys
can only be understood if both are brought into the same line of evolution.
To trace man's line of descent directly back to the old Eocene mammals,
alongside of, but with no relation to these very similar forms, is to abandon
the method of exact comparison, which, as Darwin rightly recognised, alone
justifies us in drawing up genealogical trees on the basis of resemblances
and differences. The farther down we go the more does the ground slip from
beneath our feet. Even the Lemuridae show very numerous divergent
conditions, much more so the Eocene mammals (Creodonta, Condylarthra),
the chief resemblance of which to man consists in the possession of
pentadactylous hands and feet! Thus the farther course of the line of descent
disappears in the darkness of the ancestry of the mammals. With just as
much reason we might pass by the Vertebrates altogether, and go back to

the lower Invertebrates, but in that case it would be much easier to say that
man has arisen independently, and has evolved, without relation to any
animals, from the lowest primitive form to his present isolated and
dominant position. But this would be to deny all value to classification,
which must after all be the ultimate basis of a genealogical tree. We can, as
Darwin rightly observed, only infer the line of descent from the degree of
resemblance between single forms. If we regard man as directly derived
from primitive forms very far back, we have no way of explaining the many
points of agreement between him and the monkeys in general, and the
anthropoid apes in particular. These must remain an inexplicable marvel.
I have thus, I trust, shown that the first class of special theories of descent,
which assumes that man has developed, parallel with the monkeys, but
without relation to them, from very low primitive forms cannot be upheld,
because it fails to take into account the close structural affinity of man and
monkeys. I cannot but regard this hypothesis as lamentably retrograde, for it
makes impossible any application of the facts that have been discovered in
the course of the anatomical and embryological study of man and monkeys,
and indeed prejudges investigations of that class as pointless. The whole
method is perverted; an unjustifiable theory of descent is first formulated
with the aid of the imagination, and then we are asked to declare that all
structural relations between man and monkeys, and between the different
groups of the latter, are valueless,—the fact being that they are the only true
basis on which a genealogical tree can be constructed.
So much for this most modern method of classification, which has probably
found adherents because it would deliver us from the relationship to apes
which many people so much dislike. In contrast to it we have the second
class of special hypotheses of descent, which keeps strictly to the nearest
structural relationship. This is the only basis that justifies the drawing up of
a special hypothesis of descent. If this fundamental proposition be
recognised, it will be admitted that the doctrine of special descent upheld by
Haeckel, and set forth in Darwin's Descent of Man, is still valid to-day. In
the genealogical tree, man's place is quite close to the anthropoid apes;
these again have as their nearest relatives the lower Old World monkeys,
and their progenitors must be sought among the less differentiated
Platyrrhine monkeys, whose most important characters have been handed
on to the present day New World monkeys. How the different genera are to

be arranged within the general scheme indicated depends in the main on the
classificatory value attributed to individual characters. This is particularly
true in regard to Pithecanthropus, which I consider as the root of a branch
which has sprung from the anthropoid ape root and has led up to man; the
latter I have designated the family of the Hominidae.
For the rest, there are, as we have said, various possible ways of
constructing the narrower genealogy within the limits of this branch
including men and apes, and these methods will probably continue to
change with the accumulation of new facts. Haeckel himself has modified
his genealogical tree of the Primates in certain details since the publication
of his Generelle Morphologie in 1866, but its general basis remains the
same.[123] All the special genealogical trees drawn up on the lines laid down
by Haeckel and Darwin—and that of Dubois may be specially mentioned—
are based, in general, on the close relationship of monkeys and men,
although they may vary in detail. Various hypotheses have been formulated
on these lines, with special reference to the evolution of man.
Pithecanthropus is regarded by some authorities as the direct ancestor of
man, by others as a side-track failure in the attempt at the evolution of man.
The problem of the monophyletic or polyphyletic origin of the human race
has also been much discussed. Sergi[124] inclines towards the assumption of
a polyphyletic origin of the three main races of man, the African primitive
form of which has given rise also to the gorilla and chimpanzee, the Asiatic
to the Orang, the Gibbon, and Pithecanthropus. Kollmann regards existing
human races as derived from small primitive races (pigmies), and considers
that Homo primigenius must have arisen in a secondary and degenerative
manner.
But this is not the place, nor have I the space to criticise the various special
theories of descent. One, however, must receive particular notice.
According to Ameghino, the South American monkeys (Pitheculites) from
the oldest Tertiary of the Pampas are the forms from which have arisen the
existing American monkeys on the one hand, and on the other, the extinct
South American Homunculidae, which are also small forms. From these
last, anthropoid apes and man have, he believes, been evolved. Among the
progenitors of man, Ameghino reckons the form discovered by him
(Tetraprothomo), from which a South American primitive man, Homo
pampaeus, might be directly evolved, while on the other hand all the lower

Old World monkeys may have arisen from older fossil South American
forms (Clenialitidae), the distribution of which may be explained by the
bridge formerly existing between South America and Africa, as may be the
derivation of all existing human races from Homo pampaeus.[125] The fossil
forms discovered by Ameghino deserve the most minute investigation, as
does also the fossil man from South America of which Lehmann-
Nitsche[126] has made a thorough study.
It is obvious that, notwithstanding the necessity for fitting man's line of
descent into the genealogical tree of the Primates, especially the apes,
opinions in regard to it differ greatly in detail. This could not be otherwise,
since the different Primate forms, especially the fossile forms, are still far
from being exhaustively known. But one thing remains certain,—the idea of
the close relationship between man and monkeys set forth in Darwin's
Descent of Man. Only those who deny the many points of agreement, the
sole basis of classification, and thus of a natural genealogical tree, can look
upon the position of Darwin and Haeckel as antiquated, or as standing on an
insufficient foundation. For such a genealogical tree is nothing more than a
summarised representation of what is known in regard to the degree of
resemblance between the different forms.
Darwin's work in regard to the descent of man has not been surpassed; the
more we immerse ourselves in the study of the structural relationships
between apes and man, the more is our path illumined by the clear light
radiating from him, and through his calm and deliberate investigation,
based on a mass of material in the accumulation of which he has never had
an equal. Darwin's fame will be bound up for all time with the unprejudiced
investigation of the question of all questions, the descent of the human race.

FOOTNOTES:
[75] Life and Letters of Thomas Henry Huxley, Vol. I. p. 171, London, 1900.
[76] Ibid., p. 363.
[77] No italics in original.
[78] Life and Letters of Charles Darwin, Vol. I. p. 93.
[79] Ibid. Vol. II. p. 263.

[80] Ibid. Vol. I. p. 94.
[81] Life and Letters, Vol. III. p. 175.
[82] Ibid. Vol. II. p. 109.
[83] Ibid. Vol. III. p. 112.
[84] Ibid. Vol. I. pp. 304-317.
[85] Life and Letters, Vol. I. p. 309.
[86] Loc. cit. p. 313.
[87] Ibid. Vol. II. p. 310.
[88] Ibid. Vol. III. p. 236. ["C. Ridley," Mr. Francis Darwin points out to me,
should be H. N. Ridley. A.C.S.]
[89] Descent of Man (Popular Edit., 1901), fig. 1, p. 14.
[90] G. Schwalbe, "Das Darwin'sche Spitzohr beim menschlichen Embryo,"
Anatom. Anzeiger, 1889, pp. 176-189, and other papers.
[91] Descent of Man, fig. 3, p. 24.
[92] Descent of man, p. 6.
[93] Ibid. p. 54.
[94] Descent of Man, p. 92.
[95] Ibid. p. 100.
[96] Life and letters, Vol. II. p. 161, June 22, 1859.
[97] Ibid. Vol. III. p. 15, March 17, 1863.
[98] Descent of Man, p. 132.
[99] Ibid. pp. 136, 137.
[100] Ibid. p. 143.
[101] Ibid. p. 193.
[102] Descent of Man, p. 231.
[103] Descent of Man, p. 308.
[104] Life and Letters, Vol. II. p. 23.
[105] Loc. cit. p. 39.
[106] Loc. cit. (1856), p. 82.
[107] Ibid. Vol. III p. 159.
[108] Descent of Man, p. 924.

[109] "Die Hautfarbe des Menschen," Mitteilungen der Anthropologischen
Gessellschaft in Wien, Vol. xxxiv. pp. 331-352.
[110] Ibid. p. 947.
[111] Descent of Man, p. 240.
[112] "Das geologische Alter der Pithecanthropus-Schichten bei Trinil, Ost-
Java." Neues Jahrb. f. Mineralogie. Festband, 1907.
[113] Descent of Man, p. 82.
[114] "La race humaine de Néanderthal ou de Canstatt en Belgique." Arch. de
Biologie, VII. 1887.
[115] Gorjanovič-Kramberger. Der diluviale Mensch van Krapina in Kroatien,
1906.
[116] Studien zur Vorgeschichte des Menschen, 1906, pp. 154 ff.
[117] "On the cranial and facial characters of the Neandertal Race." Trans. R.
Soc. London, vol. 199, 1908, p. 281.
[118] Since this essay was written Schoetensack has discovered near Heidelberg
and briefly described an exceedingly interesting lower jaw from rocks between
the Pliocene and Diluvial beds. This exhibits interesting differences from the
forms of lower jaw of Homo primigenius. (Schoetensack, Der Unterkiefer des
Homo heidelbergensis, Leipzig, 1908.) G. S.
[119] Life and Letters of Thomas Henry Huxley, Vol. II. p. 394.
[120] Descent of Man, p. 229.
[121] Loc. cit.
[122] Klaatsch in his last publications speaks in the main only of an ancestral
form common to men and anthropoid apes.
[123] Haeckels latest genealogical tree is to be found in his most recent work,
Unsere Ahnenreihe. Jena, 1908.
[124] Sergi, G. Europa, 1908.
[125] See Ameghino's latest paper, "Notas preliminaries sobre el Tetraprothomo
argentinus," etc. Anales del Museo nacional de Buenos Aires, xvi. pp. 107-242,
1907.
[126] "Nouvelles recherches sur la formation pampéenne et l'homme fossile de
la République Argentine." Rivista del Museo de la Plata, T. xiv. pp. 193-488.

V

CHARLES DARWIN AS AN
ANTHROPOLOGIST
By Ernst Haeckel

Professor of Zoology in the University of Jena

The great advance that anthropology has made in the second half of the
nineteenth century is due, in the first place, to Darwin's discovery of the
origin of man. No other problem in the whole field of research is so
momentous as that of "Man's place in nature," which was justly described
by Huxley (1863) as the most fundamental of all questions. Yet the
scientific solution of this problem was impossible until the theory of
descent had been established.
It is now a hundred years since the great French biologist Jean Lamarck
published his Philosophie Zoologique. By a remarkable coincidence the
year in which that work was issued, 1809, was the year of the birth of his
most distinguished successor, Charles Darwin. Lamarck had already
recognised that the descent of man from a series of other Vertebrates—that
is, from a series of Ape-like Primates—was essentially involved in the
general theory of transformation which he had erected on a broad inductive
basis; and he had sufficient penetration to detect the agencies that had been
at work in the evolution of the erect bimanous man from the arboreal and
quadrumanous ape. He had, however, few empirical arguments to advance
in support of his hypothesis, and it could not be established until the further
development of the biological sciences—the founding of comparative
embryology by Baer (1828) and of the cell-theory by Schleiden and
Schwann (1838), the advance of physiology under Johannes Müller (1833),
and the enormous progress of palaeontology and comparative anatomy
between 1820 and 1860—provided this necessary foundation. Darwin was
the first to coordinate the ample results of these lines of research. With no
less comprehensiveness than discrimination he consolidated them as a basis
of a modified theory of descent, and associated with them his own theory of

natural selection, which we take to be distinctive of "Darwinism" in the
stricter sense. The illuminating truth of these cumulative arguments was so
great in every branch of biology that, in spite of the most vehement
opposition, the battle was won within a single decade, and Darwin secured
the general admiration and recognition that had been denied to his
forerunner, Lamarck, up to the hour of his death (1829).
Before, however, we consider the momentous influence that Darwinism has
had in anthropology, we shall find it useful to glance at its history in the
course of the last half century, and notice the various theories that have
contributed to its advance. The first attempt to give extensive expression to
the reform of biology by Darwin's work will be found in my Generelle
Morphologie (1866)[127] which was followed by a more popular treatment
of the subject in my Natürliche Schöpfungsgeschichte (1868),[128] a
compilation from the earlier work. In the first volume of the Generelle
Morphologie I endeavoured to show the great importance of evolution in
settling the fundamental questions of biological philosophy, especially in
regard to comparative anatomy. In the second volume I dealt broadly with
the principle of evolution, distinguishing ontogeny and phylogeny as its two
coordinate main branches, and associating the two in the Biogenetic Law.
The Law may be formulated thus: "Ontogeny (embryology or the
development of the individual) is a concise and compressed recapitulation
of phylogeny (the palaeontological or genealogical series) conditioned by
laws of heredity and adaptation." The "Systematic introduction to general
evolution," with which the second volume of the Generelle Morphologie
opens, was the first attempt to draw up a natural system of organisms (in
harmony with the principles of Lamarck and Darwin) in the form of a
hypothetical pedigree, and was provisionally set forth in eight genealogical
tables.
In the nineteenth chapter of the Generelle Morphologie—a part of which
has been republished, without any alteration, after a lapse of forty years—I
made a critical study of Lamarck's theory of descent and of Darwin's theory
of selection, and endeavoured to bring the complex phenomena of heredity
and adaptation under definite laws for the first time. Heredity I divided into
conservative and progressive: adaptation into indirect (or potential) and
direct (or actual). I then found it possible to give some explanation of the
correlation of the two physiological functions in the struggle for life

(selection), and to indicate the important laws of divergence (or
differentiation) and complexity (or division of labor), which are the direct
and inevitable outcome of selection. Finally, I marked off dysteleology as
the science of the aimless (vestigial, abortive, atrophied, and useless) organs
and parts of the body. In all this I worked from a strictly monistic
standpoint, and sought to explain all biological phenomena on the
mechanical and naturalistic lines that had long been recognised in the study
of inorganic nature. Then (1866), as now, being convinced of the unity of
nature, the fundamental identity of the agencies at work in the inorganic and
the organic worlds, I discarded vitalism, teleology, and all hypotheses of a
mystic character.
It was clear from the first that it was essential, in the monistic conception of
evolution, to distinguish between the laws of conservative and progressive
heredity. Conservative heredity maintains from generation to generation the
enduring characters of the species. Each organism transmits to its
descendants a part of the morphological and physiological qualities that it
has received from its parents and ancestors. On the other hand, progressive
heredity brings new characters to the species—characters that were not
found in preceding generations. Each organism may transmit to its offspring
a part of the morphological and physiological features that it has itself
acquired, by adaptation, in the course of its individual career, through the
use or disuse of particular organs, the influence of environment, climate,
nutrition, etc. At that time I gave the name of "progressive heredity" to this
inheritance of acquired characters, as a short and convenient expression, but
have since changed the term to "transformative heredity" (as distinguished
from conservative). This term is preferable, as inherited regressive
modifications (degeneration, retrograde metamorphosis, etc.) come under
the same head.
Transformative heredity—or the transmission of acquired characters—is
one of the most important principles in evolutionary science. Unless we
admit it most of the facts of comparative anatomy and physiology are
inexplicable. That was the conviction of Darwin no less than of Lamarck, of
Spencer as well as Virchow, of Huxley as well as Gegenbaur, indeed of the
great majority of speculative biologists. This fundamental principle was for
the first time called in question and assailed in 1885 by August Weismann
of Freiburg, the eminent zoologist to whom the theory of evolution owes a

great deal of valuable support, and who has attained distinction by his
extension of the theory of selection. In explanation of the phenomena of
heredity he introduced a new theory, the "theory of the continuity of the
germ-plasm." According to him the living substance in all organisms
consists of two quite distinct kinds of plasm, somatic and germinal. The
permanent germ-plasm, or the active substance of the two germ-cells (egg-
cell and sperm-cell), passes unchanged through a series of generations, and
is not affected by environmental influences. The environment modifies only
the soma-plasm, the organs and tissues of the body. The modifications that
these parts undergo through the influence of the environment or their own
activity (use and habit), do not affect the germ-plasm, and cannot therefore
be transmitted.
This theory of the continuity of the germ-plasm has been expounded by
Weismann during the last twenty-four years in a number of able volumes,
and is regarded by many biologists, such as Mr. Francis Galton, Sir E. Ray
Lankester, and Professor J. Arthur Thomson (who has recently made a
thorough-going defence of it in his important work Heredity),[129] as the
most striking advance in evolutionary science. On the other hand, the theory
has been rejected by Herbert Spencer, Sir W. Turner, Gegenbaur, Kölliker,
Hertwig, and many others. For my part I have, with all respect for the
distinguished Darwinian, contested the theory from the first, because its
whole foundation seems to me erroneous, and its deductions do not seem to
be in accord with the main facts of comparative morphology and
physiology. Weismann's theory in its entirety is a finely conceived
molecular hypothesis, but it is devoid of empirical basis. The notion of the
absolute and permanent independence of the germ-plasm, as distinguished
from the soma-plasm, is purely speculative; as is also the theory of germinal
selection. The determinants, ids, and idants, are purely hypothetical
elements. The experiments that have been devised to demonstrate their
existence really prove nothing.
It seems to me quite improper to describe this hypothetical structure as
"Neodarwinism." Darwin was just as convinced as Lamarck of the
transmission of acquired characters and its great importance in the scheme
of evolution. I had the good fortune to visit Darwin at Down three times and
discuss with him the main principles of his system, and on each occasion
we were fully agreed as to the incalculable importance of what I may call

transformative inheritance. It is only proper to point out that Weismann's
theory of the germ-plasm is in express contradiction to the fundamental
principles of Darwin and Lamarck. Nor is it more acceptable in what one
may call its "ultradarwinism"—the idea that the theory of selection explains
everything in the evolution of the organic world. This belief in the
"omnipotence of natural selection" was not shared by Darwin himself.
Assuredly, I regard it as of the utmost value, as the process of natural
selection through the struggle for life affords an explanation of the
mechanical origin of the adapted organisation. It solves the great problem:
how could the finely adapted structure of the animal or plant body be
formed unless it was built on a preconceived plan? It thus enables us to
dispense with the teleology of the metaphysician and the dualist, and to set
aside the old mythological and poetic legends of creation. The idea had
occurred in vague form to the great Empedocles 2000 years before the time
of Darwin, but it was reserved for modern research to give it ample
expression. Nevertheless, natural selection does not of itself give the
solution of all our evolutionary problems. It has to be taken in conjunction
with the transformism of Lamarck, with which it is in complete harmony.
The monumental greatness of Charles Darwin, who surpasses every other
student of science in the nineteenth century by the loftiness of his monistic
conception of nature and the progressive influence of his ideas, is perhaps
best seen in the fact that not one of his many successors has succeeded in
modifying his theory of descent in any essential point or in discovering an
entirely new standpoint in the interpretation of the organic world. Neither
Nägeli nor Weismann, neither De Vries nor Roux, has done this. Nägeli, in
his Mechanisch-Physiologische Theorie der Abstammungslehre[130] which
is to a great extent in agreement with Weismann, constructed a theory of the
idioplasm, that represents it (like the germ-plasm) as developing
continuously in a definite direction from internal causes. But his internal
"principle of progress" is at the bottom just as teleological as the vital force
of the Vitalists, and the micella structure of the idioplasm is just as
hypothetical as the "dominant" structure of the germ-plasm. In 1889 Moritz
Wagner sought to explain the origin of species by migration and isolation,
and on that basis constructed a special "migration-theory." This, however, is
not out of harmony with the theory of selection. It merely elevates one
single factor in the theory to a predominant position. Isolation is only a

special case of selection, as I had pointed out in the fifteenth chapter of my
Natural history of creation. The "mutation-theory" of De Vries,[131] that
would explain the origin of species by sudden and saltatory variations rather
than by gradual modification, is regarded by many botanists as a great step
in advance, but it is generally rejected by zoologists. It affords no
explanation of the facts of adaptation, and has no causal value.
Much more important than these theories is that of Wilhelm Roux[132] of
"the struggle of parts within the organism, a supplementation of the theory
of mechanical adaptation." He explains the functional autoformation of the
purposive structure by a combination of Darwin's principle of selection with
Lamarck's idea of transformative heredity, and applies the two in
conjunction to the facts of histology. He lays stress on the significance of
functional adaptation, which I had described in 1866, under the head of
cumulative adaptation, as the most important factor in evolution. Pointing
out its influence in the cell-life of the tissues, he puts "cellular selection"
above "personal selection," and shows how the finest conceivable
adaptations in the structure of the tissue may be brought about quite
mechanically, without preconceived plan. This "mechanical teleology" is a
valuable extension of Darwin's monistic principle of selection to the whole
field of cellular physiology and histology, and is wholly destructive of
dualistic vitalism.
The most important advance that evolution has made since Darwin and the
most valuable amplification of his theory of selection is, in my opinion, the
work of Richard Semon: Die Mneme als erhaltendes Prinzip im Wechsel
des organischen Geschehens.[133] He offers a psychological explanation of
the facts of heredity by reducing them to a process of (unconscious)
memory. The physiologist Ewald Hering had shown in 1870 that memory
must be regarded as a general function of organic matter, and that we are
quite unable to explain the chief vital phenomena, especially those of
reproduction and inheritance, unless we admit this unconscious memory. In
my essay Die Perigenesis der Plastidule[134] I elaborated this far-reaching
idea, and applied the physical principle of transmitted motion to the
plastidules, or active molecules of plasm. I concluded that "heredity is the
memory of the plastidules, and variability their power of comprehension."
This "provisional attempt to give a mechanical explanation of the
elementary processes of evolution" I afterwards extended by showing that

sensitiveness is (as Carl Nägeli, Ernst Mach, and Albrecht Rau express it) a
general quality of matter. This form of panpsychism finds its simplest
expression in the "trinity of substance."
To the two fundamental attributes that Spinoza ascribed to substance—
Extension (matter as occupying space) and Cogitation (energy, force)—we
now add the third fundamental quality of Psychoma (sensitiveness, soul). I
further elaborated this trinitarian conception of substance in the nineteenth
chapter of my Die Lebenswunder (1904),[135] and it seems to me well
calculated to afford a monistic solution of many of the antitheses of
philosophy.
This important Mneme-theory of Semon and the luminous physiological
experiments and observations associated with it not only throw considerable
light on transformative inheritance, but provide a sound physiological
foundation for the biogenetic law. I had endeavoured to show in 1874, in
the first chapter of my Anthropogenie,[136] that this fundamental law of
organic evolution holds good generally, and that there is everywhere a
direct causal connection between ontogeny and phylogeny. "Phylogenesis is
the mechanical cause of ontogenesis;" in other words, "The evolution of the
stem or race is—in accordance with the laws of heredity and adaptation—
the real cause of all the changes that appear, in a condensed form, in the
development of the individual organism from the ovum, in either the
embryo or the larva."
It is now fifty years since Charles Darwin pointed out, in the thirteenth
chapter of his epoch-making Origin of Species, the fundamental importance
of embryology in connection with his theory of descent:
"The leading facts in embryology, which are second to none in importance,
are explained on the principle of variations in the many descendants from
some one ancient progenitor, having appeared at a not very early period of
life, and having been inherited at a corresponding period."[137]
He then shows that the striking resemblance of the embryos and larvae of
closely related animals, which in the mature stage belong to widely
different species and genera, can only be explained by their descent from a
common progenitor. Fritz Müller made a closer study of these important
phenomena in the instructive instance of the Crustacean larva, as given in

his able work Für Darwin[138] (1864). I then, in 1872, extended the range so
as to include all animals (with the exception of the unicellular Protozoa)
and showed, by means of the theory of the Gastraea, that all multicellular,
tissue-forming animals—all the Metazoa—develop in essentially the same
way from the primary germ-layers.
I conceived the embryonic form, in which the whole structure consists of
only two layers of cells, and is known as the gastrula, to be the ontogenetic
recapitulation, maintained by tenacious heredity, of a primitive common
progenitor of all the Metazoa, the Gastraea. At a later date (1895)
Monticelli discovered that this conjectural ancestral form is still preserved
in certain primitive Coelenterata—Pemmatodiscus, Kunstleria, and the
nearly-related Orthonectida.
The general application of the biogenetic law to all classes of animals and
plants has been proved in my Systematische Phylogenie.[139] It has,
however, been frequently challenged, both by botanists and zoologists,
chiefly owing to the fact that many have failed to distinguish its two
essential elements, palingenesis and cenogenesis. As early as 1874 I had
emphasised, in the first chapter of my Evolution of Man, the importance of
discriminating carefully between these two sets of phenomena:
"In the evolutionary appreciation of the facts of embryology we must take
particular care to distinguish sharply and clearly between the primary,
palingenetic evolutionary processes and the secondary, cenogenetic
processes. The palingenetic phenomena, or embryonic recapitulations, are
due to heredity, to the transmission of characters from one generation to
another. They enable us to draw direct inferences in regard to corresponding
structures in the development of the species (e.g. the chorda or the branchial
arches in all vertebrate embryos). The cenogenetic phenomena, on the other
hand, or the embryonic variations, cannot be traced to inheritance from a
mature ancestor, but are due to the adaption of the embryo or the larva to
certain conditions of its individual development (e.g. the amnion, the
allantois, and the vitelline arteries in the embryos of the higher vertebrates).
These cenogenetic phenomena are later additions; we must not infer from
them that there were corresponding processes in the ancestral history, and
hence they are apt to mislead."

The fundamental importance of these facts of comparative anatomy,
atavism, and the rudimentary organs, was pointed out by Darwin in the first
part of his classic work, The Descent of Man and Selection in Relation to
Sex (1871).[140] In the "General summary and conclusion" (chap. xxi.) he
was able to say, with perfect justice: "He who is not content to look, like a
savage, at the phenomena of nature as disconnected, cannot any longer
believe that man is the work of a separate act of creation. He will be forced
to admit that the close resemblance of the embryo of man to that, for
instance, of a dog—the construction of his skull, limbs, and whole frame on
the same plan with that of other mammals, independently of the uses to
which the parts may be put—the occasional reappearance of various
structures, for instance of several muscles, which man does not normally
possess, but which are common to the Quadrumana—and a crowd of
analogous facts—all point in the plainest manner to the conclusion that man
is the co-descendant with other mammals of a common progenitor."
These few lines of Darwin's have a greater scientific value than hundreds of
those so-called "anthropological treatises," which give detailed descriptions
of single organs, or mathematical tables with series of numbers and what
are claimed to be "exact analyses," but are devoid of synoptic conclusions
and a philosophical spirit.
Charles Darwin is not generally recognised as a great anthropologist, nor
does the school of modern anthropologists regard him as a leading
authority. In Germany, especially, the great majority of the members of the
anthropological societies took up an attitude of hostility to him from the
very beginning of the controversy in 1860. The Descent of Man was not
merely rejected, but even the discussion of it was forbidden on the ground
that it was "unscientific."
The centre of this inveterate hostility for thirty years—especially after 1877
—was Rudolph Virchow of Berlin, the leading investigator in pathological
anatomy, who did so much for the reform of medicine by his establishment
of cellular pathology in 1858. As a prominent representative of "exact" or
"descriptive" anthropology, and lacking a broad equipment in comparative
anatomy and ontogeny, he was unable to accept the theory of descent. In
earlier years, and especially during his splendid period of activity at
Würzburg (1848-1856), he had been a consistent free-thinker, and had in a

number of able articles (collected in his Gesammelte Abhandlungen)[141]
upheld the unity of human nature, the inseparability of body and spirit. In
later years at Berlin, where he was more occupied with political work and
sociology (especially after 1866), he abandoned the positive monistic
position for one of agnosticism and scepticism, and made concessions to the
dualistic dogma of a spiritual world apart from the material frame.
In the course of a Scientific Congress at Munich in 1877 the conflict of
these antithetic views of nature came into sharp relief. At this memorable
Congress I had undertaken to deliver the first address (September 18th) on
the subject of "Modern evolution in relation to the whole of science." I
maintained that Darwin's theory not only solved the great problem of the
origin of species, but that its implications, especially in regard to the nature
of man, threw considerable light on the whole of science, and on
anthropology in particular. The discovery of the real origin of man by
evolution from a long series of mammal ancestors threw light on his place
in nature in every respect, as Huxley had already shown in his excellent
lectures of 1863. Just as all the organs and tissues of the human body had
originated from those of the nearest related mammals, certain ape-like
forms, so we were bound to conclude that his mental qualities also had been
derived from those of his extinct primate ancestor.
This monistic view of the origin and nature of man, which is now admitted
by nearly all who have the requisite acquaintance with biology, and
approach the subject without prejudice, encountered a sharp opposition at
that time. The opposition found its strongest expression in an address that
Virchow delivered at Munich four days afterwards (September 22nd), on
"The freedom of science in the modern State." He spoke of the theory of
evolution as an unproved hypothesis, and declared that it ought not to be
taught in the schools, because it was dangerous to the State. "We must not,"
he said, "teach that man has descended from the ape or any other animal."
When Darwin, usually so lenient in his judgment, read the English
translation of Virchow's speech, he expressed his disapproval in strong
terms. But the great authority that Virchow had—an authority well founded
in pathology and sociology—and his prestige as president of the German
Anthropological Society, had the effect of preventing any member of the
Society from raising serious opposition to him for thirty years. Numbers of
journals and treatises repeated his dogmatic statement: "It is quite certain

that man has descended neither from the ape nor from any other animal." In
this he persisted till his death in 1902. Since that time the whole position of
German anthropology has changed. The question is no longer whether man
was created by a distinct supernatural act or evolved from other mammals,
but to which line of the animal hierarchy we must look for the actual series
of ancestors. The interested reader will find an account of this "battle of
Munich" (1877) in my three Berlin lectures (April, 1905), Der Kampf um
die Entwickelungs-Gedanken.[142]
The main points in our genealogical tree were clearly recognised by Darwin
in the sixth chapter of the Descent of Man. Lowly organised fishes, like the
lancelot (Amphioxus), are descended from lower invertebrates resembling
the larvae of an existing Tunicate (Appendicularia). From these primitive
fishes were evolved higher fishes of the ganoid type and others of the type
of Lepidosiren (Dipneusta). It is a very small step from these to the
Amphibia:
"In the class of animals the steps are not difficult to conceive which led
from the ancient Monotremata to the ancient Marsupials; and from these to
the early progenitors of the placental mammals. We may thus ascend to the
Lemuridae; and the interval is not very wide from these to the Simiadae.
The Simiadae then branched off into two great stems, the New World and
Old World monkeys; and from the latter, at a remote period, Man, the
wonder and glory of the Universe, proceeded."[143]
In these few lines Darwin clearly indicated the way in which we were to
conceive our ancestral series within the vertebrates. It is fully confirmed by
all the arguments of comparative anatomy and embryology, of
palaeontology and physiology; and all the research of the subsequent forty
years have gone to establish it. The deep interest in geology which Darwin
maintained throughout his life and his complete knowledge of
palaeontology enabled him to grasp the fundamental importance of the
palaeontological record more clearly than anthropologists and zoologists
usually do.
There has been much debate in subsequent decades whether Darwin himself
maintained that man was descended from the ape, and many writers have
sought to deny it. But the lines I have quoted verbatim from the conclusion

of the sixth chapter of the Descent of Man (1871) leave no doubt that he
was as firmly convinced of it as was his great precursor Jean Lamarck in
1809. Moreover, Darwin adds, with particular explicitness, in the "general
summary and conclusion" (chap. xxi.) of that standard work:[144]
"By considering the embryological structure of man—the homologies
which he presents with the lower animals,—the rudiments which he retains,
—and the reversions to which he is liable, we can partly recall in
imagination the former condition of our early progenitors; and can
approximately place them in their proper place in the zoological series. We
thus learn that man is descended from a hairy, tailed quadruped, probably
arboreal in its habits, and an inhabitant of the Old World. This creature, if
its whole structure had been examined by a naturalist, would have been
classed amongst the Quadrumana, as surely as the still more ancient
progenitor of the Old and New World monkeys."
These clear and definite lines leave no doubt that Darwin—so critical and
cautious in regard to important conclusions—was quite as firmly convinced
of the descent of man from the apes (the Catarrhinae, in particular) as
Lamarck was in 1809 and Huxley in 1863.
It is to be noted particularly that, in these and other observations on the
subject, Darwin decidedly assumes the monophyletic origin of the
mammals, including man. It is my own conviction that this is of the greatest
importance. A number of difficult questions in regard to the development of
man, in respect of anatomy, physiology, psychology, and embryology, are
easily settled if we do not merely extend our progonotaxis to our nearest
relatives, the anthropoid apes and the tailed monkeys from which these
have descended, but go further back and find an ancestor in the group of the
Lemuridae, and still further back to the Marsupials and Monotremata. The
essential identity of all the Mammals in point of anatomical structure and
embryonic development—in spite of their astonishing differences in
external appearance and habits of life—is so palpably significant that
modern zoologists are agreed in the hypothesis that they have all sprung
from a common root, and that this root may be sought in the earlier
Palaeozoic Amphibia.

The fundamental importance of this comparative morphology of the
Mammals, as a sound basis of scientific anthropology, was recognised just
before the beginning of the nineteenth century, when Lamarck first
emphasised (1794) the division of the animal kingdom into Vertebrates and
Invertebrates. Even thirteen years earlier (1781), when Goethe made a close
study of the mammal skeleton in the Anatomical Institute at Jena, he was
intensely interested to find that the composition of the skull was the same in
man as in the other mammals. His discovery of the os inter-maxillare in
man (1784), which was contradicted by most of the anatomists of the time,
and his ingenious "vertebral theory of the skull," were the splendid fruit of
his morphological studies. They remind us how Germany's greatest
philosopher and poet was for many years ardently absorbed in the
comparative anatomy of man and the mammals, and how he divined that
their wonderful identity in structure was no mere superficial resemblance,
but pointed to a deep internal connection. In my Generelle Morphologie
(1866), in which I published the first attempts to construct phylogenetic
trees, I have given a number of remarkable theses of Goethe, which may be
called "phyletic prophecies." They justify us in regarding him as a precursor
of Darwin.
In the ensuing forty years I have made many conscientious efforts to
penetrate further along that line of anthropological research that was opened
up by Goethe, Lamarck, and Darwin. I have brought together the many
valuable results that have constantly been reached in comparative anatomy,
physiology, ontogeny, and palaeontology, and maintained the effort to
reform the classification of animals and plants in an evolutionary sense. The
first rough drafts of pedigrees that were published in the Generelle
Morphologie have been improved time after time in the ten editions of my
Natürlich Schöpfungsgeschichte (1868-1902).[145] A sounded basis for my
phyletic hypotheses, derived from a discriminating combination of the three
great records—morphology, ontogeny, and palaeontology—was provided in
the three volumes of my Systematische Phylogenie[146] (1894 Protists and
Plants, 1895 Vertebrates, 1896 Invertebrates).
In my Anthropogenie[147] I endeavoured to employ all the known facts of
comparative ontogeny (embryology) for the purpose of completing my
scheme of human phylogeny (evolution). I attempted to sketch the historical
development of each organ of the body, beginning with the most elementary

structures in the germ-layers of the Gastraea. At the same time I drew up a
corrected statement of the most important steps in the line of our ancestral
series.
At the fourth International Congress of Zoology at Cambridge (August
26th, 1898) I delivered an address on "Our present knowledge of the
Descent of Man." It was translated into English, enriched with many
valuable notes and additions, by my friend and pupil in earlier days Dr.
Hans Gadow (Cambridge), and published under the title: The Last Link: our
present knowledge of the Descent of Man[148] The determination of the chief
animal forms that occur in the line of our ancestry is there restricted to
thirty types, and these are distributed in six main groups.
The first half of this "Progonotaxis hominis," which has no support from
fossil evidence, comprises three groups: (i) Protista (unicellular organisms,
1-5): (ii) Invertebrate Metazoa (Coelenteria 6-8, Vermalia 9-11): (iii)
Monorrhine Vertebrates (Acrania 12-13, Cyclostoma 14-15). The second
half, which is based on fossil records, also comprises three groups: (iv)
Palaeozoic cold-blooded Craniota (Fishes 16-18, Amphibia 19, Reptiles
20): (v) Mesozoic Mammals (Monotrema 21, Marsupialia 22, Mallotheria
23): (vi) Cenozoic Primates (Lemuridae 24-25, Tailed Apes 26-27,
Anthropomorpha 28-30). An improved and enlarged edition of this
hypothetic "Progonotaxis hominis" was published in 1908, in my essay
Unsere Ahnenreihe.[149]
If I have succeeded in furthering, in some degree, by these anthropological
works, the solution of the great problem of Man's place in nature, and
particularly in helping to trace the definite stages in our ancestral series, I
owe the success, not merely to the vast progress that biology has made in
the last half century, but largely to the luminous example of the great
investigators who have applied themselves to the problem, with so much
assiduity and genius, for a century and a quarter—I mean Goethe and
Lamarck, Gegenbaur and Huxley, but, above all, Charles Darwin. It was the
great genius of Darwin that first brought together that symmetrical temple
of scientific knowledge, the theory of descent. It was Darwin who put the
crown on the edifice by his theory of natural selection. Not until this broad
inductive law was firmly established was it possible to vindicate the special
conclusion, the descent of man from a series of other Vertebrates. By his

illuminating discovery Darwin did more for anthropology than thousands of
those writers, who are more specifically titled anthropologists, have done
by their technical treatises. We may, indeed, say that it is not merely as an
exact observer and ingenious experimenter, but as a distinguished
anthropologist and far-seeing thinker, that Darwin takes his place among the
greatest men of science of the nineteenth century.
To appreciate fully the immortal merit of Darwin in connection with
anthropology, we must remember that not only did his chief work, The
Origin of Species, which opened up a new era in natural history in 1859,
sustain the most virulent and widespread opposition for a lengthy period,
but even thirty years later, when its principles were generally recognised
and adopted, the application of them to man was energetically contested by
many high scientific authorities. Even Alfred Russel Wallace, who
discovered the principle of natural selection independently in 1858, did not
concede that it was applicable to the higher mental and moral qualities of
man. Dr. Wallace still holds a spiritualist and dualist view of the nature of
man, contending that he is composed of a material frame (descended from
the apes) and an immortal immaterial soul (infused by a higher power). This
dual conception, moreover, is still predominant in the wide circles of
modern theology and metaphysics, and has the general and influential
adherence of the more conservative classes of society.
In strict contradiction to this mystical dualism, which is generally connected
with teleology and vitalism, Darwin always maintained the complete unity
of human nature, and showed convincingly that the psychological side of
man was developed, in the same way as the body, from the less advanced
soul of the anthropoid ape, and, at a still more remote period, from the
cerebral functions of the older vertebrates. The eighth chapter of the Origin
of Species, which is devoted to instinct, contains weighty evidence that the
instincts of animals are subject, like all other vital processes, to the general
laws of historic development. The special instincts of particular species
were formed by adaptation, and the modifications thus acquired were
handed on to posterity by heredity; in their formation and preservation
natural selection plays the same part as in the transformation of every other
physiological function. The higher moral qualities of civilised man have
been derived from the lower mental functions of the uncultivated barbarians
and savages, and these in turn from the social instincts of the mammals.

This natural and monistic psychology of Darwin's was afterwards more
fully developed by his friend George Romanes in his excellent works
Mental Evolution in Animals and Mental Evolution in Man.[150]
Many valuable and most interesting contributions to this monistic
psychology of man were made by Darwin in his fine work on The Descent
of Man and Selection in Relation to Sex, and again in his supplementary
work, The Expression of the Emotions in Man and Animals. To understand
the historical development of Darwin's anthropology one must read his life
and the introduction to The Descent of Man. From the moment that he was
convinced of the truth of the principle of descent—that is to say, from his
thirtieth year, in 1838—he recognised clearly that man could not be
excluded from its range. He recognised as a logical necessity the important
conclusion that "man is the co-descendant with other species of some
ancient, lower, and extinct form." For many years he gathered notes and
arguments in support of this thesis, and for the purpose of showing the
probable line of man's ancestry. But in the first edition of The Origin of
Species (1859) he restricted himself to the single line, that by this work
"light would be thrown on the origin of man and his history." In the fifty
years that have elapsed since that time the science of the origin and nature
of man has made astonishing progress, and we are now fairly agreed in a
monistic conception of nature that regards the whole universe, including
man, as a wonderful unity, governed by unalterable and eternal laws. In my
philosophical book Die Welträtsel (1899)[151] and in the supplementary
volume Die Lebenswunder (1904)[152] I have endeavoured to show that this
pure monism is securely established, and that the admission of the all-
powerful rule of the same principle of evolution throughout the universe
compels us to formulate a single supreme law—the all-embracing "Law of
Substance," or the united laws of the constancy of matter and the
conservation of energy. We should never have reached this supreme general
conception if Charles Darwin—a "monistic philosopher" in the true sense of
the word—had not prepared the way by his theory of descent by natural
selection, and crowned the great work of his life by the association of this
theory with a naturalistic anthropology.

FOOTNOTES:
[127] Generelle Morphologie der Organismen, 2 vols., Berlin, 1866.
[128] Eng. transl.; The History of Creation, London, 1876.
[129] London, 1908.
[130] Munich, 1884.
[131] Die Mutationstheorie, Leipzig, 1903.
[132] Der Kampf der Theile im Organismus, Leipzig, 1881.
[133] Leipzig, 1904.
[134] Berlin, 1876.
[135] Wonders of Life, London and New York, 1904.
[136] Eng. transl.; The Evolution of Man, 2 vols., London, 1879 and 1905.
[137] Origin of Species (6th edit.), p. 396.
[138] Eng. transl.; Facts and Arguments for Darwin, London, 1869.
[139] 3 vols., Berlin, 1894-96.
[140] Descent of Man (Popular Edit.), p. 927.
[141] Gesammelte Abhandlungen zur wissenschaftlichen Medizin, Berlin, 1856.
[142] Eng. transl.; Last Words on Evolution, London, 1906.
[143] Descent of Man, (Popular Edit.), p. 255.
[144] Descent of Man, p. 930.
[145] Eng. transl.; The History of Creation, London, 1876.
[146] Berlin, 1894-96.
[147] Leipzig, 1874, 5th edit. 1905. Eng. transl.; The Evolution of Man, London,
1905.
[148] London, 1898.
[149] Festschrift zur 350-jährigen Jubelfeier der Thüringer Universität Jena.
Jena. 1908.
[150] London, 1885; 1888.
[151] The Riddle of the Universe, London and New York, 1900.
[152] The Wonders of Life, London and New York, 1904.

VI

MENTAL FACTORS IN EVOLUTION
By C. Lloyd Morgan, LLD., F.R.S
In developing his conception of organic evolution Charles Darwin was of
necessity brought into contact with some of the problems of mental
evolution. In The Origin of Species he devoted a chapter to "the diversities
of instinct and of the other mental faculties in animals of the same class."
[153] When he passed to the detailed consideration of The Descent of Man, it
was part of his object to show "that there is no fundamental difference
between man and the higher mammals in their mental faculties."[154] "If no
organic being excepting man," he said, "had possessed any mental power, or
if his powers had been of a wholly different nature, from those of the lower
animals, then we should never have been able to convince ourselves that
our high faculties had been gradually developed."[155] In his discussion of
The Expression of the Emotions it was important for his purpose "fully to
recognise that actions readily become associated with other actions and
with various states of the mind."[156] His hypothesis of sexual selection is
largely dependent upon the exercise of choice on the part of the female and
her preference for "not only the more attractive but at the same time the
more vigourous and vicious males."[157] Mental processes and physiological
processes were for Darwin closely correlated; and he accepted the
conclusion "that the nervous system not only regulates most of the existing
functions of the body, but has indirectly influenced the progressive
development of various bodily structures and of certain mental qualities."
[158]

Throughout his treatment, mental evolution was for Darwin incidental to
and contributory to organic evolution. For specialised research in
comparative and genetic psychology, as an independent field of
investigation, he had neither the time nor the requisite training. None the
less his writings and the spirit of his work have exercised a profound
influence on this department of evolutionary thought. And, for those who
follow Darwin's lead, mental evolution is still in a measure subservient to
organic evolution. Mental processes are the accompaniments or

concomitants of the functional activity of specially differentiated parts of
the organism. They are in some way dependent on physiological and
physical conditions. But though they are not physical in their nature, and
though it is difficult or impossible to conceive that they are physical in their
origin, they are, for Darwin and his followers, factors in the evolutionary
process in its physical or organic aspect. By the physiologist within his
special and well-defined universe of discourse they may be properly
regarded as epiphenomena; but by the naturalist in his more catholic survey
of nature they cannot be so regarded, and were not so regarded by Darwin.
Intelligence has contributed to evolution of which it is in a sense a product.
The facts of observation or of inference which Darwin accepted are these:
Conscious experience accompanies some of the modes of animal behaviour;
it is concomitant with certain physiological processes; these processes are
the outcome of development in the individual and evolution in the race; the
accompanying mental processes undergo a like development. Into the subtle
philosophical questions which arise out of the naïve acceptance of such a
creed it was not Darwin's province to enter; "I have nothing to do," he said,
[159] "with the origin of the mental powers, any more than I have with that
of life itself." He dealt with the natural history of organisms, including not
only their structure but their modes of behaviour; with the natural history of
the states of consciousness which accompany some of their actions; and
with the relation of behaviour to experience. We will endeavour to follow
Darwin in his modesty and candour in making no pretence to give ultimate
explanations. But we must note one of the implications of this self-denying
ordinance of science. Development and evolution imply continuity. For
Darwin and his followers the continuity is organic through physical
heredity. Apart from speculative hypothesis, legitimate enough in its proper
place but here out of court, we know nothing of continuity of mental
evolution as such: consciousness appears afresh in each succeeding
generation. Hence it is that for those who follow Darwin's lead, mental
evolution is and must ever be, within his universe of discourse, subservient
to organic evolution. Only in so far as conscious experience, or its neural
correlate, effects some changes in organic structure can it influence the
course of heredity; and conversely only in so far as changes in organic
structure are transmitted through heredity, is mental evolution rendered
possible. Such is the logical outcome of Darwin's teaching.

Those who abide by the cardinal results of this teaching are bound to regard
all behaviour as the expression of the functional activities of the living
tissues of the organism, and all conscious experience as correlated with
such activities. For the purposes of scientific treatment, mental processes
are one mode of expression of the same changes of which the physiological
processes accompanying behaviour are another mode of expression. This is
simply accepted as a fact which others may seek to explain. The behaviour
itself is the adaptive application of the energies of the organism; it is called
forth by some form of presentation or stimulation brought to bear on the
organism by the environment. This presentation is always an individual or
personal matter. But in order that the organism may be fitted to respond to
the presentation of the environment it must have undergone in some way a
suitable preparation. According to the theory of evolution this preparation is
primarily racial and is transmitted through heredity. Darwin's main thesis
was that the method of preparation is predominantly by natural selection.
Subordinate to racial preparation, and always dependent thereon, is
individual or personal preparation through some kind of acquisition; of
which the guidance of behaviour through individually won experience is a
typical example. We here introduce the mental factor because the facts seem
to justify the inference. Thus there are some modes of behaviour which are
wholly and solely dependent upon inherited racial preparation; there are
other modes of behaviour which are also dependent, in part at least, on
individual preparation. In the former case the behaviour is adaptive on the
first occurrence of the appropriate presentation; in the latter case
accommodation to circumstances is only reached after a greater or less
amount of acquired organic modification of structure, often accompanied
(as we assume) in the higher animals by acquired experience. Logically and
biologically the two classes of behaviour are clearly distinguishable: but the
analysis of complex cases of behaviour where the two factors coöperate, is
difficult and requires careful and critical study of life-history.
The foundations of the mental life are laid in the conscious experience that
accompanies those modes of behaviour, dependent entirely on racial
preparation, which may broadly be described as instinctive. In the eighth
chapter of The Origin of Species Darwin says,[160] "I will not attempt any
definition of instinct.... Every one understands what is meant, when it is
said that instinct impels the cuckoo to migrate and to lay her eggs in other

birds' nests. An action, which we ourselves require experience to enable us
to perform, when performed by an animal, more especially by a very young
one, without experience, and when performed by many individuals in the
same way, without their knowing for what purpose it is performed, is
usually said to be instinctive." And in the summary at the close of the
chapter he says,[161] "I have endeavoured briefly to show that the mental
qualities of our domestic animals vary, and that the variations are inherited.
Still more briefly I have attempted to show that instincts vary slightly in a
state of nature. No one will dispute that instincts are of the highest
importance to each animal. Therefore there is no real difficulty, under
changing conditions of life, in natural selection accumulating to any extent
slight modifications of instinct which are in any way useful. In many cases
habit or use and disuse have probably come into play."
Into the details of Darwin's treatment there is neither space nor need to
enter. There are some ambiguous passages; but it may be said that for him,
as for his followers to-day, instinctive behaviour is wholly the result of
racial preparation transmitted through organic heredity. For the performance
of the instinctive act no individual preparation under the guidance of
personal experience is necessary. It is true that Darwin quotes with approval
Huber's saying that "a little dose of judgment or reason often comes into
play, even with animals low in the scale of nature."[162] But we may fairly
interpret his meaning to be that in behaviour, which is commonly called
instinctive, some element of intelligent guidance is often combined. If this
be conceded the strictly instinctive performance (or part of the
performance) is the outcome of heredity and due to the direct transmission
of parental or ancestral aptitudes. Hence the instinctive response as such
depends entirely on how the nervous mechanism has been built up through
heredity; while intelligent behaviour, or the intelligent factor in behaviour,
depends also on how the nervous mechanism has been modified and
moulded by use during its development and concurrently with the growth of
individual experience in the customary situations of daily life. Of course it
is essential to the Darwinian thesis that what Sir E. Ray Lankester has
termed "educability," not less than instinct, is hereditary. But it is also
essential to the understanding of this thesis that the differentiae of the
hereditary factor should be clearly grasped.

For Darwin there were two modes of racial preparation, (1) natural
selection, and (2) the establishment of individually acquired habit. He
showed that instincts are subject to hereditary variation; he saw that
instincts are also subject to modification through acquisition in the course
of individual life. He believed that not only the variations but also, to some
extent, the modifications are inherited. He therefore held that some instincts
(the greater number) are due to natural selection but that others (less
numerous) are due, or partly due, to the inheritance of acquired habits. The
latter involve Lamarckian inheritance, which of late years has been the
centre of so much controversy. It is noteworthy however that Darwin laid
especial emphasis on the fact that many of the most typical and also the
most complex instincts—those of neuter insects—do not admit of such an
interpretation. "I am surprised," he says,[163] "that no one has hitherto
advanced this demonstrative case of neuter insects, against the well-known
doctrine of inherited habit, as advanced by Lamarck." None the less Darwin
admitted this doctrine as supplementary to that which was more
distinctively his own—for example in the case of the instincts of
domesticated animals. Still, even in such cases, "it may be doubted," he
says,[164] "whether any one would have thought of training a dog to point,
had not some one dog naturally shown a tendency in this line ... so that
habit and some degree of selection have probably concurred in civilising by
inheritance our dogs." But in the interpretation of the instincts of
domesticated animals, a more recently suggested hypothesis, that of organic
selection,[165] may be helpful. According to this hypothesis any intelligent
modification of behaviour which is subject to selection is probably
coincident in direction with an inherited tendency to behave in this fashion.
Hence in such behaviour there are two factors: (1) an incipient variation in
the line of such behaviour, and (2) an acquired modification by which the
behaviour is carried further along the same line. Under natural selection
those organisms in which the two factors coöperate are likely to survive.
Under artificial selection they are deliberately chosen out from among the
rest.
Organic selection has been termed a compromise between the more strictly
Darwinian and the Lamarckian principles of interpretation. But it is not in
any sense a compromise. The principle of interpretation of that which is
instinctive and hereditary is wholly Darwinian. It is true that some of the

facts of observation relied upon by Lamarckians are introduced. For
Lamarckians however the modifications which are admittedly factors in
survival, are regarded as the parents of inherited variations; for believers in
organic selection they are only the foster-parents or nurses. It is because
organic selection is the direct outcome of and a natural extension of
Darwin's cardinal thesis that some reference to it here is justifiable. The
matter may be put with the utmost brevity as follows: (1) Variations (V)
occur, some of which are in the direction of increased adaptation (+), others
in the direction of decreased adaptation (-).
(2) Acquired modifications (M) also occur. Some of these are in the
direction of increased accommodation to circumstances (+), while others
are in the direction of diminished accommodation (-). Four major
combinations are

(b) + V with - M, (c) - V with + M,
(a) + V with + M, (d) - V with - M.

Of these (d) must inevitably be eliminated while (a) are selected. The
predominant survival of (a) entails the survival of the adaptive variations
which are inherited. The contributory acquisitions (+ M) are not inherited;
but there are none the less factors in determining the survival of the
coincident variations. It is surely abundantly clear that this is Darwinism
and has no tincture of Lamarck's essential principle, the inheritance of
acquired characters.
Whether Darwin himself would have accepted this interpretation of some at
least of the evidence put forward by Lamarckians is unfortunately a matter
of conjecture. The fact remains that in his interpretation of instinct and in
allied questions he accepted the inheritance of individually acquired
modifications of behaviour and structure.
Darwin was chiefly concerned with instinct from the biological rather than
from the psychological point of view. Indeed it must be confessed that,
from the latter standpoint, his conception of instinct as a "mental faculty"
which "impels" an animal to the performance of certain actions, scarcely
affords a satisfactory basis for genetic treatment. To carry out the spirit of

Darwin's teaching it is necessary to link more closely biological and
psychological evolution. The first step towards this is to interpret the
phenomena of instinctive behaviour in terms of stimulation and response. It
may be well to take a particular case. Swimming on the part of a duckling
is, from the biological point of view, a typical example of instinctive
behaviour. Gently lower a recently hatched bird into water: coordinated
movements of the limbs follow in rhythmical sequence. The behaviour is
new to the individual though it is no doubt closely related to that of
walking, which is no less instinctive. There is a group of stimuli afforded by
the "presentation" which results from partial immersion: upon this there
follows as a complex response an application of the functional activities in
swimming; the sequence of adaptive application on the appropriate
presentation is determined by racial preparation. We know, it is true, but
little of the physiological details of what takes place in the central nervous
system; but in broad outline the nature of the organic mechanism and the
manner of its functioning may at least be provisionally conjectured in the
present state of physiological knowledge. Similarly in the case of the
pecking of newly-hatched chicks; there is a visual presentation, there is
probably a coöperating group of stimuli from the alimentary tract in need of
food, there is an adaptive application of the activities in a definite mode of
behaviour. Like data are afforded in a great number of cases of instinctive
procedure, sometimes occurring very early in life, not infrequently deferred
until the organism is more fully developed, but all of them dependent upon
racial preparation. No doubt there is some range of variation in the
behaviour, just such variation as the theory of natural selection demands.
But there can be no question that the higher animals inherit a bodily
organisation and a nervous system, the functional working of which gives
rise to those inherited modes of behaviour which are termed instinctive.
It is to be noted that the term "instinctive" is here employed in the adjectival
form as a descriptive heading under which may be grouped many and
various modes of behaviour due to racial preparation. We speak of these as
inherited; but in strictness what is transmitted through heredity is the
complex of anatomical and physiological conditions under which, in
appropriate circumstances, the organism so behaves. So far the term
"instinctive" has a restricted biological connotation in terms of behaviour.
But the connecting link between biological evolution and psychological

evolution is to be sought,—as Darwin fully realised,—in the phenomena of
instinct, broadly considered. The term "instinctive" has also a psychological
connotation. What is that connotation?
Let us take the case of the swimming duckling or the pecking chick, and fix
our attention on the first instinctive performance. Grant that just as there is,
strictly speaking, no inherited behaviour, but only the conditions which
render such behaviour under appropriate circumstances possible; so too
there is no inherited experience, but only the conditions which render such
experience possible; then the cerebral conditions in both cases are the same.
The biological behaviour-complex, including the total stimulation and the
total response with the intervening or resultant processes in the sensorium,
is accompanied by an experience-complex including the initial stimulation-
consciousness and resulting response-consciousness. In the experience-
complex are comprised data which in psychological analysis are grouped
under the headings of cognition, affective tone and conation. But the
complex is probably experienced as an unanalysed whole. If then we use
the term "instinctive" so as to comprise all congenital modes of behaviour
which contribute to experience, we are in a position to grasp the view that
the net result in consciousness constitutes what we may term the primary
tissue of experience. To the development of this experience each instinctive
act contributes. The nature and manner of organisation of this primary
tissue of experience are dependent on inherited biological aptitudes; but
they are from the outset onwards subject to secondary development
dependent on acquired aptitudes. Biological values are supplemented by
psychological values in terms of satisfaction or the reverse.
In our study of instinct we have to select some particular phase of animal
behaviour and isolate it so far as is possible from the life of which it is a
part. But the animal is a going concern, restlessly active in many ways.
Many instinctive performances, as Darwin pointed out,[166] are serial in
their nature. But the whole of active life is a serial and coordinated
business. The particular instinctive performance is only an episode in a life-
history, and every mode of behaviour is more or less closely correlated with
other modes. This coordination of behaviour is accompanied by a
correlation of the modes of primary experience. We may classify the
instinctive modes of behaviour and their accompanying modes of
instinctive experience under as many heads as may be convenient for our

purposes of interpretation, and label them instincts of self-preservation, of
pugnacity, of acquisition, the reproductive instincts, the parental instincts,
and so forth. An instinct, in this sense of the term (for example the parental
instinct), may be described as a specialised part of the primary tissue of
experience differentiated in relation to some definite biological end. Under
such an instinct will fall a large number of particular and often well-defined
modes of behaviour, each with its own peculiar mode of experience.
It is no doubt exceedingly difficult as a matter of observation and of
inference securely based thereon to distinguish what is primary from what
is in part due to secondary acquisition—a fact which Darwin fully
appreciated. Animals are educable in different degrees; but where they are
educable they begin to profit by experience from the first. Only, therefore,
on the occasion of the first instinctive act of a given type can the experience
gained be regarded as wholly primary; all subsequent performance is liable
to be in some degree, sometimes more, sometimes less, modified by the
acquired disposition which the initial behaviour engenders. But the early
stages of acquisition are always along the lines predetermined by instinctive
differentiation. It is the task of comparative psychology to distinguish the
primary tissue of experience from its secondary and acquired modifications.
We cannot follow up the matter in further detail. It must here suffice to
suggest that this conception of instinct as a primary form of experience
lends itself better to natural history treatment than Darwin's conception of
an impelling force, and that it is in line with the main trend of Darwin's
thought.
In a characteristic work,—characteristic in wealth of detail, in closeness and
fidelity of observation, in breadth of outlook, in candour and modesty,—
Darwin dealt with The Expression of the Emotions in Man and Animals. Sir
Charles Bell in his Anatomy of Expression had contended that many of
man's facial muscles had been specially created for the sole purpose of
being instrumental in the expression of his emotions. Darwin claimed that a
natural explanation, consistent with the doctrine of evolution, could in
many cases be given and would in other cases be afforded by an extension
of the principles he advocated. "No doubt," he said,[167] "as long as man and
all other animals are viewed as independent creations, an effectual stop is
put to our natural desire to investigate as far as possible the causes of
Expression. By this doctrine, anything and everything can be equally well

explained.... With mankind, some expressions ... can hardly be understood,
except on the belief that man once existed in a much lower and animal-like
condition. The community of certain expressions in distinct though allied
species ... is rendered somewhat more intelligible, if we believe in their
descent from a common progenitor. He who admits on general grounds that
the structure and habits of all animals have been gradually evolved, will
look at the whole subject of Expression in a new and interesting light."
Darwin relied on three principles of explanation. "The first of these
principles is, that movements which are serviceable in gratifying some
desire, or in relieving some sensation, if often repeated, become so habitual
that they are performed, whether or not of any service, whenever the same
desire or sensation is felt, even in a very weak degree."[168] The modes of
expression which fall under this head have become instinctive through the
hereditary transmission of acquired habit. "As far as we can judge, only a
few expressive movements are learnt by each individual; that is, were
consciously and voluntarily performed during the early years of life for
some definite object, or in imitation of others, and then became habitual.
The far greater number of the movements of expression, and all the more
important ones, are innate or inherited; and such cannot be said to depend
on the will of the individual. Nevertheless, all those included under our first
principle were at first voluntarily performed for a definite object,—namely,
to escape some danger, to relieve some distress, or to gratify some desire."
[169]

"Our second principle is that of antithesis. The habit of voluntarily
performing opposite movements under opposite impulses has become
firmly established in us by the practice of our whole lives. Hence, if certain
actions have been regularly performed, in accordance with our first
principle, under a certain frame of mind, there will be a strong and
involuntary tendency to the performance of directly opposite actions,
whether or not these are of any use, under the excitement of an opposite
frame of mind."[170] This principle of antithesis has not been widely
accepted. Nor is Darwin's own position easy to grasp.
"Our third principle," he says,[171] "is the direct action of the excited
nervous system on the body, independently of the will, and independently,
in large part, of habit. Experience shows that nerve-force is generated and

set free whenever the cerebro-spinal system is excited. The direction which
this nerve-force follows is necessarily determined by the lines of connection
between the nerve-cells, with each other and with various parts of the
body."
Lack of space prevents our following up the details of Darwin's treatment of
expression. Whether we accept or do not accept his three principles of
explanation we must regard his work as a masterpiece of descriptive
analysis, packed full of observations possessing lasting value. For a further
development of the subject it is essential that the instinctive factors in
expression should be more fully distinguished from those which are
individually acquired—a difficult task—and that the instinctive factors
should be rediscussed in the light of modern doctrines of heredity, with a
view to determining whether Lamarckian inheritance, on which Darwin so
largely relied, is necessary for an interpretation of the facts.
The whole subject as Darwin realised is very complex. Even the term
"expression" has a certain amount of ambiguity. When the emotion is in full
flood, the animal fights, flees, or faints. Is this full-tide effect to be regarded
as expression; or are we to restrict the term to the premonitory or residual
effects—the bared canine when the fighting mood is being roused, the
ruffled fur when reminiscent representations of the object inducing anger
cross the mind? Broadly considered both should be included. The activity of
premonitory expression as a means of communication was recognised by
Darwin; he might, perhaps, have emphasised it more strongly in dealing
with the lower animals. Man so largely relies on a special means of
communication, that of language, that he sometimes fails to realise that for
animals with their keen powers of perception, and dependent as they are on
such means of communication, the more strictly biological means of
expression are full of subtle suggestiveness. Many modes of expression,
otherwise useless, are signs of behaviour that may be anticipated,—signs
which stimulate the appropriate attitude of response. This would not,
however, serve to account for the utility of the organic accompaniments—
heart-affection, respiratory changes, vaso-motor effects and so forth,
together with heightened muscular tone,—on all of which Darwin lays
stress[172] under his third principle. The biological value of all this is,
however, of great importance, though Darwin was hardly in a position to
take it fully into account.

Having regard to the instinctive and hereditary factors of emotional
expression we may ask whether Darwin's third principle does not alone
suffice as an explanation. Whether we admit or reject Lamarckian
inheritance it would appear that all hereditary expression must be due to
pre-established connections within the central nervous system and to a
transmitted provision for coordinated response under the appropriate
stimulation. If this be so, Darwin's first and second principles are
subordinate and ancillary to the third, an expression, so far as it is
instinctive or heredity, being "the direct result of the constitution of the
nervous system."
Darwin accepted the emotions themselves as hereditary or acquired states
of mind and devoted his attention to their expression. But these emotions
themselves are genetic products and as such dependent on organic
conditions. It remained, therefore, for psychologists who accepted evolution
and sought to build on biological foundations to trace the genesis of these
modes of animal and human experience. The subject has been
independently developed by Professors Lange and James;[173] and some
modification of their view is regarded by many evolutionists as affording
the best explanation of the facts. We must fix our attention on the lower
emotions, such as anger or fear, and on their first occurrence in the life of
the individual organism. It is a matter of observation that if a group of
young birds which have been hatched in an incubator are frightened by an
appropriate presentation, auditory or visual, they instinctively respond in
special ways. If we speak of this response as the expression, we find that
there are many factors. There are certain visible modes of behaviour,
crouching at once, scattering and then crouching, remaining motionless, the
braced muscles sustaining an attitude of arrest, and so forth, There are also
certain visceral or organic effects, such as affections of the heart and
respiration. These can be readily observed by taking the young bird in the
hand. Other effects cannot be readily observed; vaso-motor changes,
affections of the alimentary canal, the skin and so forth. Now the essence of
the James-Lange view, as applied to these congenital effects, is that though
we are justified in speaking of them as effects of the stimulation, we are not
justified, without further evidence, in speaking of them as effects of the
emotional state. May it not rather be that the emotion as a primary mode of
experience is the concomitant of the net result of the organic situation—the

initial presentation, the instinctive mode of behaviour, the visceral
disturbances?
According to this interpretation the primary tissue of experience of the
emotional order, felt as an unanalysed complex, is generated by the
stimulation of the sensorium by afferent or incoming physiological
impulses from the special senses, from the organs concerned in the
responsive behaviour, from the viscere and vaso-motor system.
Some psychologists, however, contend that the emotional experience is
generated in the sensorium prior to, and not subsequent to, the behaviour-
response and the visceral disturbances. It is a direct and not an indirect
outcome of the presentation to the special senses. Be this as it may, there is
a growing tendency to bring into the closest possible relation, or even to
identify, instinct and emotion in their primary genesis. The central core of
all such interpretations is that instinctive behaviour and experience, its
emotional accompaniments, and its expression, are but different aspects of
the outcome of the same organic occurrences. Such emotions are, therefore,
only a distinguishable aspect of the primary tissue of experience and exhibit
a like differentiation. Here again a biological foundation is laid for a
psychological doctrine of the mental development of the individual.
The intimate relation between emotion as a psychological mode of
experience and expression as a group of organic conditions has an
important bearing on biological interpretation. The emotion, as the
psychological accompaniment of orderly disturbances in the central nervous
system, profoundly influences behaviour and often renders it more
vigourous and more effective. The utility of the emotions in the struggle for
existence can, therefore, scarcely be over-estimated. Just as keenness of
perception has survival-value; just as it is obviously subject to variation;
just as it must be enhanced under natural selection, whether individually
acquired increments are inherited or not; and just as its value lies not only
in this or that special perceptive act but in its importance for life as a whole;
so the vigourous effectiveness of activity has survival-value; it is subject to
variation; it must be enhanced under natural selection; and its importance
lies not only in particular modes of behaviour but in its value for life as a
whole. If emotion and its expression as a congenital endowment are but
different aspects of the same biological occurrence; and if this is a powerful

supplement to vigour effectiveness and persistency of behaviour, it must on
Darwin's principles be subject to natural selection.
If we include under the expression of the emotions not only the premonitory
symptoms of the initial phases of the organic and mental state, not only the
signs or conditions of half-tide emotion, but the full-tide manifestation of an
emotion which dominates the situation, we are naturally led on to the
consideration of many of the phenomena which are discussed under the
head of sexual selection. The subject is difficult and complex, and it was
treated by Darwin with all the strength he could summon to the task. It can
only be dealt with here from a special point of view—that which may serve
to illustrate the influence of certain mental factors on the course of
evolution. From this point of view too much stress can scarcely be laid on
the dominance of emotion during the period of courtship and pairing in the
more highly organised animals. It is a period of maximum vigour,
maximum activity, and, correlated with special modes of behaviour and
special organic and visceral accompaniments, a period also of maximum
emotional excitement. The combats of males, their dances and aerial
evolutions, their elaborate behaviour and display, or the flood of song in
birds, are emotional expressions which are at any rate coincident in time
with sexual periodicity. From the combat of the males there follows on
Darwin's principles the elimination of those which are deficient in bodily
vigour, deficient in special structures, offensive or protective, which
contribute to success, deficient in the emotional supplement of which
persistent and whole-hearted fighting is the expression, and deficient in
alertness and skill which are the outcome of the psychological development
of the powers of perception. Few biologists question that we have here a
mode of selection of much importance, though its influence on
psychological evolution often fails to receive its due emphasis. Mr.
Wallace[174] regards it as "a form of natural selection"; "to it," he says, "we
must impute the development of the exceptional strength, size, and activity
of the male, together with the possession of special offensive and defensive
weapons, and of all other characters which arise from the development of
these or are correlated with them." So far there is little disagreement among
the followers of Darwin—for Mr. Wallace, with fine magnanimity, has
always preferred to be ranked as such, notwithstanding his right, on which a
smaller man would have constantly insisted, to the claim of independent

originator of the doctrine of natural selection. So far with regard to sexual
selection Darwin and Mr. Wallace are agreed; so far and no farther. For
Darwin, says Mr. Wallace,[175] "has extended the principle into a totally
different field of action, which has none of that character of constancy and
of inevitable result that attaches to natural selection, including male rivalry;
for by far the larger portion of the phenomena, which he endeavours to
explain by the direct action of sexual selection, can only be so explained on
the hypothesis that the immediate agency is female choice or preference. It
is to this that he imputes the origin of all secondary sexual characters other
than weapons of offence and defence.... In this extension of sexual selection
to include the action of female choice or preference, and in the attempt to
give to that choice such wide-reaching effects, I am unable to follow him
more than a very little way."
Into the details of Mr. Wallace's criticisms it is impossible to enter here. We
cannot discuss either the mode of origin of the variations in structure which
have rendered secondary sexual characters possible or the modes of
selection other than sexual which have rendered them, within narrow limits,
specifically constant. Mendelism and mutation theories may have
something to say on the subject when these theories have been more fully
correlated with the basal principles of selection. It is noteworthy that Mr.
Wallace says:[176] "Besides the acquisition of weapons by the male for the
purpose of fighting with other males, there are some other sexual characters
which may have been produced by natural selection. Such are the various
sounds and odours which are peculiar to the male, and which serve as a call
to the female or as an indication of his presence. These are evidently a
valuable addition to the means of recognition of the two sexes, and are a
further indication, that the pairing season has arrived; and the production,
intensification, and differentiation of these sounds and odours are clearly
within the power of natural selection. The same remark will apply to the
peculiar calls of birds, and even to the singing of the males." Why the same
remark should not apply to their colours and adornments is not obvious.
What is obvious is that "means of recognition" and "indication that the
pairing season has arrived" are dependent on the perceptive powers of the
female who recognises and for whom the indication has meaning. The
hypothesis of female preference, stripped of the aesthetic surplusage which
is psychologically both unnecessary and unproven, is really only different

in degree from that which Mr. Wallace admits in principle when he says that
it is probable that the female is pleased or excited by the display.
Let us for our present purpose leave on one side and regard as sub judice
the question whether the specific details of secondary sexual characters are
the outcome of female choice. For us the question is whether certain
psychological accompaniments of the pairing situation have influenced the
course of evolution and whether these psychological accompaniments are
themselves the outcome of evolution. As a matter of observation, specially
differentiated modes of behaviour, often very elaborate, frequently
requiring highly developed skill, and apparently highly charged with
emotional tone, are the precursors of pairing. They are generally confined to
the males, whose fierce combats during the period of sexual activity are part
of the emotional manifestation. It is inconceivable that they have no
biological meaning; and it is difficult to conceive that they have any other
biological end than to evoke in the generally more passive female the
pairing impulse. They, are based on instinctive foundations ingrained in the
nervous constitution through natural (or may we not say sexual?) selection
in virtue of their profound utility. They are called into play by a specialised
presentation such as the sight or the scent of the female at, or a little in
advance of, a critical period of the physiological rhythm. There is no
necessity that the male should have any knowledge of the end to which his
strenuous activity leads up. In presence of the female there is an elaborate
application of all the energies of behaviour, just because ages of racial
preparation have made him biologically and emotionally what he is—a
functionally sexual male that must dance or sing or go through hereditary
movements of display, when the appropriate stimulation comes. Of course
after the first successful courtship his future behaviour will be in some
degree modified by his previous experience. No doubt during his first
courtship he is gaining the primary data of a peculiarly rich experience,
instinctive and emotional. But the biological foundations of the behaviour
of courtship are laid in the hereditary coordinations. It would seem that in
some cases, not indeed in all, perhaps especially in those cases in which
secondary sexual behaviour is most highly evolved,—correlative with the
ardour of the male is a certain amount of reluctance in the female. The
pairing act on her part only takes place after prolonged stimulation, for
affording which the behaviour of male courtship is the requisite

presentation. The most vigourous, defiant and mettlesome male is preferred
just because he alone affords a contributory stimulation adequate to evoke
the pairing impulse with its attendant emotional tone.
It is true that this places female preference or choice on a much lower
psychological plane than Darwin in some passages seems to contemplate
where, for example, he says that the female appreciates the display of the
male and places to her credit a taste for the beautiful. But Darwin himself
distinctly states[177] that "it is not probable that she consciously deliberates;
but she is most excited or attracted by the most beautiful, or melodious, or
gallant males." The view here put forward, which has been developed by
Prof. Groos,[178] therefore seems to have Darwin's own sanction. The
phenomena are not only biological; there are psychological elements as
well. One can hardly suppose that the female is unconscious of the male's
presence; the final yielding must surely be accompanied by heightened
emotional tone. Whether we call it choice or not is merely a matter of
definition of terms. The behaviour is in part determined by supplementary
psychological values. Prof. Groos regards the coyness of females as "a most
efficient means of preventing the too early and too frequent yielding to the
sexual impulse."[179] Be that as it may, it is, in any case, if we grant the
facts, a means through which male sexual behaviour with all its biological
and psychological implications, is raised to a level otherwise perhaps
unattainable by natural means, while in the female it affords opportunities
for the development in the individual and evolution in the race of what we
may follow Darwin in calling appreciation, if we empty this word of the
aesthetic implications which have gathered round it in the mental life of
man.
Regarded from this standpoint of sexual selection, broadly considered, has
probably been of great importance. The psychological accompaniments of
the pairing situation have profoundly influenced the course of biological
evolution and are themselves the outcome of that evolution.
Darwin makes only passing reference to those modes of behaviour in
animals which go by the name of play. "Nothing," he says,[180] "is more
common than for animals to take pleasure in practising whatever instinct
they follow at other times for some real good." This is one of the very
numerous cases in which a hint of the master has served to stimulate

research in his disciples. It was left to Prof. Groos to develop this subject on
evolutionary lines and to elaborate in a masterly manner Darwin's
suggestion. "The utility of play," he says,[181] "is incalculable. This utility
consists in the practice and exercise it affords for some of the more
important duties of life,"—that is to say, for the performance of activities
which will in adult life be essential to survival. He urges[182] that "the play
of young animals has its origin in the fact that certain very important
instincts appear at a time when the animal does not seriously need them." It
is, however, questionable whether any instincts appear at a time when they
are not needed. And it is questionable whether the instinctive and emotional
attitude of the play-fight, to take one example, can be identified with those
which accompany fighting in earnest, though no doubt they are closely
related and have some common factors. It is probable that play, as
preparatory behaviour, differs in biological detail (as it almost certainly
does in emotional attributes) from the earnest of after-life and that it has
been evolved through differentiation and integration of the primary tissue of
experience, as a preparation through which certain essential modes of skill
may be acquired—those animals in which the preparatory play-propensity
was not inherited in due force and requisite amount being subsequently
eliminated in the struggle for existence. In any case there is little question
that Prof. Groos is right in basing the play-propensity on instinctive
foundations.[183] None the less, as he contends, the essential biological
value of play is that it is a means of training the educable nerve-tissue, of
developing that part of the brain which is modified by experience and
which thus acquires new characters, of elaborating the secondary tissue of
experience on the predetermined lines of instinctive differentiation and thus
furthering the psychological activities which are included under the
comprehensive term "intelligent."
In The Descent of Man Darwin dealt at some length with intelligence and
the higher mental faculties.[184] His object, he says, is to show that there is
no fundamental difference between man and the higher mammals in their
mental faculties; that these faculties are variable and the variations tend to
be inherited; and that under natural selection beneficial variations of all
kinds will have been preserved and injurious ones eliminated.
Darwin was too good an observer and too honest a man to minimise the
"enormous difference" between the level of mental attainment of civilised

man and that reached by any animal. His contention was that the difference,
great as it is, is one of degree and not of kind. He realised that, in the
development of the mental faculties of man, new factors in evolution have
supervened—factors which play but a subordinate and subsidiary part in
animal intelligence. Intercommunication by means of language, approbation
and blame, and all that arises out of reflective thought, are but
foreshadowed in the mental life of animals. Still he contends that these may
be explained on the doctrine of evolution. He urges[185] "that man is
variable in body and mind; and that the variations are induced, either
directly or indirectly, by the same general causes, and obey the same
general laws, as with the lower animals." He correlates mental development
with the evolution of the brain.[186] "As the various mental faculties
gradually developed themselves, the brain would almost certainly become
larger. No one, I presume, doubts that the large proportion which the size of
man's brain bears to his body, compared to the same proportion in the
gorilla or orang, is closely connected with his higher mental powers." "With
respect to the lower animals," he says,[187] "M. E. Lartet,[188] by comparing
the crania of tertiary and recent mammals belonging to the same groups, has
come to the remarkable conclusion that the brain is generally larger and the
convolutions are more complex in the more recent form."
Sir E. Ray Lankester has sought to express in the simplest terms the
implications of the increase in size of the cerebrum. "In what," he asks,
"does the advantage of a larger cerebral mass consist?" "Man," he replies,
"is born with fewer ready-made tricks of the nerve-centres—these
performances of an inherited nervous mechanism so often called by the ill-
defined term 'instincts'—than are the monkeys or any other animal.
Correlated with the absence of inherited ready-made mechanism, man has a
greater capacity of developing in the course of his individual growth similar
nervous mechanisms (similar to but not identical with those of 'instinct')
than any other animal.... The power of being educated—'educability' as we
may term it—is what man possesses in excess as compared with the apes. I
think we are justified in forming the hypothesis that it is this 'educability'
which is the correlative of the increased size of the cerebrum." There has
been natural selection of the more educable animals, for "the character
which we describe as 'educability' can be transmitted, it is a congenital
character. But the results of education can not be transmitted. In each

generation they have to be acquired afresh, and with increased 'educability'
they are more readily acquired and a larger variety of them.... The fact is
that there is no community between the mechanisms of instinct and the
mechanisms of intelligence, and that the latter are later in the history of the
evolution of the brain than the former and can only develop in proportion as
the former become feeble and defective."[189]
In this statement we have a good example of the further development of
views which Darwin foreshadowed but did not thoroughly work out. It
states the biological case clearly and tersely. Plasticity of behaviour in
special accommodation to special circumstances is of survival value; it
depends upon acquired characters; it is correlated with increase in size and
complexity of the cerebrum; under natural selection therefore the larger and
more complex cerebrum as the organ of plastic behaviour has been the
outcome of natural selection. We have thus the biological foundations for a
further development of genetic psychology.
There are diversities of opinion, as Darwin showed, with regard to the range
of instinct in man and the higher animals as contrasted with lower types.
Darwin himself said[190] that "Man, perhaps, has somewhat fewer instincts
than those possessed by the animals which come next to him in the series."
On the other hand, Prof. Wm. James says[191] that man is probably the
animal with most instincts. The true position is that man and the higher
animals have fewer complete and self-sufficing instincts than those which
stand lower in the scale of mental evolution, but that they have an equally
large or perhaps larger mass of instinctive raw material which may furnish
the stuff to be elaborated by intelligent processes. There is, perhaps, a
greater abundance of the primary tissue of experience to be refashioned and
integrated by secondary modification; there is probably the same
differentiation in relation to the determining biological ends, but there is at
the outset less differentiation of the particular and specific modes of
behaviour. The specialised instinctive performances and their concomitant
experience-complexes are at the outset more indefinite. Only through
acquired connections, correlated with experience, do they become definitely
organised.
The full working-out of the delicate and subtle relationship of instinct and
educability—that is, of the hereditary and the acquired factors in the mental

life—is the task which lies before genetic and comparative psychology.
They interact throughout the whole of life, and their interactions are very
complex. No one can read the chapters of The Descent of Man which
Darwin devotes to a consideration of the mental characters of man and
animals without noticing, on the one hand, how sedulous he is in his search
for hereditary foundations, and, on the other hand, how fully he realises the
importance of acquired habits of mind. The fact that educability itself has
innate tendencies—is in fact a partially differentiated educability—renders
the unravelling of the factors of mental progress all the more difficult.
In his comparison of the mental powers of men and animals it was essential
that Darwin should lay stress on points of similarity rather than on points of
difference. Seeking to establish a doctrine of evolution, with its basal
concept of continuity of process and community of character, he was bound
to render clear and to emphasise the contention that the difference in mind
between man and the higher animals, great as it is, is one of degree and not
of kind. To this end Darwin not only recorded a large number of valuable
observations of his own, and collected a considerable body of information
from reliable sources, he presented the whole subject in a new light and
showed that a natural history of mind might be written and that this method
of study offered a wide and rich field for investigation. Of course those who
regarded the study of mind only as a branch of metaphysics smiled at the
philosophical ineptitude of the mere man of science. But the investigation,
on natural history lines, has been prosecuted with a large measure of
success. Much indeed still remains to be done; for special training is
required, and the workers are still few. Promise for the future is however
afforded by the fact that investigation is prosecuted on experimental lines
and that something like organised methods of research are taking form.
There is now but little reliance on casual observations recorded by those
who have not undergone the necessary discipline in these methods. There is
also some change of emphasis in formulating conclusions. Now that the
general evolutionary thesis is fully and freely accepted by those who carry
on such researches, more stress is laid on the differentiation of the stages of
evolutionary advance than on the fact of their underlying community of
nature. The conceptual intelligence which is especially characteristic of the
higher mental procedure of man is more firmly distinguished from the
perceptual intelligence which he shares with the lower animals—

distinguished now as a higher product of evolution, no longer as differing in
origin or different in kind. Some progress has been made, on the one hand
in rendering an account of intelligent profiting by experience under the
guidance of pleasure and pain in the perceptual field, on lines
predetermined by instinctive differentiation for biological ends, and on the
other hand in elucidating the method of conceptual thought employed, for
example, by the investigator himself in interpreting the perceptual
experience of the lower animals.
Thus there is a growing tendency to realise more fully that there are two
orders of educability—first an educability of the perceptual intelligence
based on the biological foundation of instinct, and secondly an educability
of the conceptual intelligence which refashions and rearranges the data
afforded by previous inheritance and acquisition. It is in relation to this
second and higher order of educability that the cerebrum of man shows so
large an increase of mass and a yet larger increase of effective surface
through its rich convolutions. It is through educability of this order that the
human child is brought intellectually and affectively into touch with the
ideal constructions by means of which man has endeavoured, with more or
less success, to reach an interpretation of nature, and to guide the course of
the further evolution of his race—ideal constructions which form part of
man's environment.
It formed no part of Darwin's purpose to consider, save in broad outline, the
methods, or to discuss in any fulness of detail the results of the process by
which a differentiation of the mental faculties of man from those of the
lower animals has been brought about—a differentiation the existence of
which he again and again acknowledges. His purpose was rather to show
that, notwithstanding this differentiation, there is basal community in kind.
This must be remembered in considering his treatment of the biological
foundations on which man's systems of ethics are built. He definitely stated
that he approached the subject "exclusively from the side of natural
history."[192] His general conclusion is that the moral sense is fundamentally
identical with the social instincts, which have been developed for the good
of the community; and he suggests that the concept which thus enables us to
interpret the biological ground-plan of morals also enables us to frame a
rational ideal of the moral end. "As the social instincts," he says,[193] "both
of man and the lower animals have no doubt been developed by nearly the

same steps, it would be advisable, if found practicable, to use the same
definition in both cases, and to take as the standard of morality, the general
good or welfare of the community, rather than the general happiness." But
the kind of community for the good of which the social instincts of animals
and primitive men were biologically developed may be different from that
which is the product of civilisation, as Darwin no doubt realised. Darwin's
contention was that conscience is a social instinct and has been evolved
because it is useful to the tribe in the struggle for existence against other
tribes. One the other hand J. S. Mill urged that the moral feelings are not
innate but acquired, and Bain held the same view, believing that the moral
sense is acquired by each individual during his life-time. Darwin, who
notes[194] their opinion with his usual candour, adds that "on the general
theory of evolution this is at least extremely improbable." It is impossible to
enter into the question here: much turns on the exact connotation of the
terms "conscience" and "moral sense," and on the meaning we attach to the
statement that the moral sense is fundamentally identical with the social
instincts.
Presumably the majority of those who approach the subjects discussed in
the third, fourth, and fifth chapters of The Descent of Man in the full
conviction that mental phenomena, not less than organic phenomena, have a
natural genesis, would, without hesitation, admit that the intellectual and
moral systems of civilised man are ideal constructions, the products of
conceptual thought, and that as such they are, in their developed form,
acquired. The moral sentiments are the emotional analogues of highly
developed concepts. This does not however imply that they are outside the
range of natural history treatment. Even though it may be desirable to
differentiate the moral conduct of men from the social behaviour of animals
(to which some such term as "pre-moral" or "quasi-moral" may be applied),
still the fact remains that, as Darwin showed, there is abundant evidence of
the occurrence of such social behaviour—social behaviour which, even
granted that it is in large part intelligently acquired, and is itself so far a
product of educability, is of survival value. It makes for that integration
without which no social group could hold together and escape elimination.
Furthermore, even if we grant that such behaviour is intelligently acquired,
that is to say arises through the modification of hereditary instincts and
emotions, the fact remains that only through these instinctive and emotional

data is afforded the primary tissue of the experience which is susceptible of
such modification.
Darwin sought to show, and succeeded in showing, that for the intellectual
and moral life there are instinctive foundations which a biological treatment
alone can disclose. It is true that he did not in all cases analytically
distinguish the foundations from the superstructure. Even to-day we are
scarcely in a position to do so adequately. But his treatment was of great
value in giving an impetus to further research. This value indeed can
scarcely be over-estimated. And when the natural history of the mental
operations shall have been written, the cardinal fact will stand forth, that the
instinctive and emotional foundations are the outcome of biological
evolution and have been ingrained in the race through natural selection. We
shall more clearly realise that educability itself is a product of natural
selection, though the specific results acquired through cerebral
modifications are not transmitted through heredity. It will, perhaps, also be
realised that the instinctive foundations of social behaviour are, for us,
somewhat out of date and have undergone but little change throughout the
progress of civilisation, because natural selection has long since ceased to
be the dominant factor in human progress. The history of human progress
has been mainly the history of man's higher educability, the products of
which he has projected on to his environment. This educability remains on
the average what it was a dozen generations ago; but the thought-woven
tapestry of his surroundings is refashioned and improved by each
succeeding generation. Few men have in greater measure enriched the
thought-environment with which it is the aim of education to bring educable
human beings into vital contact, than has Charles Darwin. His special field
of work was the wide province of biology; but he did much to help us to
realise that mental factors have contributed to organic evolution and that in
man, the highest product of Evolution, they have reached a position of
unquestioned supremacy.

FOOTNOTES:
[153] Origin of Species (6th edit.), p. 205.
[154] Descent of man (2nd edit. 1888), Vol. I. p. 99; Popular edit. p. 99.

[155] Ibid. p. 99.
[156] The Expression of the Emotions (2nd edit.), p. 32.
[157] Descent of Man, Vol. II. p. 435.
[158] Ibid. 437, 438.
[159] Origin of Species (6th edit.), p. 205.
[160] Origin of Species (6th edit.), p. 205.
[161] Ibid. p. 233.
[162] Ibid. p. 205.
[163] Origin of Species (6th edit.), p. 233.
[164] Origin of Species, pp. 210, 211.
[165] Independently suggested, on somewhat different lines, by Profs. J. Mark
Baldwin, Henry F. Osborn and the writer.
[166] Origin of Species (6th edit.), p. 206.
[167] Expression of the Emotions, p. 13. The passage is here somewhat
condensed.
[168] Ibid. p. 368.
[169] Expression of the Emotions, pp. 373, 374.
[170] Ibid. p. 368.
[171] Ibid. p. 369.
[172] Expression of the Emotions, pp. 65 ff.
[173] Cf. William James, Principles of Psychology, Vol. ii. Chap. xxv, New
York, 1890.
[174] Darwinism, pp. 282, 283, London, 1889.
[175] Ibid. p. 283.
[176] Darwinism, pp. 283, 284.
[177] Descent of Man (2nd edit.), Vol. ii. pp. 136, 137; (Popular edit.), pp. 642,
643.
[178] The Play of Animals, p. 244, London, 1898.
[179] Ibid. p. 283.
[180] Descent of Man, Vol. ii. p. 60; (Popular edit.), p. 566.
[181] The Play of Animals, p. 76.
[182] Ibid. p. 75.

[183] The Play of Animals p. 24.
[184] Descent of Man (1st edit.), Chaps. ii, iii, v; (2nd edit.), Chaps. iii, iv, v.
[185] Descent of Man, Vol. i. pp. 70, 71; (Popular edit.), pp. 70, 71.
[186] Ibid. p. 81.
[187] Ibid. (Popular edit.), p. 82.
[188] Comptes Rendus des Sciences, June 1, 1868.
[189] Nature, Vol. lxi. pp. 624, 625 (1900).
[190] Descent of Man, Vol. i. p. 100.
[191] Principles of Psychology, Vol. ii. p. 289.
[192] Descent of Man, Vol. i. p. 149.
[193] Descent of Man, p. 185.
[194] Ibid. p. 150 (footnote).

VII

THE INFLUENCE OF THE CONCEPTION OF
EVOLUTION ON MODERN PHILOSOPHY
By H. Höffding

Professor of Philosophy in the University of Copenhagen

I

It is difficult to draw a sharp line between philosophy and natural science.
The naturalist who introduces a new principle, or demonstrates a fact which
throws a new light on existence, not only renders an important service to
philosophy but is himself a philosopher in the broader sense of the word.
The aim of philosophy in the stricter sense is to attain points of view from
which the fundamental phenomena and the principles of the special
sciences can be seen in their relative importance and connection. But
philosophy in this stricter sense has always been influenced by philosophy
in the broader sense. Greek philosophy came under the influence of logic
and mathematics, modern philosophy under the influence of natural science.
The name of Charles Darwin stands with those of Galileo, Newton, and
Robert Mayer—names which denote new problems and great alterations in
our conception of the universe.
First of all we must lay stress on Darwin's own personality. His deep love of
truth, his indefatigable inquiry, his wide horizon, and his steady self-
criticism make him a scientific model, even if his results and theories
should eventually come to possess mainly an historical interest. In the
intellectual domain the primary object is to reach high summits from which
wide surveys are possible, to reach them toiling honestly upwards by the
way of experience, and then not to turn dizzy when a summit is gained.
Darwinians have sometimes turned dizzy, but Darwin never. He saw from
the first the great importance of his hypothesis, not only because of its
solution of the old problem as to the value of the concept of species, not
only because of the grand picture of natural evolution which it unrolls, but

also because of the life and inspiration its method would impart to the study
of comparative anatomy, of instinct and of heredity, and finally because of
the influence it would exert on the whole conception of existence. He wrote
in his note-book in the year 1837: "My theory would give zest to recent and
fossil comparative anatomy; it would lead to the study of instinct, heredity,
and mind-heredity, whole [of] metaphysics."[195]
We can distinguish four main points in which Darwin's investigations
possess philosophical importance.
The evolution hypothesis is much older than Darwin; it is, indeed, one of
the oldest guessings of human thought. In the eighteenth century is was put
forward by Diderot and Lamettrie and suggested by Kant (1786). As we
shall see later, it was held also by several philosophers in the first half of the
nineteenth century. In his preface to The Origin of Species, Darwin
mentions the naturalists who were his forerunners. But he has set forth the
hypothesis of evolution in so energetic and thorough a manner that it
perforce attracts the attention of all thoughtful men in a much higher degree
than it did before the publication of the Origin.
And further, the importance of his teaching rests on the fact that he, much
more than his predecessors, even than Lamarck, sought a foundation for his
hypothesis in definite facts. Modern science began by demanding—with
Kepler and Newton—evidence of varae causae; this demand Darwin
industriously set himself to satisfy—hence the wealth of material which he
collected by his observations and his experiments. He not only revived an
old hypothesis, but he saw the necessity of verifying it by facts. Whether
the special cause on which he founded the explanation of the origin of
species—Natural Selection—is sufficient, is now a subject of discussion.
He himself had some doubt in regard to this question, and the criticisms
which are directed against his hypothesis hit Darwinism rather than Darwin.
In his indefatigable search for empirical evidence he is a model even for his
antagonists: he has compelled them to approach the problems of life along
other lines than those which were formerly followed.
Whether the special cause to which Darwin appealed is sufficient or not, at
least to it is probably due the greater part of the influence which he has
exerted on the general trend of thought. "Struggle for existence" and

"natural selection" are principles which have been applied, more or less, in
every department of thought. Recent research, it is true, has discovered
greater empirical discontinuity—leaps, "mutations"—whereas Darwin
believed in the importance of small variations slowly accumulated. It has
also been shown by the experimental method, which in recent biological
work has succeeded Darwin's more historical method, that types once
constituted possess great permanence, the fluctuations being restricted
within clearly defined boundaries. The problem has become more precise,
both as to variation and as to heredity. The inner conditions of life have in
both respects shown a greater independence than Darwin had supposed in
his theory, though he always admitted that the cause of variation was to him
a great enigma, "a most perplexing problem," and that the struggle for life
could only occur where variation existed. But, at any rate, it was of the
greatest importance that Darwin gave a living impression of the struggle for
life which is everywhere going on, and to which even the highest forms of
existence must be amenable. The philosophical importance of these ideas
does not stand or fall with the answer to the question, whether natural
selection is a sufficient explanation of the origin of species or not it has an
independent, positive value for everyone who will observe life and reality
with an unbiased mind.
In accentuating the struggle for life Darwin stands as a characteristically
English thinker: he continues a train of ideas which Hobbes and Malthus
had already begun. Moreover in his critical views as to the conception of
species he had English forerunners; in the middle ages Occam and Duns
Scotus, in the eighteenth century Berkeley and Hume. In his moral
philosophy, as we shall see later, he is an adherent of the school which is
represented by Hutcheson, Home and Adam Smith. Because he is no
philosopher in the stricter sense of the term, it is of great interest to see that
his attitude of mind is that of the great thinkers of his nation.
In considering Darwin's influence on philosophy we will begin with an
examination of the attitude of philosophy to the conception of evolution at
the time when The Origin of Species appeared. We will then examine the
effects which the theory of evolution, and especially the idea of the struggle
for life, has had, and naturally must have, on the discussion of philosophical
problems.

II
When The Origin of Species appeared fifty years ago Romantic speculation,
Schelling's and Hegel's philosophy, still reigned on the continent, while in
England Positivism, the philosophy of Comte and Stuart Mill, represented
the most important trend of thought. German speculation had much to say
on evolution, it even pretended to be a philosophy of evolution. But then the
word "evolution" was to be taken in an ideal, not in a real, sense. To
speculative thought the forms and types of nature formed a system of ideas,
within which any form could lead us by continuous transitions to any other.
It was a classificatory system which was regarded as a divine world of
thought or images, within which metamorphoses could go on—a condition
comparable with that in the mind of the poet when one image follows
another with imperceptible changes. Goethe's ideas of evolution, as
expressed in his Metamorphosen der Pflanzen und der Thiere, belong to
this category; it is, therefore, incorrect to call him a forerunner of Darwin.
Schelling and Hegel held the same idea; Hegel expressly rejected the
conception of a real evolution in time as coarse and materialistic. "Nature,"
he says, "is to be considered as a system of stages, the one necessarily
arising from the other, and being the nearest truth of that from which it
proceeds; but not in such a way that the one is naturally generated by the
other; on the contrary [their connection lies] in the inner idea which is the
ground of nature. The metamorphosis can be ascribed only to the notion as
such, because it alone is evolution.... It has been a clumsy idea in the older
as well as in the newer philosophy of nature, to regard the transformation
and the transition from one natural form and sphere to a higher as an
outward and actual production."[196]
The only one of the philosophers of Romanticism who believed in a real,
historical evolution, a real production of new species, was Oken.[197]
Danish philosophers, such as Treschow (1812) and Sibbern (1846), have
also broached the idea of an historical evolution of all living beings from
the lowest to the highest. Schopenhauer's philosophy has a more realistic
character than that of Schelling's and Hegel's, his diametrical opposites,
although he also belongs to the romantic school of thought. His
philosophical and psychological views were greatly influenced by French
naturalists and philosophers, especially by Cabanis and Lamarck. He
praises the "ever memorable Lamarck," because he laid so much stress on

the "will to live." But he repudiates as a "wonderful error" the idea that the
organs of animals should have reached their present perfection through a
development in time, during the course of innumerable generations. It was,
he said, a consequence of the low standard of contemporary French
philosophy, that Lamarck came to the idea of the construction of living
beings in time through succession![198]
The positivistic stream of thought was not more in favour of a real
evolution than was the Romantic school. Its aim was to adhere to positive
facts: it looked with suspicion on far-reaching speculation. Comte laid great
stress on the discontinuity found between the different kingdoms of nature,
as well as within each single kingdom. As he regarded as unscientific every
attempt to reduce the number of physical forces, so he rejected entirely the
hypothesis of Lamarck concerning the evolution of species; the idea of
species would in his eyes absolutely lose its importance if a transition from
species to species under the influence of conditions of life were admitted.
His disciples (Littré, Robin) continued to direct against Darwin the
polemics which their master had employed against Lamarck. Stuart Mill,
who, in the theory of knowledge, represented the empirical or positivistic
movement in philosophy—like his English forerunners from Locke to
Hume—founded his theory of knowledge and morals on the experience of
the single individual. He sympathised with the theory of the original
likeness of all individuals and derived their differences, on which he
practically and theoretically laid much stress, from the influence both of
experience and education, and, generally, of physical and social causes. He
admitted an individual evolution, and, in the human species, an evolution
based on social progress; but no physiological evolution of species. He was
afraid that the hypothesis of heredity would carry us back to the old theory
of "innate" ideas.
Darwin was more empirical than Comte and Mill; experience disclosed to
him a deeper continuity than they could find; closer than before the nature
and fate of the single individual were shown to be interwoven in the great
web binding the life of the species with nature as a whole. And the
continuity which so many idealistic philosophers could find only in the
world of thought, he showed to be present in the world of reality.
III

Darwin's energetic renewal of the old idea of evolution has its chief
importance in strengthening the conviction of this real continuity in the
world, of continuity in the series of form and events. It was a great support
for all those who were prepared to base their conception of life on scientific
grounds. Together with the recently discovered law of the conservation of
energy, it helped to produce the great realistic movement which
characterises the last third of the nineteenth century. After the decline of the
Romantic movement people wished to have firmer ground under their feet
and reality now asserted itself in a more emphatic manner than in the period
of Romanticism. It was easy for Hegel to proclaim that "the real" was "the
rational," and that "the rational" was "the real": reality itself existed for him
only in the interpretation of ideal reason, and if there was anything which
could not be merged in the higher unity of thought, then it was only an
example of the "impotence of nature to hold to the idea." But now concepts
are to be founded on nature and not on any system of categories too
confidently deduced à priori. The new devotion to nature had its
recompense in itself, because the new points of view made us see that
nature could indeed "hold to ideas," though perhaps not to those which we
had cogitated beforehand.
A most important question for philosophers to answer was whether the new
views were compatible with an idealistic conception of life and existence.
Some proclaimed that we have now no need of any philosophy beyond the
principles of the conservation of matter and energy and the principle of
natural evolution: existence should and could be definitely and completely
explained by the laws of material nature. But abler thinkers saw that the
thing was not so simple. They were prepared to give the new views their
just place and to examine what alterations the old views must undergo in
order to be brought into harmony with the new data.
The realistic character of Darwin's theory was shown not only in the idea of
natural continuity, but also, and not least, in the idea of the cause whereby
organic life advances step by step. This idea—the idea of the struggle for
life—implied that nothing could persist, if it had no power to maintain itself
under the given conditions. Inner value alone does not decide. Idealism was
here put to its hardest trial. In continuous evolution it could perhaps still
find an analogy to the inner evolution of ideas in the mind; but in the
demand for power in order to struggle with outward conditions Realism

seemed to announce itself in its most brutal form. Every form of Idealism
had to ask itself seriously how it was going to "struggle for life" with this
new Realism.
We will now give a short account of the position which leading thinkers in
different countries have taken up in regard to this question.
I. Herbert Spencer was the philosopher whose mind was best prepared by
his own previous thinking to admit the theory of Darwin to a place in his
conception of the world. His criticism of the arguments which had been put
forward against the hypothesis of Lamarck, showed that Spencer, as a
young man, was an adherent to the evolution idea. In his Social Statics
(1850) he applied this idea to human life and moral civilisation. In 1852 he
wrote an essay on The Development Hypothesis, in which he definitely
stated his belief that the differentiation of species, like the differentiation
within a single organism, was the result of development. In the first edition
of his Psychology (1855) he took a step which put him in opposition to the
older English school (from Locke to Mill): he acknowledged "innate ideas"
so far as to admit the tendency of acquired habits to be inherited in the
course of generations, so that the nature and functions of the individual are
only to be understood through its connection with the life of the species. In
1857, in his essay on Progress, he propounded the law of differentiation as
a general law of evolution, verified by examples from all regions of
experience, the evolution of species being only one of these examples. On
the effect which the appearance of The Origin of Species had on his mind he
writes in his Autobiography: "Up to that time ... I held that the sole cause of
organic evolution is the inheritance of functionally-produced modifications.
The Origin of Species made it clear to me that I was wrong, and that the
larger part of the facts cannot be due to any such cause.... To have the
theory of organic evolution justified was of course to get further support for
that theory of evolution at large with which ... all my conceptions were
bound up."[199] Instead of the metaphorical expression "natural selection,"
Spencer introduced the term "survival of the fittest," which found favour
with Darwin as well as with Wallace.
In working out his ideas of evolution, Spencer found that differentiation
was not the only form of evolution. In its simplest form evolution is mainly
a concentration, previously scattered elements being integrated and losing

independent movement. Differentiation is only forthcoming when minor
wholes arise within a greater whole. And the highest form of evolution is
reached when there is a harmony between concentration and differentiation,
a harmony which Spencer calls equilibration and which he defines as a
moving equilibrium. At the same time this definition enables him to
illustrate the expression "survival of the fittest." "Every living organism
exhibits such a moving equilibrium—a balanced set of functions
constituting its life; and the overthrow of this balanced set of functions or
moving equilibrium is what we call death. Some individuals in a species are
so constituted that their moving equilibria are less easily overthrown than
those of other individuals; and these are the fittest which survive, or, in Mr.
Darwin's language, they are the select which nature preserves."[200] Not
only in the domain of organic life, but in all domains, the summit of
evolution is, according to Spencer, characterised by such a harmony—by a
moving equilibrium.
Spencer's analysis of the concept of evolution, based on a great variety of
examples, has made this concept clearer and more definite than before. It
contains the three elements; integration, differentiation and equilibration. It
is true that a concept which is to be valid for all domains of experience must
have an abstract character, and between the several domains there is, strictly
speaking, only a relation of analogy. So there is only analogy between
psychical and physical evolution. But this is no serious objection, because
general concepts do not express more than analogies between the
phenomena which they represent. Spencer takes his leading forms from the
material world in defining evolution (in the simplest form) as integration of
matter and dissipation of movement; but as he—not always quite
consistently[201]—assumed a correspondence of mind and matter, he could
very well give these terms an indirect importance for psychical evolution.
Spencer has always, in my opinion with full right, repudiated the ascription
of materialism. He is no more a materialist than Spinoza. In his Principles
of Psychology (§ 63) he expressed himself very clearly: "Though it seems
easier to translate so-called matter into so-called spirit, than to translate so-
called spirit into so-called matter—which latter is indeed wholly impossible
—yet no translation can carry us beyond our symbols." These words lead us
naturally to a group of thinkers whose starting-point was psychical
evolution. But we have still one aspect of Spencer's philosophy to mention.

Spencer founded his "laws of evolution" on an inductive basis, but he was
convinced that they could be deduced from the law of the conservation of
energy. Such a deduction is, perhaps, possible for the more elementary
forms of evolution, integration and differentiation; but it is not possible for
the highest form, the equilibration, which is a harmony of integration and
differentiation. Spencer can no more deduce the necessity for the eventual
appearance of "moving equilibria" of harmonious totalities than Hegel
could guarantee the "higher unities" in which all contradictions should be
reconciled. In Spencer's hands the theory of evolution acquired a more
decidedly optimistic character than in Darwin's; but I shall deal later with
the relation of Darwin's hypothesis to the opposition of optimism and
pessimism.
II. While the starting-point of Spencer was biological or cosmological,
psychical evolution being conceived as in analogy with physical, a group of
eminent thinkers—in Germany Wundt, in France Fouillée, in Italy Ardigò—
took, each in his own manner, their starting-point in psychical evolution as
an original fact and as a type of all evolution, the hypothesis of Darwin
coming in as a corroboration and as a special example. They maintain the
continuity of evolution; they find this character most prominent in psychical
evolution, and this is for them a motive to demand a corresponding
continuity in the material, especially in the organic domain.
To Wundt and Fouillée the concept of will is prominent. They see the type
of all evolution in the transformation of the life of will from blind impulse
to conscious choice; the theories of Lamarck and Darwin are used to
support the view that there is in nature a tendency to evolution in steady
reciprocity with external conditions. The struggle for life is here only a
secondary fact. Its apparent prominence is explained by the circumstance
that the influence of external conditions is easily made out, while inner
conditions can be verified only through their effects. For Ardigò the
evolution of thought was the starting-point and the type: in the evolution of
a scientific hypothesis we see a progress from the indefinite (indistinto) to
the definite (distinto), and this is a characteristic of all evolution, as Ardigò
has pointed out in a series of works. The opposition between indistinto and
distinto corresponds to Spencer's opposition between homogeneity and
heterogeneity. The hypothesis of the origin of differences of species from
more simple forms is a special example of the general law of evolution.

In the views of Wundt and Fouillée we find the fundamental idea of
idealism psychical phenomena as expressions of the innermost nature of
existence. They differ from the older Idealism in the great stress which they
lay on evolution as a real, historical process which is going on through
steady conflict with external conditions. The Romantic dread of reality is
broken. It is beyond doubt that Darwin's emphasis on the struggle for life as
a necessary condition of evolution has been a very important factor in
carrying philosophy back to reality from the heaven of pure ideas. The
philosophy of Ardigò, on the other side, appears more as a continuation and
deepening of positivism, though the Italian thinker arrived at his point of
view independently of French-English positivism. The idea of continuous
evolution is here maintained in opposition to Comte's and Mill's philosophy
of discontinuity. From Wundt and Fouillée Ardigò differs in conceiving
psychical evolution not as an immediate revelation of the innermost nature
of existence, but only as a single, though the most accessible example, of
evolution.
III. To the French philosophers Boutroux and Bergson, evolution proper is
continuous and qualitative, while outer experience and physical science
give us fragments only, sporadic processes and mechanical combinations.
To Bergson, in his recent work L'Evolution Créatrice, evolution consists in
an élan de vie which to our fragmentary observation and analytic reflexion
appears as broken into a manifold of elements and processes. The concept
of matter in its scientific form is the result of this breaking asunder,
essential for all scientific reflexion. In these conceptions the strongest
opposition between inner and outer conditions of evolution is expressed: in
the domain of internal conditions spontaneous development of qualitative
forms—in the domain of external conditions discontinuity and mechanical
combination.
We see, then, that the theory of evolution has influenced philosophy in a
variety of forms. It has made idealistic thinkers revise their relation to the
real world; it has led positivistic thinkers to find a closer connection
between the facts on which they based their views; it has made us all open
our eyes for new possibilities to arise through the prima facie inexplicable
"spontaneous" variations which are the condition of all evolution. This last
point is one of peculiar interest. Deeper than speculative philosophy and
mechanical science saw in the days of their triumph, we catch sight of new

streams, whose sources and laws we have still to discover. Most sharply
does this appear in the theory of mutation, which is only a stronger
accentuation of a main point in Darwinism. It is interesting to see that an
analogous problem comes into the foreground in physics through the
discovery of radioactive phenomena, and in psychology through the
assumption of psychical new formations (as held by Boutroux, William
James and Bergson). From this side, Darwin's ideas, as well as the
analogous ideas in other domains, incite us to renewed examination of our
first principles, their rationality and their value. On the other hand, his
theory of the struggle for existence challenges us to examine the conditions
and discuss the outlook as to the persistence of human life and society and
of the values that belong to them. It is not enough to hope (or fear?) the
rising of new forms; we have also to investigate the possibility of upholding
the forms and ideals which have hitherto been the bases of human life.
Darwin has here given his age the most earnest and most impressive lesson.
This side of Darwin's theory is of peculiar interest to some special
philosophical problems to which I now pass.
IV
Among philosophical problems the problem of knowledge has in the last
century occupied a foremost place. It is natural, then, to ask how Darwin
and the hypothesis whose most eminent representative he is, stand to this
problem.
Darwin started an hypothesis. But every hypothesis is won by inference
from certain presuppositions, and every inference is based on the general
principles of human thought. The evolution hypothesis presupposes, then,
human thought and its principles. And not only the abstract logical
principles are thus pre-supposed. The evolution hypothesis purports to be
not only a formal arrangement of phenomena, but to express also the law of
a real process. It supposes, then, that the real data—all that in our
knowledge which we do not produce ourselves, but which we in the main
simply receive—are subject to laws which are at least analogous to the
logical relations of our thoughts; in other words, it assumes the validity of
the principle of causality. If organic species could arise without cause there
would be no use in framing hypotheses. Only if we assume the principle of
causality, is there a problem to solve.

Though Darwinism has had a great influence on philosophy considered as a
striving after a scientific view of the world, yet here is a point of view—the
epistemological—where philosophy is not only independent but reaches
beyond any result of natural science. Perhaps it will be said: the powers and
functions of organic beings only persist (perhaps also only arise) when they
correspond sufficiently to the conditions under which the struggle of life is
to go on. Human thought itself is, then, a variation (or a mutation) which
has been able to persist and to survive. Is not, then, the problem of
knowledge solved by the evolution hypothesis? Spencer had given an
affirmative answer to this question before the appearance of The Origin of
Species. For the individual, he said, there is an à priori, original, basis (or
Anlage) for all mental life; but in the species all powers have developed in
reciprocity with extendal conditions. Knowledge is here considered from
the practical point of view, as a weapon in the struggle for life, as an
"organon" which has been continuously in use for generations. In recent
years the economic or pragmatic epistemology, as developed by Avenarius
and Mach in Germany, and by James in America, points in the same
direction. Science, it is said, only maintains those principles and
presuppositions which are necessary to the simplest and clearest orientation
be applied to experience and to practical work, will successively be
eliminated.
In these views a striking and important application is made of the idea of
struggle for life to the development of human thought. Thought must, as all
other things in the world, struggle for life. But this whole consideration
belongs to psychology, not to the theory of knowledge (epistemology),
which is concerned only with the validity of knowledge, not with its
historical origin. Every hypothesis to explain the origin of knowledge must
submit to cross-examination by the theory of knowledge, because it works
with the fundamental forms and principles of human thought. We cannot go
further back than these forms and principles, which it is the aim of
epistemology to ascertain and for which no further reason can be given.[202]
But there is another side of the problem which is, perhaps, of more
importance and which epistemology generally overlooks. If new variations
can arise, not only in organic but perhaps also in inorganic nature, new
tasks are placed before the human mind. The question is, then, if it has
forms in which there is room for the new matter? We are here touching a

possibility which the great master of epistemology did not bring to light.
Kant supposed confidently that no other matter of knowledge could stream
forth from the dark source which he called "the thing-in-itself," than such as
could be synthesised in our existing forms of knowledge. He mentions the
possibility of other forms than the human, and warns us against the
dogmatic assumption that the human conception of existence should be
absolutely adequate. But he seems to be quite sure that the thing-in-itself
works constantly, and consequently always gives us only what our powers
can master. This assumption was a consequence of Kant's rationalistic
tendency, but one for which no warrant can be given. Evolutionism and
systematism are opposing tendencies which can never be absolutely
harmonised one with the other. Evolution may at any time break some form
which the system-monger regards as finally established. Darwin himself felt
a great difference in looking at variation as an evolutionist and as a
systematist. When he was working at his evolution theory, he was very glad
to find variations; but they were a hindrance to him when he worked as a
systematist, in preparing his work on Cirripedia. He says in a letter: "I had
thought the same parts of the same species more resemble (than they do
anyhow in Cirripedia) objects cast in the same mould. Systematic work
would be easy were it not for this confounded variation, which, however, is
pleasant to me as a speculatist, though odious to me as a systematist."[203]
He could indeed be angry with variations even as an evolutionist; but then
only because he could not explain them, not because he could not classify
them. "If, as I must think, external conditions produce little direct effect,
what the devil determines each particular variation?"[204] What Darwin
experienced in this particular domain holds good of all knowledge. All
knowledge is systematic, in so far as it strives to put phenomena in quite
definite relations, one to another. But the systematisation can never be
complete. And here Darwin has contributed much to widen the world, for
us. He has shown us forces and tendencies in nature which make absolute
systems impossible, at the same time that they give us new objects and
problems. There is still a place for what Lessing called "the unceasing
striving after truth," while "absolute truth" (in the sense of a closed system)
is unattainable so long as life and experience are going on.
There is here a special remark to be made. As we have seen above, recent
research has shown that natural selection or struggle for life is no

explanation of variations. Hugo de Vries distinguishes between partial and
embryonal variations, or between variations and mutations, only the last-
named being heritable, and therefore of importance for the origin of new
species. But the existence of variations is not only of interest for the
problem of the origin of species; it has also a more general interest. An
individual does not lose its importance for knowledge, because its qualities
are not heritable. On the contrary, in higher beings at least, individual
peculiarities will become more and more independent objects of interest.
Knowledge takes account of the biographies not only of species, but also of
individuals: it seeks to find the law of development of the single individual.
[205] As Leibnitz said long ago, individuality consists in the law of the
changes of a being: "La loi du changement fait l'individualité de chaque
substance." Here is a world which is almost new for science, which till now
has mainly occupied itself with general laws and forms. But these are
ultimately only means to understand the individual phenomena, in whose
nature and history a manifold of laws and forms always coöperate. The
importance of this remark will appear in the sequel.
V
To many people the Darwinian theory of natural selection or struggle for
existence seemed to change the whole conception of life, and particularly
all the conditions on which the validity of ethical ideas depends. If only that
has persistence which can be adapted to a given condition, what will then
be the fate of our ideals, of our standards of good and evil? Blind force
seems to reign, and the only thing that counts seems to be the most heedless
use of power. Darwinism, it was said, has proclaimed brutality. No other
difference seems permanent save that between the sound, powerful and
happy on the one side, the sick, feeble and unhappy on the other; and every
attempt to alleviate this difference seems to lead to general enervation.
Some of those who interpreted Darwinism in this manner felt an aesthetic
delight in contemplating the heedlessness and energy of the great struggle
for existence and anticipated the realisation of a higher human type as the
outcome of it: so Nietzsche and his followers. Others recognising the same
consequences in Darwinism regarded these as one of the strongest
objections against it; so Dühring and Kropotkin (in his earlier works).

This interpretation of Darwinism was frequent in the interval between the
two main works of Darwin—The Origin of Species and The Descent of
Man. But even during this interval it was evident to an attentive reader that
Darwin himself did not found his standard of good and evil on the features
of the life of nature he had emphasised so strongly. He did not justify the
ways along which nature reached its ends; he only pointed them out. The
"real" was not to him, as to Hegel, one with the "rational." Darwin has,
indeed, by his whole conception of nature, rendered a great service to ethics
in making the difference between the life of nature and the ethical life
appear in so strong a light. The ethical problem could now be stated in a
sharper form than before. But this was not the first time that the idea of the
struggle for life was put in relation to the ethical problem. In the
seventeenth century Thomas Hobbes gave the first impulse to the whole
modern discussion of ethical principles in his theory of bellum omnium
contra omnes. Men, he taught, are in the state of nature enemies one of
another, and they live either in fright or in the glory of power. But it was not
the opinion of Hobbes that this made ethics impossible. On the contrary, he
found a standard for virtue and vice in the fact that some qualities and
actions have a tendency to bring us out of the state of war and to secure
peace, while other qualities have a contrary tendency. In the eighteenth
century even Immanuel Kant's ideal ethics had—so far as can be seen—a
similar origin. Shortly before the foundation of his definitive ethics, Kant
wrote his Idee zu einer allgemeinen Weltgeschichte (1784), where—in a
way which reminds us of Hobbes, and is prophetic of Darwin—he describes
the forward-driving power of struggle in the human world. It is here as with
the struggle of the trees for light and air, through which they compete with
one another in height. Anxiety about war can only be allayed by an
ordinance which gives everyone his full liberty under acknowledgment of
the equal liberty of others. And such ordinance and acknowledgment are
also attributes of the content of the moral law, as Kant proclaimed it in the
year after the publication of his essay (1785).[206] Kant really came to his
ethics by the way of evolution, though he afterwards disavowed it.
Similarly the same line of thought may be traced in Hegel though it has
been disguised in the form of speculative dialectics.[207] And in
Schopenhauer's theory of the blind will to live and its abrogation by the
ethical feeling, which is founded on universal sympathy, we have a more
individualistic form of the same idea.

It was, then, not entirely a foreign point of view which Darwin introduced
into ethical thought, even if we take no account of the poetical character of
the word "struggle" and of the more direct adaptation, through the use and
non-use of power, which Darwin also emphasised. In The Descent of Man
he has devoted a special chapter[208] to a discussion of the origin of the
ethical consciousness. The characteristic expression of this consciousness
he found, just as Kant did, in the idea of "ought"; it was the origin of this
new idea which should be explained. His hypothesis was that the ethical
"ought" has its origin in the social and parental instincts, which, as well as
other instincts (e.g. the instinct of self-preservation), lie deeper than
pleasure and pain. In many species, not least in the human species, these
instincts are fostered by natural selection; and when the powers of memory
and comparison are developed, so that single acts can be valued according
to the claims of the deep social instinct, then consciousness of duty and
remorse are possible. Blind instinct has developed to conscious ethical will.
As already stated, Darwin, as a moral philosopher belongs to the school that
was founded by Shaftesbury, and was afterwards represented by Hutcheson,
Hume, Adam Smith, Comte and Spencer. His merit is, first, that he has
given this tendency of thought a biological foundation, and that he has
stamped on it a doughty character in showing that ethical ideas and
sentiments, rightly conceived, are forces which are at work in the struggle
for life.
There are still many questions to solve. Not only does the ethical
development within the human species contain features still unexplained;
[209] but we are confronted by the great problem whether after all a genetic
historical theory can be of decisive importance here. To every consequent
ethical consciousness there is a standard of value, a primordial value which
determines the single ethical judgments as their last presupposition, and the
"rightness" of this basis, the "value" of this value can as little be discussed
as the "rationality" of our logical principles. There is here revealed a
possibility of ethical scepticism which evolutionistic ethics (as well as
intuitive or rationalistic ethics) has overlooked. No demonstration can show
that the results of the ethical development are definitive and universal. We
meet here again with the important opposition of systematisation and
evolution. There will, I think, always be an open question here, though

comparative ethics, of which we have so far only the first attempts, can do
much to throw light on it.
It would carry us too far to discuss all the philosophical works on ethics,
which have been influenced directly or indirectly by evolutionism. I may,
however, here refer to the book of C. M. Williams, A Review of the Systems
of Ethics founded on the Theory of Evolution,[210] in which, besides Darwin,
the following authors are reviewed: Wallace, Haeckel, Spencer, Fiske,
Rolph, Barratt, Stephen, Carneri, Höffding, Gizycki, Alexander, Rée. As
works which criticise evolutionistic ethics from an intuitive point of view
and in an instructive way, may be cited: Guyau, La morale anglaise
contemporaine,[211] and Sorley, Ethics of Naturalism. I will only mention
some interesting contributions to ethical discussion which can be found in
Darwinism besides the idea of struggle for life.
The attention which Darwin has directed to variations has opened our eyes
to the differences in human nature as well as in nature generally. There is
here a fact of great importance for ethical thought, no matter from what
ultimate premiss it starts. Only from a very abstract point of view can
different individuals be treated in the same manner. The most eminent
ethical thinkers, men such as Jeremy Bentham and Immanuel Kant, who
discussed ethical questions from very opposite standpoints, agreed in
regarding all men as equal in respect of ethical endowment. In regard to
Bentham, Leslie Stephen remarks: "He is determined to be thoroughly
empirical, to take men as he found them. But his utilitarianism supposed
that men's views of happiness and utility were uniform and clear, and that
all that was wanted was to show them the means by which their ends could
be reached."[212] And Kant supposed that every man would find the
"categorical imperative" in his consciousness, when he came to sober
reflexion, and that all would have the same qualifications to follow it. But if
continual variations, great or small, are going on in human nature, it is the
duty of ethics to make allowance for them, both in making claims, and in
valuing what is done. A new set of ethical problems have their origin here.
[213] It is an interesting fact that Stuart Mill's book On Liberty appeared in
the same year as The Origin of Species. Though Mill agreed with Bentham
about the original equality of all men's endowments, he regarded individual
differences as a necessary result of physical and social influences, and he
claimed that free play shall be allowed to differences of character so far as

is possible without injury to other men. It is a condition of individual and
social progress that a man's mode of action should be determined by his
own character and not by tradition and custom, nor by abstract rules. This
view was to be corroborated by the theory of Darwin.
But here we have reached a point of view from which the criticism, which
in recent years has often been directed against Darwin—that small
variations are of no importance in the struggle for life—is of no weight.
From an ethical standpoint, and particularly from the ethical standpoint of
Darwin himself, it is a duty to foster individual differences that can be
valuable, even though they can neither be of service for physical
preservation nor be physically inherited. The distinction between variation
and mutation is here without importance. It is quite natural that biologists
should be particularly interested in such variations as can be inherited and
produce new species. But in the human world there is not only a physical,
but also a mental and social heredity. When an ideal human character has
taken form, then there is shaped a type, which through imitation and
influence can become an important factor in subsequent development, even
if it cannot form a species in the biological sense of the word. Spiritually
strong men often succumb in the physical struggle for life; but they can
nevertheless be victorious through the typical influence they exert, perhaps
on very distant generations, if the remembrance of them is kept alive, be it
in legendary or in historical form. Their very failure can show that a type
has taken form which is maintained at all risks, a standard of life which is
adhered to in spite of the strongest opposition. The question "to be or not to
be" can be put from very different levels of being: it has too often been
considered a consequence of Darwinism that this question is only to be put
from the lowest level. When a stage is reached, where ideal (ethical,
intellectual, aesthetic) interests are concerned, the struggle for life is a
struggle for the preservation of this stage. The giving up of a higher
standard of life is a sort of death; for there is not only a physical, there is
also a spiritual, death.
VI
The Socratic character of Darwin's mind appears in his wariness in drawing
the last consequences of his doctrine, in contrast both with the audacious
theories of so many of his followers and with the consequences which his

antagonists were busy in drawing. Though he, as we have seen, saw from
the beginning that his hypothesis would occasion "a whole of metaphysics,"
he was himself very reserved as to the ultimate questions, and his answers
to such questions were extorted from him.
As to the question of optimism and pessimism, Darwin held that though
pain and suffering were very often the ways by which animals were led to
pursue that course of action which is most beneficial to the species, yet
pleasurable feelings were the most habitual guides. "We see this in the
pleasure from exertion, even occasionally from great exertion of the body
or mind, in the pleasure of our daily meals, and especially in the pleasure
derived from sociability, and from loving our families." But there was to
him so much suffering in the world that it was a strong argument against the
existence of an intelligent First Cause.[214]
It seems to me that Darwin was not so clear on another question, that of the
relation between improvement and adaptation. He wrote to Lyell: "When
you contrast natural selection and 'improvement,' you seem always to
overlook ... that every step in the natural selection of each species implies
improvement in that species in relation to its condition of life....
Improvement implies, I suppose, each form obtaining many parts or
organs, all excellently adapted for their functions." "All this," he adds,
"seems to me quite compatible with certain forms fitted for simple
conditions, remaining unaltered, or being, degraded."[215] But the great
question is, if the conditions of life will in the long run favour
"improvement" in the sense of differentiation (or harmony of differentiation
and integration). Many beings are best adapted to their conditions of life if
they have few organs and few necessities. Pessimism would not only be the
consequence, if suffering outweighed happiness, but also if the most
elementary forms of happiness were predominant, or if there were a
tendency to reduce the standard of life to the simplest possible, the
contentment of inertia or stable equilibrium. There are animals which are
very highly differentiated and active in their young state, but later lose their
complex organisation and concentrate themselves on the one function of
nutrition. In the human world analogies to this sort of adaptation are not
wanting. Young "idealists" very often end as old "Philistines." Adaptation
and progress are not the same.

Another question of great importance in respect to human evolution is,
whether there will be always a possibility for the existence of an impulse to
progress, an impulse to make great claims on life, to be active and to alter
the conditions of life instead of adapting to them in a passive manner. Many
people do not develop because they have too few necessities, and because
they have no power to imagine other conditions of life than those under
which they live. In his remarks on "the pleasure from exertion" Darwin has
a point of contact with the practical idealism of former times—with the
ideas of Lessing and Goethe, of Condorcet and Fichte. The continual
striving which was the condition of salvation to Faust's soul, is also the
condition of salvation to mankind. There is a holy fire which we ought to
keep burning, if adaptation is really to be improvement. If, as I have tried to
show in my Philosophy of Religion, the innermost core of all religion is
faith in the persistence of value in the world, and if the highest values
express themselves in the cry "Excelsior!" then the capital point is, that this
cry should always be heard and followed. We have here a corollary of the
theory of evolution in its application to human life.
Darwin declared himself an agnostic, not only because he could not
harmonise the large amount of suffering in the world with the idea of a God
as its first cause, but also because he "was aware that if we admit a first
cause, the mind still craves to know whence it came and how it arose."[216]
He saw, as Kant had seen before him and expressed in his Kritik der
Urtheilskraft, that we cannot accept either of the only two possibilities
which we are able to conceive: chance (or brute force) and design. Neither
mechanism nor teleology can give an absolute answer to ultimate questions.
The universe, and especially the organic life in it, can neither be explained
as a mere combination of absolute elements nor as the effect of a
constructing thought. Darwin concluded, as Kant, and before him Spinoza,
that the oppositions and distinctions which our experience presents, cannot
safely be regarded as valid for existence in itself. And, with Kant and
Fichte, he found his stronghold in the conviction that man has something to
do, even if he cannot solve all enigmas. "The safest conclusion seems to me
that the whole subject is beyond the scope of man's intellect; but man can
do his duty."[217]
Is this the last word of human thought? Does not the possibility, that man
can do his duty, suppose that the conditions of life allow of continuous

ethical striving, so that there is a certain harmony between cosmic order and
human ideals? Darwin himself has shown how the consciousness of duty
can arise as a natural result of evolution. Moreover there are lines of
evolution which have their end in ethical idealism, in a kingdom of values,
which must struggle for life as all things in the world must do, but a
kingdom which has its firm foundation in reality.

FOOTNOTES:
[195] Life and Letters of Charles Darwin, Vol. i. p. 8.
[196] Encyclopädie der philosophischen Wissenschaften (4th edit.), Berlin,
1845, § 249.
[197] Lehrbuch der Naturphilosophie, Jena, 1809.
[198] Ueber den Willen in der Natur (2nd edit.), Frankfurt a. M., 1854, pp. 41-
43.
[199] Spencer, Autobiography, Vol. ii. p. 50, London and New York, 1904.
[200] Autobiography, Vol. ii. p. 100.
[201] Cf. my letter to him 1876, now printed in Duncan's Life and Letters of
Herbert Spencer, p. 178. London, 1908.
[202] The present writer, many years ago, in his Psychology (Copenhagen, 1882;
Eng. transl. London, 1891), criticised the evolutionistic treatment of the problem
of knowledge from the Kantian point of view.
[203] Life and Letters, Vol. II. p. 37.
[204] Ibid. p. 232.
[205] The new science of Ecology occupies an intermediate position between the
biography of species and the biography of individuals. Compare Congress of
Arts and Science, St. Louis, Vol. v. 1906 (The Reports of Drude and Robinson)
and the work of my colleague, E. Warming.
[206] Cf. my History of Modern Philosophy (Eng. transl. London, 1900), i. pp.
76-79.
[207] "Herrschaft und Knechtschaft," Phönomenologie des Geistes, iv. A.,
Leiden, 1907.
[208] The Descent of Man, Vol. i. Ch. iii.
[209] The works of Westermarck and Hobhouse throw new light on many of
these features.
[210] New York and London, 1893.

[211] Paris, 1879.
[212] English literature and society in the eighteenth century, London, 1904, p.
187.
[213] Cf. my paper, "The law of relativity in Ethics," International Journal of
Ethics, Vol. i. 1891, pp. 37-62.
[214] Life and Letters, Vol. I. p. 310.
[215] Ibid. Vol. II. p. 177.
[216] Life and Letters, Vol. 1. p. 306.
[217] Life and Letters, p. 307.

VIII

THE INFLUENCE OF DARWIN UPON
RELIGIOUS THOUGHT
By P. N. Waggett, M.A., S.S.J.E.

I

The object of this paper is first to point out certain elements of the
Darwinian influence upon Religious thought, and then to show reason for
the conclusion that it has been, from a Christian point of view, satisfactory. I
shall not proceed further to urge that the Christian apologetic in relation to
biology has been successful. A variety of opinions may be held on this
question, without disturbing the conclusion that the movements of
readjustment have been beneficial to those who remain Christians, and this
by making them more Christian and not only more liberal. The theologians
may sometimes have retreated, but there has been an advance of theology. I
know that this account incurs the charge of optimism. It is not the worst that
could be made. The influence has been limited in personal range, unequal,
even divergent, in operation, and accompanied by the appearance of waste
and mischievous products. The estimate which follows requires for due
balance a full development of many qualifying considerations. For this I
lack space, but I must at least distinguish my view from the popular one that
our difficulties about religion and natural science have come to an end.
Concerning the older questions about origins—the origin of the world, of
species, of man, of reason, conscience, religion—a large measure of
understanding has been reached by some thoughtful men. But meanwhile
new questions have arisen, questions about conduct, regarding both the
reality of morals and the rule of right action for individuals and societies.
And these problems, still far from solution, may also be traced to the
influence of Darwin. For they arise from the renewed attention to heredity,
brought about by the search for the causes of variation, without which the
study of the selection of variations has no sufficient basis.

Even the existing understanding about origins is very far from universal. On
these points there were always thoughtful men who denied the necessity of
conflict, and there are still thoughtful men who deny the possibility of a
truce.
It must further be remembered that the earlier discussion now, as I hope to
show, producing favourable results, created also for a time grave damage,
not only in the disturbance of faith and the loss of men—a loss not repaired
by a change in the currents of debate—but in what I believe to be a still
more serious respect. I mean the introduction of a habit of facile and
untested hypothesis in religious as in other departments of thought.
Darwin is not responsible for this, but he is in part the cause of it. Great
ideas are dangerous guests in narrow minds; and thus it has happened that
Darwin—the most patient of scientific workers, in whom hypothesis waited
upon research, or if it provisionally outstepped it did so only with the most
scrupulously careful acknowledgment—has led smaller and less
conscientious men in natural science, in history, and in theology to an over-
eager confidence in probable conjecture and a loose grip upon the facts of
experience. It is not too much to say that in many quarters the age of
materialism was the least matter-of-fact age conceivable, and the age of
science the age which showed least of the patient temper of inquiry.
I have indicated, as shortly as I could, some losses and dangers which in a
balanced account of Darwin's influence would be discussed at length.
One other loss must be mentioned. It is a defect in our thought which, in
some quarters, has by itself almost cancelled all the advantages secured. I
mean the exaggerated emphasis on uniformity or continuity; the
unwillingness to rest any part of faith or of our practical expectation upon
anything that from any point of view can be called exceptional. The high
degree of success reached by naturalists in tracing, or reasonably
conjecturing, the small beginnings of great differences, has led the
inconsiderate to believe that anything may in time become anything else.
It is true that this exaggeration of the belief in uniformity has produced in
turn its own perilous reaction. From refusing to believe whatever can be
called exceptional, some have come to believe whatever can be called
wonderful.

But, on the whole, the discontinuous or highly various character of
experience received for many years too little deliberate attention. The
conception of uniformity which is a necessity of scientific description has
been taken for the substance of history. We have accepted a postulate of
scientific method as if it were a conclusion of scientific demonstration. In
the name of a generalisation which, however just on the lines of a particular
method, is the prize of a difficult exploit of reflexion, we have discarded the
direct impressions of experience; or, perhaps it is more true to say, we have
used for the criticism of alleged experiences a doctrine of uniformity which
is only valid in the region of abstract science. For every science depends for
its advance upon limitation of attention, upon the selection out of the whole
content of consciousness of that part or aspect which is measurable by the
method of the science. Accordingly there is a science of life which rightly
displays the unity underlying all its manifestations. But there is another
view of life, equally valid, and practically sometimes more important,
which recognises the immediate and lasting effect of crisis, difference, and
revolution. Our ardour for the demonstration of uniformity of process and
of minute continuous change needs to be balanced by a recognition of the
catastrophic element in experience, and also by a recognition of the
exceptional significance for us of events which may be perfectly regular
from an impersonal point of view.
An exorbitant jealousy of miracle, revelation, and ultimate moral
distinctions has been imported from evolutionary science into religious
thought. And it has been a damaging influence, because it has taken men's
attention from facts, and fixed them upon theories.
II
With this acknowledgment of important drawbacks, requiring many words
for their proper description, I proceed to indicate certain results of Darwin's
doctrine which I believe to be in the long run wholly beneficial to Christian
thought. These are:
The encouragement in theology of that evolutionary method of observation
and study, which has shaped all modern research:
The recoil of Christian apologetics towards the ground of religious
experience, a recoil produced by the pressure of scientific criticism upon

other supports of faith:
The restatement, or the recovery of ancient forms of statement, of the
doctrines of Creation and of divine Design in Nature, consequent upon the
discussion of evolution and of natural selection as its guiding factor.
(1) The first of these is quite possibly the most important of all. It was well
defined in a notable paper read by Dr. Gore, now Bishop of Birmingham, to
the Church Congress at Shrewsbury in 1896. We have learnt a new caution
both in ascribing and in denying significance to items of evidence, in
utterance or in event. There has been, as in art, a study of values, which
secures perspective and solidity in our representation of facts. On the one
hand, a given utterance or event cannot be drawn into evidence as if all
items were of equal consequence, like sovereigns in a bag. The question
whence and whither must be asked, and the particular thing measured as
part of a series. Thus measured it is not less truly important, but it may be
important in a lower degree. On the other hand, and for exactly the same
reason, nothing that is real is unimportant. The "failures" are not mere
mistakes. We see them, in St. Augustine's words, as "scholar's faults which
men praise in hope of fruit."
We cannot safely trace the origin of the evolutionistic method to the
influence of natural science. The view is tenable that theology led the way.
Probably this is a case of alternate and reciprocal debt. Quite certainly the
evolutionist method in theology, in Christian history and in the estimate of
scripture, has received vast reinforcement from biology, in which evolution
has been the ever present and ever victorious conception.
(2) The second effect named is the new willingness of Christian thinkers to
take definite account of religious experience. This is related to Darwin
through the general pressure upon religious faith of scientific criticism. The
great advance of our knowledge of organisms has been an important
element in the general advance of science. It has acted, by the varied
requirements of the theory of organisms, upon all other branches of natural
inquiry, and it held for a long time that leading place in public attention
which is now occupied by speculative physics. Consequently it contributed
largely to our present estimation of science as the supreme judge in all
matters of inquiry,[218] to the supposed destruction of mystery and the

disparagement of metaphysics which marked the last age, as well as to the
just recommendation of scientific method in branches of learning where the
direct acquisitions of natural science had no place.
Besides this, the new application of the idea of law and mechanical
regularity to the organic world seemed to rob faith of a kind of refuge. The
romantics had, as Berthelot[219] shows, appealed to life to redress the
judgments drawn from mechanism. Now, in Spencer, evolution gave us a
vitalist mechanic or mechanical vitalism, and the appeal seemed cut off. We
may return to this point later when we consider evolution; at present I only
endeavour to indicate that general pressure of scientific criticism which
drove men of faith to seek the grounds of reassurance in a science of their
own; in a method of experiment, of observation, of hypothesis checked by
known facts. It is impossible for me to do more than glance across the
threshold of this subject. But it is necessary to say that the method is in an
elementary stage of revival. The imposing success that belongs to natural
science is absent: we fall short of the unchallengeable unanimity of the
Biologists on fundamentals. The experimental method with its sure
repetitions cannot be applied to our subject-matter. But we have something
like the observational method of palaeontology and geographical
distribution; and in biology there are still men who think that the large
examination of varieties by way of geography and the search of strata is as
truly scientific, uses as genuinely the logical method of difference, and is as
fruitful in sure conclusions as the quasi-chemical analysis of Mendelian
laboratory work, of which last I desire to express my humble admiration.
Religion also has its observational work in the larger and possibly more
arduous manner.
But the scientific work in religion makes its way through difficulties and
dangers. We are far from having found the formula of its combination with
the historical elements of our apologetic. It is exposed, therefore, to a
damaging fire not only from unspiritualist psychology and pathology but
also from the side of scholastic dogma. It is hard to admit on equal terms a
partner to the old undivided rule of books and learning. With Charles Lamb,
we cry in some distress, "must knowledge come to me, if it come at all, by
some awkward experiment of intuition, and no longer by this familiar
process of reading?"[220] and we are answered that the old process has an
imperishable value, only we have not yet made clear its connection with

other contributions. And all the work is young, liable to be drawn into
unprofitable excursions, side-tracked by self-deceit and pretence; and it
fatally attracts, like the older mysticism, the curiosity and the expository
powers of those least in sympathy with it, ready writers who, with all the air
of extended research, have been content with narrow grounds for induction.
There is a danger, besides, which accompanies even the most genuine work
of this science and must be provided against by all its serious students. I
mean the danger of unbalanced introspection both for individuals and for
societies; of a preoccupation comparable to our modern social
preoccupation with bodily health; of reflexion upon mental states not
accompanied by exercise and growth of the mental powers; the danger of
contemplating will and neglecting work, of analysing conviction and not
criticising evidence.
Still, in spite of dangers and mistakes, the work remains full of hopeful
indications, and, in the best examples,[221] it is truly scientific in its
determination to know the very truth, to tell what we think, not what we
think we ought to think,[222] truly scientific in its employment of hypothesis
and verification, and in growing conviction of the reality of its subject-
matter through the repeated victories of a mastery which advances, like
science, in the Baconian road of obedience. It is reasonable to hope that
progress in this respect will be more rapid and sure when religious study
enlists more men affected by scientific desire and endowed with scientific
capacity.
The class of investigating minds is a small one, possibly even smaller than
that of reflecting minds. Very few persons at any period are able to find out
anything whatever. There are few observers, few discoverers, few who even
wish to discover truth. In how many societies the problems of philology
which face every person who speaks English are left unattempted! And if
the inquiring or the successfully inquiring class of minds is small, much
smaller, of course, is the class of those possessing the scientific aptitude in
an eminent degree. During the last age this most distinguished class was to
a very great extent absorbed in the study of phenomena, a study which had
fallen into arrears. For we stood possessed, in rudiment, of means of
observation, means for travelling and acquisition, qualifying men for a
larger knowledge than had yet been attempted. These were now to be
directed with new accuracy and ardour upon the fabric and behaviour of the

world of sense. Our debt to the great masters in physical science who
overtook and almost outstripped the task cannot be measured; and, under
the honourable leadership of Ruskin, we may all well do penance if we
have failed "in the respect due to their great powers of thought, or in the
admiration due to the far scope of their discovery."[223] With what
miraculous mental energy and divine good fortune—as Romans said of
their soldiers—did our men of curiosity face the apparently impenetrable
mysteries of nature! And how natural it was that immense accessions of
knowledge, unrelated to the spiritual facts of life, should discredit Christian
faith, by the apparent superiority of the new work to the feeble and
unprogressive knowledge of Christian believers! The day is coming when
men of this mental character and rank, of this curiosity, this energy and this
good fortune in investigation, will be employed in opening mysteries of a
spiritual nature. They will silence with masterful witness the over-confident
denials of naturalism. They will be in danger of the widespread recognition
which thirty years ago accompanied every utterance of Huxley, Tyndall,
Spencer. They will contribute, in spite of adulation, to the advance of sober
religious and moral science.
And this result will be due to Darwin, first because by raising the dignity of
natural science, he encouraged the development of the scientific mind;
secondly because he gave to religious students the example of patient and
ardent investigation; and thirdly because by the pressure of naturalistic
criticism the religious have been driven to ascertain the causes of their own
convictions, a work in which they were not without the sympathy of men of
science.[224]
In leaving the subject of scientific religious inquiry, I will only add that I do
not believe it receives any important help—and certainly it suffers
incidentally much damaging interruption—from the study of abnormal
manifestations or abnormal conditions of personality.
(3) Both of the above effects seem to me of high, perhaps the very highest,
importance to faith and to thought. But, under the third head, I name two
which are more directly traceable to the personal work of Darwin, and more
definitely characteristic of the age in which his influence was paramount:
viz. the influence of the two conceptions of evolution and natural selection
upon the doctrine of creation and of design respectively.

It is impossible here, though it is necessary for a complete sketch of the
matter, to distinguish the different elements and channels of this Darwinian
influence; in Darwin's own writings, in the vigourous polemic of Huxley,
and strangely enough, but very actually for popular thought, in the teaching
of the definitely anti-Darwinian evolutionist Spencer.
Under the head of the directly and purely Darwinian elements I should class
as preeminent the work of Wallace and of Bates; for no two sets of facts
have done more to fix in ordinary intelligent minds a belief in organic
evolution and in natural selection as its guiding factor than the facts of
geographical distribution and of protective colour and mimicry. The facts of
geology were difficult to grasp and the public and theologians heard more
often of the imperfection than of the extent of the geological record. The
witness of embryology, depending to a great extent upon microscopic work,
was and is beyond the appreciation of persons occupied in fields of work
other than biology.
III
From the influence in religion of scientific modes of thought we pass to the
influence of particular biological conceptions. The former effect comes by
way of analogy, example, encouragement and challenge; inspiring or
provoking kindred or similar modes of thought in the field of theology; the
latter by a collision of opinions upon matters of fact or conjecture which
seem to concern both science and religion.
In the case of Darwinism the story of this collision is familiar, and falls
under the heads of evolution and natural selection, the doctrine of descent
with modification, and the doctrine of its guidance or determination by the
struggle for existence between related varieties. These doctrines, though
associated and interdependent, and in popular thought not only combined
but confused, must be considered separately. It is true that the ancient
doctrine of Evolution, in spite of the ingenuity and ardour of Lamarck,
remained a dream tantalising the intellectual ambition of naturalists, until
the day when Darwin made it conceivable by suggesting the machinery of
its guidance. And, further, the idea of natural selection has so effectively
opened the door of research and stimulated observation in a score of
principal directions that, even if the Darwinian explanation became one day

much less convincing than, in spite of recent criticism, it now is, yet its
passing, supposing it to pass, would leave the doctrine of Evolution
immeasurably and permanently strengthened. For in the interests of the
theory of selection, "Für Darwin," as Müller wrote, facts have been
collected which remain in any case evidence of the reality of descent with
modification.
But still, though thus united in the modern history of convictions, though
united and confused in the collision of biological and traditional opinion,
yet evolution and natural selection must be separated in theological no less
than in biological estimation. Evolution seemed inconsistent with Creation;
natural selection with Providence and Divine design.
Discussion was maintained about these points for many years and with
much dark heat. It ranged over many particular topics and engaged minds
different in tone, in quality, and in accomplishment. There was at most
times a degree of misconception. Some naturalists attributed to theologians
in general a poverty of thought which belonged really to men of a particular
temper or training. The "timid theism" discerned in Darwin by so cautious a
theologian as Liddon[225] was supposed by many biologists to be the
necessary foundation of an honest Christianity. It was really more
characteristic of devout naturalists like Philip Henry Gosse, than of
religious believers as such.[226] The study of theologians more considerable
and even more typically conservative than Liddon does not confirm the
description of religious intolerance given in good faith, but in serious
ignorance, by a disputant so acute, so observant and so candid as Huxley.
Something hid from each other's knowledge the devoted pilgrims in two
great ways of thought. The truth may be, that naturalists took their view of
what creation was from Christian men of science who naturally looked in
their own special studies for the supports and illustrations of their religious
belief. Of almost every labourious student it may be said: "Hic ab arte sua
non recessit." And both the believing and the denying naturalists, confining
habitual attention to a part of experience, are apt to affirm and deny with
trenchant vigour and something of a narrow clearness "Qui respiciunt ad
pauca, de facili pronunciant."[227]
Newman says of some secular teachers that "they persuade the world of
what is false by urging upon it what is true." Of some early opponents of

Darwin it might be said by a candid friend that, in all sincerity of devotion
to truth, they tried to persuade the world of what is true by urging upon it
what is false. If naturalists took their version of orthodoxy from amateurs in
theology, some conservative Christians, instead of learning what evolution
meant to its regular exponents, took their view of it from celebrated
persons, not of the front rank in theology or in thought, but eager to take
account of public movements and able to arrest public attention.
Cleverness and eloquence on both sides certainly had their share in
producing the very great and general disturbance of men's minds in the
early days of Darwinian teaching. But by far the greater part of that
disturbance was due to the practical novelty and the profound importance of
the teaching itself, and to the fact that the controversy about evolution
quickly became much more public than any controversy of equal
seriousness had been for many generations.
We must not think lightly of that great disturbance because it has, in some
real sense, done its work, and because it is impossible in days of more
coolness and light, to recover a full sense of its very real difficulties.
Those who would know them better should add to the calm records of
Darwin[228] and to the story of Huxley's impassioned championship, all that
they can learn of George Romanes.[229] For his life was absorbed in this
very struggle and reproduced its stages. It began in a certain assured
simplicity of biblical interpretation; it went on, through the glories and
adventures of a paladin in Darwin's train, to the darkness and dismay of a
man who saw all his most cherished beliefs rendered, as he thought,
incredible.[230] He lived to find the freer faith for which process and
purpose are not irreconcilable, but necessary to one another. His
development, scientific, intellectual and moral, was itself of high
significance; and its record is of unique value to our own generation, so
near the age of that doubt and yet so far from it; certainly still much in need
of the caution and courage by which past endurance prepares men for new
emergencies. We have little enough reason to be sure that in the discussions
awaiting us we shall do as well as our predecessors in theirs. Remembering
their endurance of mental pain, their ardour in mental labour, the heroic
temper and the high sincerity of controversialists on either side, we may
well speak of our fathers in such words of modesty and self-judgment as

Drayton used when he sang the victors of Agincourt. The progress of
biblical study, in the departments of Introduction and Exegesis, resulting in
the recovery of a point of view anciently tolerated if not prevalent, has
altered some of the conditions of that discussion. In the years near 1858, the
witness of Scripture was adduced both by Christian advocates and their
critics as if unmistakably irreconcilable with Evolution.
Huxley[231] found the path of the blameless naturalist everywhere blocked
by "Moses": the believer in revelation was generally held to be forced to a
choice between revealed cosmogony and the scientific account of origins. It
is not clear how far the change in Biblical interpretation is due to natural
science, and how far to the vital movements of theological study which
have been quite independent of the controversy about species. It belongs to
a general renewal of Christian movement, the recovery of a heritage.
"Special Creation"—really a biological rather than a theological conception,
—seems in its rigid form to have been a recent element even in English
biblical orthodoxy.
The Middle Ages had no suspicion that religious faith forbad inquiry into
the natural origination of the different forms of life. Bartholomaeus
Anglicus, an English Franciscan of the thirteenth century, was a mutationist
in his way, as Aristotle, "the Philosopher" of the Christian Schoolmen, had
been in his. So late as the seventeenth century, as we learn not only from
early proceedings of the Royal Society, but from a writer so homely and so
regularly pious as Walton, the variation of species and "spontaneous"
generations had no theological bearing, except as instances of that various
wonder of the world which in devout minds is food for devotion.
It was in the eighteenth century that the harder statement took shape.
Something in the preciseness of that age, its exaltation of law, its cold
passion for a stable and measured universe, its cold denial, its cold
affirmation of the power of God, a God of ice, is the occasion of that
rigidity of religious thought about the living world which Darwin by
accident challenged, or rather by one of those movements of genius which,
Goethe[232] declares, are "elevated above all earthly control."
If religious thought in the eighteenth century was aimed at a fixed and
nearly finite world of spirit, it followed in all these respects the secular and

critical lead. "La philosophie réformatrice du XVIIIe siècle[233] ramenait la
nature et la société à des mécanismes que la pensée réfléchie peut concevoir
et récomposer." In fact, religion in a mechanical age is condemned if it
takes any but a mechanical tone. Butler's thought was too moving, too vital,
too evolutionary, for the sceptics of his time. In a rationalist, encyclopaedic
period, religion also must give hard outline to its facts, it must be able to
display its secret to any sensible man in the language used by all sensible
men. Milton's prophetic genius furnished the eighteenth century, out of the
depth of the passionate age before it, with the theological tone it was to
need. In spite of the austere magnificence of his devotion, he gives to
smaller souls a dangerous lead. The rigidity of Scripture exegesis belonged
to this stately but imperfectly sensitive mode of thought. It passed away
with the influence of the older rationalists whose precise denials matched
the precise and limited affirmations of the static orthodoxy.
I shall, then, leave the specially biblical aspect of the debate—interesting as
it is and even useful, as in Huxley's correspondence with the Duke of Argyll
and others in 1892[234]—in order to consider without complication the
permanent elements of Christian thought brought into question by the
teaching of evolution.
Such permanent elements are the doctrine of God as Creator of the
universe, and the doctrine of man as spiritual and unique. Upon both the
doctrine of evolution seemed to fall with crushing force.
With regard to Man I leave out, acknowledging a grave omission, the
doctrine of the Fall and of Sin. And I do so because these have not yet, as I
believe, been adequately treated: here the fruitful reaction to the stimulus of
evolution is yet to come. The doctrine of sin, indeed, falls principally within
the scope of that discussion which has followed or displaced the Darwinian;
and without it the Fall cannot be usefully considered. For the question about
the Fall is a question not merely of origins, but of the interpretation of
moral facts whose moral reality must first be established.
I confine myself therefore to Creation and the dignity of man.
The meaning of evolution, in the most general terms, is that the
differentiation of forms is not essentially separate from their behaviour and
use; that if these are within the scope of study, that is also; that the world

has taken the form we see by movements not unlike those we now see in
progress; that what may be called proximate origins are continuous in the
way of force and matter, continuous in the way of life, with actual
occurrences and actual characteristics. All this has no revolutionary bearing
upon the question of ultimate origins. The whole is a statement about
process. It says nothing to metaphysicians about cause. It simply brings
within the scope of observation or conjecture that series of changes which
has given their special characters to the different parts of the world we see.
In particular, evolutionary science aspires to the discovery of the process or
order of the appearance of life itself: if it were to achieve its aim it could
say nothing of the cause of this or indeed of the most familiar occurrences.
We should have become spectators or convinced historians of an event
which, in respect of its cause and ultimate meaning, would be still
impenetrable.
With regard to the origin of species, supposing life already established,
biological science has the well founded hopes and the measure of success
with which we are all familiar. All this has, it would seem, little chance of
collision with a consistent theism, a doctrine which has its own difficulties
unconnected with any particular view of order or process. But when it was
stated that species had arisen by processes through which new species were
still being made, evolutionism came into collision with a statement,
traditionally religious, that species were formed and fixed once for all and
long ago.
What is the theological import of such a statement when it is regarded as
essential to belief in God? Simply that God's activity, with respect to the
formation of living creatures, ceased at some point in past time.
"God rested" is made the touchstone of orthodoxy. And when, under the
pressure of the evidences, we found ourselves obliged to acknowledge and
assert the present and persistent power of God, in the maintenance and in
the continued formation of "types," what happened was the abolition of a
time-limit. We were forced only to a bolder claim, to a theistic language less
halting, more consistent, more thorough in its own line, as well as better
qualified to assimilate and modify such schemes as Von Hartmann's
philosophy of the Unconscious—a philosophy, by the way, quite intolerant
of a merely mechanical evolution.[235]

Here was not the retrenchment of an extravagant assertion, but the
expansion of one which was faltering and inadequate. The traditional
statement did not need paring down so as to pass the meshes of a new and
exacting criticism. It was itself a net meant to surround and enclose
experience; and we must increase its size and close its mesh to hold newly
disclosed facts of life. The world, which had seemed a fixed picture or
model, gained first perspective and then solidity and movement. We had a
glimpse of organic history; and Christian thought became more living and
more assured as it met the larger view of life.
However unsatisfactory the new attitude might be to our critics, to
Christians the reform was positive. What was discarded was a limitation, a
negation. The movement was essentially conservative, even actually
reconstructive. For the language disused was a language inconsistent with
the definitions of orthodoxy; it set bounds to the infinite, and by implication
withdrew from the creative rule all such processes as could be brought
within the descriptions of research. It ascribed fixity and finality to that
"creature" in which an apostle taught us to recognise the birth-struggles of
an unexhausted progress. It tended to banish mystery from the world we
see, and to confine it to a remote first age.
In the reformed, the restored, language of religion, Creation became again
not a link in a rational series to complete a circle of the sciences, but the
mysterious and permanent relation between the infinite and the finite,
between the moving changes we know in part, and the Power, after the
fashion of that observation, unknown, which is itself "unmoved all motion's
source."[236]
With regard to man it is hardly necessary, even were it possible, to illustrate
the application of this bolder faith. When the record of his high extraction
fell under dispute, we were driven to a contemplation of the whole of his
life, rather than of a part and that part out of sight. We remembered again,
out of Aristotle, that the result of a process interprets its beginnings. We
were obliged to read the title of such dignity as we may claim, in results and
still more in aspirations.
Some men still measure the value of great present facts in life—reason and
virtue and sacrifice—by what a self-disparaged reason can collect of the

meaner rudiments of these noble gifts. Mr. Balfour has admirably displayed
the discrepancy, in this view, between the alleged origin and the alleged
authority of reason. Such an argument ought to be used not to discredit the
confident reason, but to illuminate and dignify its dark beginnings, and to
show that at every step in the long course of growth a Power was at work
which is not included in any term or in all the terms of the series.
I submit that the more men know of actual Christian teaching, its fidelity to
the past, and its sincerity in face of discovery, the more certainly they will
judge that the stimulus of the doctrine of evolution has produced in the long
run vigour as well as flexibility in the doctrine of Creation and of man.
I pass from Evolution in general to Natural Selection.
The character in religious language which I have for short called
mechanical was not absent in the argument from design as stated before
Darwin. It seemed to have reference to a world conceived as fixed. It
pointed, not to the plastic capacity and energy of living matter, but to the
fixed adaptation of this and that organ to an unchanging place or function.
Mr. Hobhouse has given us the valuable phrase "a niche of organic
opportunity." Such a phrase would have borne a different sense in non-
evolutionary thought. In that thought, the opportunity was an opportunity
for the Creative Power, and Design appeared in the preparation of the
organism to fit the niche. The idea of the niche and its occupant growing
together from simpler to more complex mutual adjustment was unwelcome
to this teleology. If the adaptation was traced to the influence, through
competition, of the environment, the old teleology lost an illustration and a
proof. For the cogency of the proof in every instance depended upon the
absence of explanation. Where the process of adaptation was discerned, the
evidence of Purpose or Design was weak. It was strong only when the
natural antecedents were not discovered, strongest when they could be
declared undiscoverable.
Paley's favourite word is "Contrivance"; and for him contrivance is most
certain where production is most obscure. He points out the physiological
advantage of the valvulae conniventes to man, and the advantage for
teleology of the fact that they cannot have been formed by "action and
pressure." What is not due to pressure may be attributed to design, and

when a "mechanical" process more subtle than pressure was suggested, the
case for design was so far weakened. The cumulative proof from the
multitude of instances began to disappear when, in selection, a natural
sequence was suggested in which all the adaptations might be reached by
the motive power of life, and especially when, as in Darwin's teaching,
there was full recognition of the reactions of life to the stimulus of
circumstance. "The organism fits the niche," said the teleologist, "because
the Creator formed it so as to fit." "The organism fits the niche," said the
naturalist, "because unless it fitted it could not exist." "It was fitted to
survive," said the theologian. "It survives because it fits," said the
selectionist. The two forms of statement are not incompatible; but the new
statement, by provision of an ideally universal explanation of process, was
hostile to a doctrine of purpose which relied upon evidences always
exceptional however numerous. Science persistently presses on to find the
universal machinery of adaptation in this planet; and whether this be found
in selection, or in direct-effect, or in vital reactions resulting in large
changes, or in a combination of these and other factors, it must always be
opposed to the conception of a Divine Power here and there but not
everywhere active.
For science, the Divine must be constant, operative everywhere and in
every quality and power, in environment and in organism, in stimulus and
in reaction, in variation and in struggle, in hereditary equilibrium, and in
"the unstable state of species"; equally present on both sides of every strain,
in all pressures and in all resistances, in short in the general wonder of life
and the world. And this is exactly what the Divine Power must be for
religious faith.
The point I wish once more to make is that the necessary readjustment of
teleology, so as to make it depend upon the contemplation of the whole
instead of a part, is advantageous quite as much to theology as to science.
For the older view failed in courage. Here again our theism was not
sufficiently theistic.
Where results seemed inevitable, it dared not claim them as God-given. In
the argument from Design it spoke not of God in the sense of theology, but
of a Contriver, immensely, not infinitely wise and good, working within a
world, the scene, rather than the ever dependent outcome, of His Wisdom;

working in such emergencies and opportunities as occurred, by forces not
altogether within His control, towards an end beyond Himself. It gave us,
instead of the awful reverence due to the Cause of all substance and form,
all love and wisdom, a dangerously detached appreciation of an ingenuity
and benevolence meritorious in aim and often surprisingly successful in
contrivance.
The old teleology was more useful to science than to religion, and the
design-naturalists ought to be gratefully remembered by Biologists. Their
search for evidences led them to an eager study of adaptations and of
minute forms, a study such as we have now an incentive to in the theory of
Natural Selection. One hardly meets with the same ardour in microscopical
research until we come to modern workers. But the argument from Design
was never of great importance to faith. Still, to rid it of this character was
worth all the stress and anxiety of the gallant old war. If Darwin had done
nothing else for us, we are to-day deeply in his debt for this. The world is
not less venerable to us now, not less eloquent of the causing mind, rather
much more eloquent and sacred. But our wonder is not that "the underjaw
of the swine works under the ground" or in any or all of those particular
adaptations which Paley collected with so much skill, but that a purpose
transcending, though resembling, our own purposes, is everywhere
manifest; that what we live in is a whole, mutually sustaining, eventful and
beautiful, where the "dead" forces feed the energies of life, and life sustains
a stranger existence, able in some real measure to contemplate the whole, of
which, mechanically considered, it is a minor product and a rare ingredient.
Here, again, the change was altogether positive. It was not the escape of a
vessel in a storm with loss of spars and rigging, not a shortening of sail to
save the masts and make a port of refuge. It was rather the emergence from
narrow channels to an open sea. We had propelled the great ship, finding
purchase here and there for slow and uncertain movement. Now, in deep
water, we spread large canvas to a favouring breeze.
The scattered traces of design might be forgotten or obliterated. But the
broad impression of Order became plainer when seen at due distance and in
sufficient range of effect, and the evidence of love and wisdom in the
universe could be trusted more securely for the loss of the particular
calculation of their machinery.

Many other topics of faith are affected by modern biology. In some of these
we have learnt at present only a wise caution, a wise uncertainty. We stand
before the newly unfolded spectacle of suffering, silenced; with faith not
scientifically reassured but still holding fast certain other clues of
conviction. In many important topics we are at a loss. But in others, and
among them those I have mentioned, we have passed beyond this negative
state and find faith positively strengthened and more fully expressed.
We have gained also a language and a habit of thought more fit for the great
and dark problems that remain, less liable to damaging conflicts, equipped
for more rapid assimilation of knowledge. And by this change biology itself
is a gainer. For, relieved of fruitless encounters with popular religion, it may
advance with surer aim along the path of really scientific life-study which
was reopened for modern men by the publication of The Origin of Species.
Charles Darwin regretted that, in following science, he had not done "more
direct good"[237] to his fellow-creatures. He has, in fact, rendered
substantial service to interests bound up with the daily conduct and hopes of
common men; for his work has led to improvements in the preaching of the
Christian faith.

FOOTNOTES:
[218] F. R. Tennant: "The Being of God in the light of Physical Science," in
Essays on some theological questions of the day. London, 1905.
[219] Evolutionisme et Platonisme, pp. 45, 46, 47. Paris, 1908.
[220] Essays of Elia, "New Year's Eve," p. 41; Ainger's edition. London, 1899.
[221] Such an example is given in Baron F. von Hügel's recently finished book,
the result of thirty years' research: The Mystical Element of Religion, as studied
in Saint Catherine of Genoa and her Friends. London, 1908.
[222] G. Tyrrell, in Mediaevalism, has a chapter which is full of the important
moral element in a scientific attitude. "The only infallible guardian of truth is the
spirit of truthfulness." Mediaevalism, p. 182, London, 1908.
[223] Queen of the Air, Preface, p. vii. London, 1906.
[224] The scientific rank of its writer justifies the insertion of the following letter
from the late Sir John Burdon-Sanderson to me. In the lecture referred to I had
described the methods of Professor Moseley in teaching Biology as affording a
suggestion of the scientific treatment of religion.
Oxford,
April 30, 1902.
Dear Sir:
I feel that I must express to you my thanks for the discourse which I
had the pleasure of listening to yesterday afternoon.
I do not mean to say that I was able to follow all that you said as to
the identity of Method in the two fields of Science and Religion, but
I recognise that the "mysticism" of which you spoke gives us the
only way by which the two fields can be brought into relation.
Among much that was memorable, nothing interested me more than
what you said of Moseley.
No one, I am sure, knew better than you the value of his teaching
and in what that value consisted.
Yours faithfully,
J. BURDON-SANDERSON.
[225] H. P. Liddon, The Recovery of S. Thomas; a sermon preached in St. Paul's,
London, on April 23rd, 1882 (the Sunday after Darwin's death).
[226] Dr. Pusey (Unscience not Science adverse to Faith, 1878) writes: "The
questions as to 'species,' of what variations the animal world is capable, whether
the species be more or fewer, whether accidental variations may become

hereditary ... and the like, naturally fall under the province of science. In all
these questions Mr. Darwin's careful observations gained for him a deserved
approbation and confidence."
[227] Aristotle, in Bacon, quoted by Newman in his Idea of a University, p. 78.
London, 1873.
[228] Life and Letters and More Letters of Charles Darwin.
[229] Life and Letters, London, 1896. Thoughts on Religion, London, 1895.
Candid Examination of Theism, London, 1878.
[230] "Never in the history of man has so terrific a calamity befallen the race as
that which all who look may now (viz. in consequence of the scientific victory of
Darwin) behold advancing as a deluge black with destruction, resistless in might,
uprooting our most cherished hopes, engulphing our most precious creed, and
burying our highest life in mindless destruction."—A Candid Examination of
Theism, p. 51.
[231] Science and Christian Tradition. London, 1904.
[232] "No productiveness of the highest kind ... is in the power of anyone."—
Conversations of Goethe with Eckermann and Soret. London, 1850.
[233] Berthelot, Evolutionisme et Platonisme, Paris, 1908, p. 45.
[234] Times, 1892, passim.
[235] See Von Hartmann's Wahrheit und Irrthum in Darwinismus. Berlin, 1875.
[236] Hymn of the Church—

Rerum Deus tenax vigor,
Immotus in te permanens.

[237] Life and Letters, Vol. iii. p. 359.

IX

DARWINISM AND HISTORY
By J. B. Bury, LITT.D., LL.D.

Regius Professor of Modern History in the University of Cambridge

1. Evolution, and the principles associated with the Darwinian theory, could
not fail to exert a considerable influence on the studies connected with the
history of civilised man. The speculations which are known as "philosophy
of history," as well as the sciences of anthropology, ethnography, and
sociology (sciences which though they stand on their own feet are for the
historian auxiliary), have been deeply affected by these principles.
Historiographers, indeed, have with few exceptions made little attempt to
apply them; but the growth of historical study in the nineteenth century has
been determined and characterised by the same general principle which has
underlain the simultaneous developments of the study of nature, namely the
genetic idea. The "historical" conception of nature, which has produced the
history of the solar system, the story of the earth, the genealogies of telluric
organisms, and has revolutionised natural science, belongs to the same
order of thought as the conception of human history as a continuous,
genetic, causal process—a conception which has revolutionised historical
research and made it scientific. Before proceeding to consider the
application of evolutional principles, it will be pertinent to notice the rise of
this new view.
2. With the Greeks and Romans history had been either a descriptive record
or had been written in practical interests. The most eminent of the ancient
historians were pragmatical; that is, they regarded history as an instructress
in statesmanship, or in the art of war, or in morals. Their records reached
back such a short way, their experience was so brief, that they never
attained to the conception of continuous process, or realised the significance
of time; and they never viewed the history of human societies as a
phenomenon to be investigated for its own sake. In the middle ages there
was still less chance of the emergence of the ideas of progress and

development. Such notions were excluded by the fundamental doctrines of
the dominant religion which bounded and bound men's minds. As the
course of history was held to be determined from hour to hour by the
arbitrary will of an extra cosmic person, there could be no self-contained
causal development, only a dispensation imposed from without. And as it
was believed that the world was within no great distance from the end of
this dispensation, there was no motive to take much interest in
understanding the temporal, which was to be only temporary.
The intellectual movements of the fifteenth and sixteenth centuries prepared
the way for a new conception, but it did not emerge immediately. The
historians of the Renaissance period simply reverted to the ancient
pragmatical view. For Machiavelli, exactly as for Thucydides and Polybius,
the use of studying history was instruction in the art of politics. The
Renaissance itself was the appearance of a new culture, different from
anything that had gone before; but at the time men were not conscious of
this; they saw clearly that the traditions of classical antiquity had been lost
for a long period, and they were seeking to revive them, but otherwise they
did not perceive that the world had moved, and that their own spirit, culture,
and conditions were entirely unlike those of the thirteenth century. It was
hardly till the seventeenth century that the presence of a new age, as
different from the middle ages as from the ages of Greece and Rome, was
fully realised. It was then that the triple division of ancient, medieval, and
modern was first applied to the history of western civilisation. Whatever
objections may be urged against this division, which has now become
almost a category of thought, it marks a most significant advance in man's
view of his own past. He has become conscious of the immense changes in
civilisation which have come about slowly in the course of time, and history
confronts him with a new aspect. He has to explain how those changes have
been produced, how the transformations were effected. The appearance of
this problem was almost simultaneous with the rise of rationalism, and the
great historians and thinkers of the eighteenth century, such as
Montesquieu, Voltaire, Gibbon, attempted to explain the movement of
civilisation by purely natural causes. These brilliant writers prepared the
way for the genetic history of the following century. But in the spirit of the
Aufklärung, that eighteenth-century Enlightenment to which they belonged,
they were concerned to judge all phenomena before the tribunal of reason;

and the apotheosis of "reason" tended to foster a certain superior a priori
attitude, which was not favourable to objective treatment and was
incompatible with a "historical sense." Moreover the traditions of
pragmatical historiography had by no means disappeared.
3. In the first quarter of the nineteenth century the meaning of genetic
history was fully realised. "Genetic" perhaps is as good a word as can be
found for the conception which in this century was applied to so many
branches of knowledge in the spheres both of nature and of mind. It does
not commit us to the doctrine proper of evolution, nor yet to any
teleological hypothesis such as is implied in "progress." For history it meant
that the present condition of the human race is simply and strictly the result
of a causal series (or set of causal series)—a continuous succession of
changes, where each state arises causally out of the preceding; and that the
business of historians is to trace this genetic process, to explain each
change, and ultimately to grasp' the complete development of the life of
humanity. Three influential writers, who appeared at this stage and helped
to initiate a new period of research, may specially be mentioned. Ranke in
1824 definitely repudiated the pragmatical view which ascribes to history
the duties of an instructress, and with no less decision renounced the
function, assumed by the historians of the Aufklärung, to judge the past; it
was his business, he said, merely to show how things really happened.
Niebuhr was already working in the same spirit and did more than any other
writer to establish the principle that historical transactions must be related
to the ideas and conditions of their age. Savigny about the same time
founded the "historical school" of law. He sought to show that law was not
the creation of an enlightened will, but grew out of custom and was
developed by a series of adaptations and rejections, thus applying the
conception of evolution. He helped to diffuse the notion that all the
institutions of a society or a nation are as closely interconnected as the parts
of a living organism.
4. The conception of the history of man as a causal development meant the
elevation of historical inquiry to the dignity of a science. Just as the study of
bees cannot become scientific so long as the student's interest in them is
only to procure honey or to derive moral lessons from the labours of "the
little busy bee," so the history of human societies cannot become the object
of pure scientific investigation so long as man estimates its value in

pragmatical scales. Nor can it become a science until it is conceived as
lying entirely within a sphere in which the law of cause and effect has
unreserved and unrestricted dominion. On the other hand, once history is
envisaged as a causal process, which contains within itself the explanation
of the development of man from his primitive state to the point which he
has reached, such a process necessarily becomes the object of scientific
investigation and the interest in it is scientific curiosity.
At the same time, the instruments were sharpened and refined. Here Wolf, a
philologist with historical instinct, was a pioneer. His Prolegomena to
Homer (1795) announced new modes of attack. Historical investigation was
soon transformed by the elaboration of new methods.
5. "Progress" involves a judgment of value, which is not involved in the
conception of history as a genetic process. It is also an idea distinct from
that of evolution. Nevertheless it is closely related to the ideas which
revolutionised history at the beginning of the last century; it swam into
men's ken simultaneously; and it helped effectively to establish the notion
of history as a continuous process and to emphasise the significance of
time. Passing over earlier anticipations, I may point to a Discours of Turgot
(1750), where history is presented as a process in which "the total mass of
the human race" "marches continually though sometimes slowly to an ever
increasing perfection." That is a clear statement of the conception which
Turgot's friend Condorcet elaborated in the famous work, published in
1795, Esquisse d'un tableau historique des progrès de l'esprit humain. This
work first treated with explicit fulness the idea to which a leading role was
to fall in the ideology of the nineteenth century. Condorcet's book reflects
the triumphs of the Tiers état, whose growing importance had also inspired
Turgot; it was the political changes in the eighteenth century which led to
the doctrine, emphatically formulated by Condorcet, that the masses are the
most important element in the historical process. I dwell on this because,
though Condorcet had no idea of evolution, the predominant importance of
the masses was the assumption which made it possible to apply evolutional
principles to history. And it enabled Condorcet himself to maintain that the
history of civilisation, a progress still far from being complete, was a
development conditioned by general laws.

6. The assimilation of society to an organism, which was a governing notion
in the school of Savigny, and the conception of progress, combined to
produce the idea of an organic development, in which the historian has to
determine the central principle or leading character. This is illustrated by the
apotheosis of democracy in Tocqueville's Démocratie en Amérique, where
the theory is maintained that "the gradual and progressive development of
equality is at once the past and the future of the history of men." The same
two principles are combined in the doctrine of Spencer (who held that
society is an organism, though he also contemplated its being what he calls
a "super-organic aggregate"),[238] that social evolution is a progressive
change from militarism to industrialism.
7. The idea of development assumed another form in the speculations of
German idealism. Hegel conceived the successive periods of history as
corresponding to the ascending phases or ideas in the self-evolution of his
Absolute Being. His Lectures on the Philosophy of History were published
in 1837 after his death. His philosophy had a considerable effect, direct and
indirect, on the treatment of history by historians, and although he was
superficial and unscientific himself in dealing with historical phenomena,
he contributed much towards making the idea of historical development
familiar. Ranke was influenced, if not by Hegel himself, at least by the
Idealistic philosophies of which Hegel's was the greatest. He was inclined
to conceive the stages in the process of history as marked by incarnations,
as it were, of ideas, and sometimes speaks as if the ideas were independent
forces, with hands and feet. But while Hegel determined his ideas by a
priori logic, Ranke obtained his by induction—by a strict investigation of
the phenomena; so that he was scientific in his method and work, and was
influenced by Hegelian prepossessions only in the kind of significance
which he was disposed to ascribe to his results. It is to be noted that the
theory of Hegel implied a judgment of value; the movement was a progress
towards perfection.
8. In France, Comte approached the subject from a different side, and
exercised, outside Germany, a far wider influence than Hegel. The 4th
volume of his Cours de philosophie positive, which appeared in 1839,
created sociology and treated history as a part of this new science, namely
as "social dynamics." Comte sought the key for unfolding historical
development, in what he called the social-psychological point of view, and

he worked out the two ideas which had been enunciated by Condorcet: that
the historian's attention should be directed not, as hitherto, principally to
eminent individuals, but to the collective behaviour of the masses, as being
the most important element in the process; and that, as in nature, so in
history, there are general laws, necessary and constant, which condition the
development. The two points are intimately connected, for it is only when
the masses are moved into the foreground that regularity, uniformity, and
law can be conceived as applicable. To determine the social-psychological
laws which have controlled the development is, according to Comte, the
task of sociologists and historians.
9. The hypothesis of general laws operative in history was carried further in
a book which appeared in England twenty years later and exercised an
influence in Europe far beyond its intrinsic merit, Buckle's History of
Civilisation in England (1857-61). Buckle owed much to Comte, and
followed him, or rather outdid him, in regarding intellect as the most
important factor conditioning the upward development of man, so that
progress, according to him, consisted in the victory of the intellectual over
the moral laws.
10. The tendency of Comte and Buckle to assimilate history to the sciences
of nature by reducing it to general "laws," derived stimulus and plausibility
from the vista offered by the study of statistics, in which the Belgian
Quetelet, whose book Sur l'homme appeared in 1835, discerned endless
possibilities. The astonishing uniformities which statistical inquiry
disclosed led to the belief that it was only a question of collecting a
sufficient amount of statistical material, to enable us to predict how a given
social group will act in a particular case. Bourdeau, a disciple of this school,
looks forward to the time when historical science will become entirely
quantitative. The actions of prominent individuals, which are generally
considered to have altered or determined the course of things, are obviously
not amenable to statistical computation or explicable by general laws.
Thinkers like Buckle sought to minimise their importance or explain them
away.
11. These indications may suffice to show that the new efforts to interpret
history which marked the first half of the nineteenth century were governed
by conceptions closely related to those which were current in the field of

natural science and which resulted in the doctrine of evolution. The genetic
principle, progressive development, general laws, the significance of time,
the conception of society as an organic aggregate, the metaphysical theory
of history as the self-evolution of spirit,—all these ideas show that
historical inquiry had been advancing independently on somewhat parallel
lines to the sciences of nature. It was necessary to bring this out in order to
appreciate the influence of Darwinism.
12. In the course of the dozen years which elapsed between the appearances
of The Origin of Species (observe that the first volume of Buckle's work
was published just two years before) and of The Descent of Man (1871), the
hypothesis of Lamarck that man is the co-descendant with other species of
some lower extinct form was admitted to have been raised to the rank of an
established fact by most thinkers whose brains were not working under the
constraint of theological authority.
One important effect of the discovery of this fact (I am not speaking now of
the Darwinian explanation) was to assign to history a definite place in the
coordinated whole of knowledge, and relate it more closely to other
sciences. It had indeed a defined logical place in systems such as Hegel's
and Comte's; but Darwinism certified its standing convincingly and without
more ado. The prevailing doctrine that man was created ex abrupto had
placed history in an isolated position, disconnected with the sciences of
nature. Anthropology, which deals with the animal anthropos, now comes
into line with zoology, and brings it into relation with history.[239] Man's
condition at the present day is the result of a series of transformations,
going back to the most primitive phase of society, which is the ideal
(unattainable) beginning of history. But that beginning had emerged without
any breach of continuity from a development which carries us back to a
quadrimane ancestor, still further back (according to Darwin's conjecture) to
a marine animal of the ascidian type, and then through remoter periods to
the lowest form of organism. It is essential in this theory that though links
have been lost there was no break in the gradual development; and this
conception of a continuous progress in the evolution of life, resulting in the
appearance of uncivilised Anthropos, helped to reinforce, and increase a
belief in, the conception of the history of civilised Anthropos as itself also a
continuous progressive development.

13. Thus the diffusion of the Darwinian theory of the origin of man, by
emphasising the idea of continuity and breaking down the barriers between
the human and animal kingdoms, has had an important effect in establishing
the position of history among the sciences which deal with telluric
development. The perspective of history is merged in a larger perspective of
development. As one of the objects of biology is to find the exact steps in
the genealogy of man from the lowest organic form, so the scope of history
is to determine the stages in the unique causal series from the most
rudimentary to the present state of human civilisation.
It is to be observed that the interest in historical research implied by this
conception need not be that of Comte. In the Positive Philosophy history is
part of sociology; the interest in it is to discover the sociological laws. In
the view of which I have just spoken, history is permitted to be an end in
itself; the reconstruction of the genetic process is an independent interest.
For the purpose of the reconstruction, sociology, as well as physical
geography, biology, psychology, is necessary; the sociologist and the
historian play into each other's hands; but the object of the former is to
establish generalisations; the aim of the latter is to trace in detail a singular
causal sequence.
14. The success of the evolutional theory helped to discredit the assumption
or at least the invocation of transcendent causes. Philosophically of course
it is compatible with theism, but historians have for the most part desisted
from invoking the naive conception of a "god in history" to explain
historical movements. A historian may be a theist; but, so far as his work is
concerned, this particular belief is otiose. Otherwise indeed (as was
remarked above) history could not be a science; for with a deus ex machina
who can be brought on the stage to solve difficulties scientific treatment is a
farce. The transcendent element had appeared in a more subtle form through
the influence of German philosophy. I noticed how Ranke is prone to refer
to ideas as if they were transcendent existences manifesting themselves in
the successive movements of history. It is intelligible to speak of certain
ideas as controlling, in a given period,—for instance, the idea of nationality;
but from the scientific point of view, such ideas have no existence outside
the minds of individuals and are purely psychical forces; and a historical
"idea," if it does not exist in this form, is merely a way of expressing a
synthesis of the historian himself.

15. From the more general influence of Darwinism on the place of history
in the system of human knowledge, we may turn to the influence of the
principles and methods by which Darwin explained development. It had
been recognised even by ancient writers (such as Aristotle and Polybius)
that physical circumstances (geography, climate) were factors conditioning
the character and history of a race or society. In the sixteenth century Bodin
emphasised these factors, and many subsequent writers took them into
account. The investigations of Darwin, which brought them into the
foreground, naturally promoted attempts to discover in them the chief key
to the growth of civilisation. Comte had expressly denounced the notion
that the biological methods of Lamarck could be applied to social man.
Buckle had taken account of natural influences, but had relegated them to a
secondary plane, compared with psychological factors. But the Darwinian
theory made it tempting to explain the development of civilisation in terms
of "adaptation to environment," "struggle for existence," "natural selection,"
"survival of the fittest," etc.[240]
The operation of these principles cannot be denied. Man is still an animal,
subject to zoological as well as mechanical laws. The dark influence of
heredity continues to be effective; and psychical development had begun in
lower organic forms,—perhaps with life itself. The organic and the social
struggles for existence are manifestations of the same principle.
Environment and climatic influence must be called in to explain not only
the differentiation of the great racial sections of humanity, but also the
varieties within these sub-species and, it may be, the assimilation of distinct
varieties. Ritter's Anthropogeography has opened a useful line of research.
But on the other hand, it is urged that, in explaining the course of history,
these principles do not take us very far, and that it is chiefly for the
primitive ultra-prehistoric period that they can account for human
development. It may be said that, so far as concerns the actions and
movements of men which are the subject of recorded history, physical
environment has ceased to act mechanically, and in order to affect their
actions must affect their wills first; and that this psychical character of the
causal relations substantially alters the problem. The development of human
societies, it may be argued, derives a completely new character from the
dominance of the conscious psychical element, creating as it does new
conditions (inventions, social institutions, etc.) which limit and counteract

the operation of natural selection, and control and modify the influence of
physical environment. Most thinkers agree now that the chief clews to the
growth of civilisation must be sought in the psychological sphere. Imitation,
for instance, is a principle which is probably more significant for the
explanation of human development than natural selection. Darwin himself
was conscious that his principles had only a very restricted application in
this sphere, as is evident from his cautious and tentative remarks in the 5th
chapter of his Descent of Man. He applied natural selection to the growth of
the intellectual faculties and of the fundamental social instincts, and also to
the differentiation of the great races or "sub-species" (Caucasian, African,
etc.) which differ in anthropological character.[241]
16. But if it is admitted that the governing factors which concern the student
of social development are of the psychical order, the preliminary success of
natural science in explaining organic evolution by general principles
encouraged sociologists to hope that social evolution could be explained on
general principles also. The idea of Condorcet, Buckle, and others, that
history could be assimilated to the natural sciences was powerfully
reinforced, and the notion that the actual historical process, and every social
movement involved in it, can be accounted for by sociological
generalisations, so-called "laws," is still entertained by many, in one form or
another. Dissentients from this view do not deny that the generalisations at
which the sociologist arrives by the comparative method, by the analysis of
social factors, and by psychological deduction may be an aid to the
historian; but they deny that such uniformities are laws or contain an
explanation of the phenomena. They can point to the element of chance
coincidence. This element must have played a part in the events of organic
evolution, but it has probably in a larger measure helped to determine
events in social evolution. The collision of two unconnected sequences may
be fraught with great results. The sudden death of a leader or a marriage
without issue, to take simple cases, has again and again led to permanent
political consequences. More emphasis is laid on the decisive actions of
individuals, which cannot be reduced under generalisations and which
deflect the course of events. If the significance of the individual will had
been exaggerated to the neglect of the collective activity of the social
aggregate before Condorcet, his doctrine tended to eliminate as unimportant
the roles of prominent men, and by means of this elimination it was

possible to found sociology. But it may be urged that it is patent on the face
of history that its course has constantly been shaped and modified by the
wills of individuals,[242] which are by no means always the expression of
the collective will; and that the appearance of such personalities at the given
moments is not a necessary outcome of the conditions and cannot be
deduced. Nor is there any proof that, if such and such an individual had not
been born, some one else would have arisen to do what he did. In some
cases there is no reason to think that what happened need ever have come to
pass. In other cases, it seems evident that the actual change was inevitable,
but in default of the man who initiated and guided it, it might have been
postponed, and, postponed or not, might have borne a different cachet. I
may illustrate by an instance which has just come under my notice. Modern
painting was founded by Giotto, and the Italian expedition of Charles VIII,
near the close of the sixteenth century, introduced into France the fashion of
imitating Italian painters. But for Giotto and Charles VIII, French painting
might have been very different. It may be said that "if Giotto had not
appeared, some other great imitator would have played a role analogous to
his, and that without Charles VIII there would have been the commerce
with Italy, which in the long run would have sufficed to place France in
relation with Italian artists. But the equivalent of Giotto might have been
deferred for a century and probably would have been different; and
commercial relations would have required ages to produce the rayonnement
imitatif of Italian art in France, which the expedition of the royal adventurer
provoked in a few years."[243] Instances furnished by political history are
simply endless. Can we conjecture how events would have moved if the son
of Philip of Macedon had been an incompetent? The aggressive action of
Prussia which astonished Europe in 1740 determined the subsequent history
of Germany; but that action was anything but inevitable; it depended
entirely on the personality of Frederick the Great.
Hence it may be argued that the action of individual wills is a determining
and disturbing factor, too significant and effective to allow history to be
grasped by sociological formulae. The types and general forms of
development which the sociologist attempts to disengage can only assist the
historian in understanding the actual course of events. It is in the special
domains of economic history and Culturgeschichte which have come to the