EVOLUTION IN MODERN
BY HAECKEL, THOMSON, WEISMANN
THE MODERN LIBRARY
PUBLISHERS :: :: NEW YORK
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I DARWIN'S PREDECESSORS
J. Arthur Thomson, Professor of Natural History in the University of Aberdeen
II The Selection Theory
August Weismann, Professor of Zoology in the University of Freiburg (Baden)
III HEREDITY AND VARIATION IN MODERN LIGHTS
W. Bateson, Professor of Biology in the University of Cambridge
IV "THE DESCENT OF MAN"
G. Schwalbe, Professor of Anatomy in the University of Strassburg
V CHARLES DARWIN AS AN ANTHROPOLOGIST
Ernst Haeckel, Professor of Zoology in the University of Jena
VI MENTAL FACTORS IN EVOLUTION
C. Lloyd Morgan, Professor of Psychology at University College, Bristol
VII THE INFLUENCE OF THE CONCEPTION OF EVOLUTION ON MODERN PHILOSOPHY
H. Hoeffding, Professor of Philosophy in the University of Copenhagen
VIII THE INFLUENCE OF DARWIN UPON RELIGIOUS THOUGHT
Rev. P. H. Waggett
IX DARWINISM AND HISTORY
J. B. Bury, Regious Professor of Modern History in the University of Cambridge
X DARWINISM AND SOCIOLOGY
C. Bougle, Professor of Social Philosophy in the University of Toulouse, and Deputy-Professor at the Sorbonne, Paris
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EVOLUTION IN MODERN THOUGHT
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 AEtiology 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, 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." But the fundamental idea of one stage giving origin to another was absent. As the blue AEgean 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 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, 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, "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 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.
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 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.
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 Linnaeus (1707-1778), protagonist of the counter-doctrine of the fixity of species, 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'enchainment des etres."
Erasmus Darwin (1731-1802), probably influenced by Buffon, was another firm evolutionist, and the outline of his argument in the Zoonomia 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 [Greek: proton oon], or first great egg, produced by night, that is, whose origin is involved in obscurity, and animated by [Greek: Eros], 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."
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.
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; 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—"pensees de la jeunesse, executees par l'age mur"—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.
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 a 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.
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 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:
"Le cours ordinaire de la nature vivante, est en general toujours constant, toujours le meme; son mouvement, toujours regulier, roule sur deux points inebranlables: l'un, la fecondite sans bornes donnee a toutes les especes; l'autre, les obstacles sans nombre qui reduisent cette fecondite a une mesure determinee et ne laissent en tout temps qu'a peu pres la meme quantite d'individus de chaque espece" ... "Les especes les moins parfaites, les plus delicates, les plus pesantes, les moins agissantes, les moins armees, etc., ont deja disparu ou disparaitront.".
Erasmus Darwin 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."
Lamarck (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 (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 (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. Etienne 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." 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."
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." 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." 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." 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.'" 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."
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 Etienne 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 (a la convenance) the new environment."
Prof. E. B. Poulton 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, 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.
[Footnote 1: Columbia University Biological Series, Vol. I. New York and London, 1894. We must acknowledge our great indebtedness to this fine piece of work.]
[Footnote 2: op. cit. p. 41.]
[Footnote 3: See G. J. Romanes, "Aristotle as a Naturalist," Contemporary Review, Vol. lix. p. 275, 1891; G. Pouchet, La Biologie Aristotelique, Paris, 1885; E. Zeller, A History of Greek Philosophy, London, 1881, and "Ueber die griechischen Vorgaenger Darwin's," Abhandl. Berlin Akad. 1878, pp. 111-124.]
[Footnote 4: op. cit. p. 81.]
[Footnote 5: op. cit. p. 87.]
[Footnote 6: See Brock, "Die Stellung Kant's zur Deszendenztheorie," Biol. Centralbl. viii. 1889, pp. 641-648. Fritz Schultze, Kant und Darwin, Jena, 1875.]
[Footnote 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.]
[Footnote 8: Experimental Evolution. London, 1892. Chap. I. p. 14.]
[Footnote 9: See J. Arthur Thomson, The Science of Life. London, 1899, Chap. XVI. "Evolution of Evolution Theory."]
[Footnote 10: See Carus Sterne (Ernst Krause), Die allgemeine Weltanschauung in ihrer historischen Entwickelung. Stuttgart, 1889. Chapter entitled "Bestaendigkeit oder Veraenderlichkeit der Naturwesen."]
[Footnote 11: Zoonomia, or the Laws of Organic Life, 2 vols. London, 1794; Osborn, op. cit. p. 145.]
[Footnote 12: See Alpheus S. Packard, Lamarck, the Founder of Evolution, His Life and Work, with Translations of his writings on Organic Evolution. London, 1901.]
[Footnote 13: See Edward Clodd, Pioneers of Evolution, London, p. 161, 1897.]
[Footnote 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.]
[Footnote 15: See Prof. W. A. Locy's Biology and its Makers. New York, 1908. Part II. "The Doctrine of Organic Evolution."]
[Footnote 16: Presidential Address to the British Association meeting at Dublin in 1908.]
[Footnote 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.]
[Footnote 18: op. cit. p. 136.]
[Footnote 19: See Ernest Krause and Charles Darwin, Erasmus Darwin, London, 1879.]
[Footnote 20: Osborn, op. cit. p. 142.]
[Footnote 21: See E. Perrier, La Philosophie Zoologique avant Darwin, Paris, 1884; A. de Quatrefages, Darwin et ses Precurseurs Francais, Paris, 1870; Packard, op. cit.; also Claus, Lamarck als Begruender der Descendenzlehre, Wien, 1888; Haeckel, Natural History of Creation, Eng. transl. London, 1879; Lang, Zur Charakteristik der Forschungswege von Lamarck und Darwin, Jena, 1889.]
[Footnote 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.]
[Footnote 23: See Haeckel, Die Naturanschauung von Darwin, Goethe und Lamarck, Jena, 1882.]
[Footnote 24: Origin of Species (6th edit.), p. xvii.]
[Footnote 25: The Life and Letters of Charles Darwin, Vol. 1. p. 83. London, 1887.]
[Footnote 26: A. R. Wallace, My Life, a Record of Events and Opinions, London, 1905, Vol. 1, p. 232.]
[Footnote 27: My Life, Vol. 1. p. 361.]
[Footnote 28: P. Geddes. article "Biology." Chambers's Encyclopaedia.]
[Footnote 29: Origin of Species (6th edit.), p. xv.]
[Footnote 30: Life and Letters, II, p. 301.]
[Footnote 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.]
[Footnote 32: See Prof. Patrick Geddes's article "Variation and Selection," Encyclopaedia Britannica (9th edit.) 1888.]
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 cooeperation 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. Koelliker, 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, and I am the less inclined to return to it here that it now appears 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 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 (Haecker). The same thing has been shown by Schuett 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 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 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 cooeperation 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.
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 role 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 cooeperated 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?
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 cooeperation 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 cooeperation 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.