By J. Arthur Thomson Professor of Natural History in the University of AberdeenIn 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 (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 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, Aristotle's views of Nature 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, 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, 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 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 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, Erasmus Darwin (1731-1802), probably influenced by Buffon, was another firm evolutionist, and the outline of his argument in the Zoonomia 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; 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. 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. 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 "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 Lamarck Treviranus Goethe 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." 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." 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 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 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 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 Hints of the idea of Natural Selection have been detected elsewhere. James Watt, 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 |