Part 14

"In the evolutionary appreciation of the facts of embryology we must take particular care to distinguish sharply and clearly between the primary, palingenetic evolutionary processes and the secondary, cenogenetic processes. The palingenetic phenomena, or embryonic recapitulations, are due to heredity, to the transmission of characters from one generation to another. They enable us to draw direct inferences in regard to corresponding structures in the development of the species (e.g. the chorda or the branchial arches in all vertebrate embryos). The cenogenetic phenomena, on the other hand, or the embryonic variations, cannot be traced to inheritance from a mature ancestor, but are due to the adaption of the embryo or the larva to certain conditions of its individual development (e.g. the amnion, the allantois, and the vitelline arteries in the embryos of the higher vertebrates). These cenogenetic phenomena are later additions; we must not infer from them that there were corresponding processes in the ancestral history, and hence they are apt to mislead."

The fundamental importance of these facts of comparative anatomy, atavism, and the rudimentary organs, was pointed out by Darwin in the first part of his classic work, The Descent of Man and Selection in Relation to Sex (1871). In the "General summary and conclusion" (chap. xxi.) he was able to say, with perfect justice: "He who is not content to look, like a savage, at the phenomena of nature as disconnected, cannot any longer believe that man is the work of a separate act of creation. He will be forced to admit that the close resemblance of the embryo of man to that, for instance, of a dog--the construction of his skull, limbs, and whole frame on the same plan with that of other mammals, independently of the uses to which the parts may be put--the occasional reappearance of various structures, for instance of several muscles, which man does not normally possess, but which are common to the Quadrumana--and a crowd of analogous facts--all point in the plainest manner to the conclusion that man is the co-descendant with other mammals of a common progenitor."

These few lines of Darwin's have a greater scientific value than hundreds of those so-called "anthropological treatises," which give detailed descriptions of single organs, or mathematical tables with series of numbers and what are claimed to be "exact analyses," but are devoid of synoptic conclusions and a philosophical spirit.

Charles Darwin is not generally recognised as a great anthropologist, nor does the school of modern anthropologists regard him as a leading authority. In Germany, especially, the great majority of the members of the anthropological societies took up an attitude of hostility to him from the very beginning of the controversy in 1860. The Descent of Man was not merely rejected, but even the discussion of it was forbidden on the ground that it was "unscientific."

The centre of this inveterate hostility for thirty years--especially after 1877--was Rudolph Virchow of Berlin, the leading investigator in pathological anatomy, who did so much for the reform of medicine by his establishment of cellular pathology in 1858. As a prominent representative of "exact" or "descriptive" anthropology, and lacking a broad equipment in comparative anatomy and ontogeny, he was unable to accept the theory of descent. In earlier years, and especially during his splendid period of activity at Wuerzburg (1848-1856), he had been a consistent free-thinker, and had in a number of able articles (collected in his Gesammelte Abhandlungen) upheld the unity of human nature, the inseparability of body and spirit. In later years at Berlin, where he was more occupied with political work and sociology (especially after 1866), he abandoned the positive monistic position for one of agnosticism and scepticism, and made concessions to the dualistic dogma of a spiritual world apart from the material frame.

In the course of a Scientific Congress at Munich in 1877 the conflict of these antithetic views of nature came into sharp relief. At this memorable Congress I had undertaken to deliver the first address (September 18th) on the subject of "Modern evolution in relation to the whole of science." I maintained that Darwin's theory not only solved the great problem of the origin of species, but that its implications, especially in regard to the nature of man, threw considerable light on the whole of science, and on anthropology in particular. The discovery of the real origin of man by evolution from a long series of mammal ancestors threw light on his place in nature in every respect, as Huxley had already shown in his excellent lectures of 1863. Just as all the organs and tissues of the human body had originated from those of the nearest related mammals, certain ape-like forms, so we were bound to conclude that his mental qualities also had been derived from those of his extinct primate ancestor.

This monistic view of the origin and nature of man, which is now admitted by nearly all who have the requisite acquaintance with biology, and approach the subject without prejudice, encountered a sharp opposition at that time. The opposition found its strongest expression in an address that Virchow delivered at Munich four days afterwards (September 22nd), on "The freedom of science in the modern State." He spoke of the theory of evolution as an unproved hypothesis, and declared that it ought not to be taught in the schools, because it was dangerous to the State. "We must not," he said, "teach that man has descended from the ape or any other animal." When Darwin, usually so lenient in his judgment, read the English translation of Virchow's speech, he expressed his disapproval in strong terms. But the great authority that Virchow had--an authority well founded in pathology and sociology--and his prestige as president of the German Anthropological Society, had the effect of preventing any member of the Society from raising serious opposition to him for thirty years. Numbers of journals and treatises repeated his dogmatic statement: "It is quite certain that man has descended neither from the ape nor from any other animal." In this he persisted till his death in 1902. Since that time the whole position of German anthropology has changed. The question is no longer whether man was created by a distinct supernatural act or evolved from other mammals, but to which line of the animal hierarchy we must look for the actual series of ancestors. The interested reader will find an account of this "battle of Munich" (1877) in my three Berlin lectures (April, 1905), Der Kampf um die Entwickelungs-Gedanken.

The main points in our genealogical tree were clearly recognised by Darwin in the sixth chapter of the Descent of Man. Lowly organised fishes, like the lancelot (Amphioxus), are descended from lower invertebrates resembling the larvae of an existing Tunicate (Appendicularia). From these primitive fishes were evolved higher fishes of the ganoid type and others of the type of Lepidosiren (Dipneusta). It is a very small step from these to the Amphibia:

"In the class of animals the steps are not difficult to conceive which led from the ancient Monotremata to the ancient Marsupials; and from these to the early progenitors of the placental mammals. We may thus ascend to the Lemuridae; and the interval is not very wide from these to the Simiadae. The Simiadae then branched off into two great stems, the New World and Old World monkeys; and from the latter, at a remote period, Man, the wonder and glory of the Universe, proceeded."

In these few lines Darwin clearly indicated the way in which we were to conceive our ancestral series within the vertebrates. It is fully confirmed by all the arguments of comparative anatomy and embryology, of palaeontology and physiology; and all the research of the subsequent forty years have gone to establish it. The deep interest in geology which Darwin maintained throughout his life and his complete knowledge of palaeontology enabled him to grasp the fundamental importance of the palaeontological record more clearly than anthropologists and zoologists usually do.

There has been much debate in subsequent decades whether Darwin himself maintained that man was descended from the ape, and many writers have sought to deny it. But the lines I have quoted verbatim from the conclusion of the sixth chapter of the Descent of Man (1871) leave no doubt that he was as firmly convinced of it as was his great precursor Jean Lamarck in 1809. Moreover, Darwin adds, with particular explicitness, in the "general summary and conclusion" (chap. xxi.) of that standard work:

"By considering the embryological structure of man--the homologies which he presents with the lower animals,--the rudiments which he retains,--and the reversions to which he is liable, we can partly recall in imagination the former condition of our early progenitors; and can approximately place them in their proper place in the zoological series. We thus learn that man is descended from a hairy, tailed quadruped, probably arboreal in its habits, and an inhabitant of the Old World. This creature, if its whole structure had been examined by a naturalist, would have been classed amongst the Quadrumana, as surely as the still more ancient progenitor of the Old and New World monkeys."

These clear and definite lines leave no doubt that Darwin--so critical and cautious in regard to important conclusions--was quite as firmly convinced of the descent of man from the apes (the Catarrhinae, in particular) as Lamarck was in 1809 and Huxley in 1863.

It is to be noted particularly that, in these and other observations on the subject, Darwin decidedly assumes the monophyletic origin of the mammals, including man. It is my own conviction that this is of the greatest importance. A number of difficult questions in regard to the development of man, in respect of anatomy, physiology, psychology, and embryology, are easily settled if we do not merely extend our progonotaxis to our nearest relatives, the anthropoid apes and the tailed monkeys from which these have descended, but go further back and find an ancestor in the group of the Lemuridae, and still further back to the Marsupials and Monotremata. The essential identity of all the Mammals in point of anatomical structure and embryonic development--in spite of their astonishing differences in external appearance and habits of life--is so palpably significant that modern zoologists are agreed in the hypothesis that they have all sprung from a common root, and that this root may be sought in the earlier Palaeozoic Amphibia.

The fundamental importance of this comparative morphology of the Mammals, as a sound basis of scientific anthropology, was recognised just before the beginning of the nineteenth century, when Lamarck first emphasised (1794) the division of the animal kingdom into Vertebrates and Invertebrates. Even thirteen years earlier (1781), when Goethe made a close study of the mammal skeleton in the Anatomical Institute at Jena, he was intensely interested to find that the composition of the skull was the same in man as in the other mammals. His discovery of the os inter-maxillare in man (1784), which was contradicted by most of the anatomists of the time, and his ingenious "vertebral theory of the skull," were the splendid fruit of his morphological studies. They remind us how Germany's greatest philosopher and poet was for many years ardently absorbed in the comparative anatomy of man and the mammals, and how he divined that their wonderful identity in structure was no mere superficial resemblance, but pointed to a deep internal connection. In my Generelle Morphologie (1866), in which I published the first attempts to construct phylogenetic trees, I have given a number of remarkable theses of Goethe, which may be called "phyletic prophecies." They justify us in regarding him as a precursor of Darwin.

In the ensuing forty years I have made many conscientious efforts to penetrate further along that line of anthropological research that was opened up by Goethe, Lamarck, and Darwin. I have brought together the many valuable results that have constantly been reached in comparative anatomy, physiology, ontogeny, and palaeontology, and maintained the effort to reform the classification of animals and plants in an evolutionary sense. The first rough drafts of pedigrees that were published in the Generelle Morphologie have been improved time after time in the ten editions of my Natuerlich Schoepfungsgeschichte (1868-1902). A sounded basis for my phyletic hypotheses, derived from a discriminating combination of the three great records--morphology, ontogeny, and palaeontology--was provided in the three volumes of my Systematische Phylogenie (1894 Protists and Plants, 1895 Vertebrates, 1896 Invertebrates).

In my Anthropogenie I endeavoured to employ all the known facts of comparative ontogeny (embryology) for the purpose of completing my scheme of human phylogeny (evolution). I attempted to sketch the historical development of each organ of the body, beginning with the most elementary structures in the germ-layers of the Gastraea. At the same time I drew up a corrected statement of the most important steps in the line of our ancestral series.

At the fourth International Congress of Zoology at Cambridge (August 26th, 1898) I delivered an address on "Our present knowledge of the Descent of Man." It was translated into English, enriched with many valuable notes and additions, by my friend and pupil in earlier days Dr. Hans Gadow (Cambridge), and published under the title: The Last Link: our present knowledge of the Descent of Man The determination of the chief animal forms that occur in the line of our ancestry is there restricted to thirty types, and these are distributed in six main groups.

The first half of this "Progonotaxis hominis," which has no support from fossil evidence, comprises three groups: (i) Protista (unicellular organisms, 1-5): (ii) Invertebrate Metazoa (Coelenteria 6-8, Vermalia 9-11): (iii) Monorrhine Vertebrates (Acrania 12-13, Cyclostoma 14-15). The second half, which is based on fossil records, also comprises three groups: (iv) Palaeozoic cold-blooded Craniota (Fishes 16-18, Amphibia 19, Reptiles 20): (v) Mesozoic Mammals (Monotrema 21, Marsupialia 22, Mallotheria 23): (vi) Cenozoic Primates (Lemuridae 24-25, Tailed Apes 26-27, Anthropomorpha 28-30). An improved and enlarged edition of this hypothetic "Progonotaxis hominis" was published in 1908, in my essay Unsere Ahnenreihe.

If I have succeeded in furthering, in some degree, by these anthropological works, the solution of the great problem of Man's place in nature, and particularly in helping to trace the definite stages in our ancestral series, I owe the success, not merely to the vast progress that biology has made in the last half century, but largely to the luminous example of the great investigators who have applied themselves to the problem, with so much assiduity and genius, for a century and a quarter--I mean Goethe and Lamarck, Gegenbaur and Huxley, but, above all, Charles Darwin. It was the great genius of Darwin that first brought together that symmetrical temple of scientific knowledge, the theory of descent. It was Darwin who put the crown on the edifice by his theory of natural selection. Not until this broad inductive law was firmly established was it possible to vindicate the special conclusion, the descent of man from a series of other Vertebrates. By his illuminating discovery Darwin did more for anthropology than thousands of those writers, who are more specifically titled anthropologists, have done by their technical treatises. We may, indeed, say that it is not merely as an exact observer and ingenious experimenter, but as a distinguished anthropologist and far-seeing thinker, that Darwin takes his place among the greatest men of science of the nineteenth century.

To appreciate fully the immortal merit of Darwin in connection with anthropology, we must remember that not only did his chief work, The Origin of Species, which opened up a new era in natural history in 1859, sustain the most virulent and widespread opposition for a lengthy period, but even thirty years later, when its principles were generally recognised and adopted, the application of them to man was energetically contested by many high scientific authorities. Even Alfred Russel Wallace, who discovered the principle of natural selection independently in 1858, did not concede that it was applicable to the higher mental and moral qualities of man. Dr. Wallace still holds a spiritualist and dualist view of the nature of man, contending that he is composed of a material frame (descended from the apes) and an immortal immaterial soul (infused by a higher power). This dual conception, moreover, is still predominant in the wide circles of modern theology and metaphysics, and has the general and influential adherence of the more conservative classes of society.

In strict contradiction to this mystical dualism, which is generally connected with teleology and vitalism, Darwin always maintained the complete unity of human nature, and showed convincingly that the psychological side of man was developed, in the same way as the body, from the less advanced soul of the anthropoid ape, and, at a still more remote period, from the cerebral functions of the older vertebrates. The eighth chapter of the Origin of Species, which is devoted to instinct, contains weighty evidence that the instincts of animals are subject, like all other vital processes, to the general laws of historic development. The special instincts of particular species were formed by adaptation, and the modifications thus acquired were handed on to posterity by heredity; in their formation and preservation natural selection plays the same part as in the transformation of every other physiological function. The higher moral qualities of civilised man have been derived from the lower mental functions of the uncultivated barbarians and savages, and these in turn from the social instincts of the mammals. This natural and monistic psychology of Darwin's was afterwards more fully developed by his friend George Romanes in his excellent works Mental Evolution in Animals and Mental Evolution in Man.

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

FOOTNOTES:

MENTAL FACTORS IN EVOLUTION

BY C. LLOYD MORGAN, LLD., F.R.S

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

Throughout his treatment, mental evolution was for Darwin incidental to and contributory to organic evolution. For specialised research in comparative and genetic psychology, as an independent field of investigation, he had neither the time nor the requisite training. None the less his writings and the spirit of his work have exercised a profound influence on this department of evolutionary thought. And, for those who follow Darwin's lead, mental evolution is still in a measure subservient to organic evolution. Mental processes are the accompaniments or concomitants of the functional activity of specially differentiated parts of the organism. They are in some way dependent on physiological and physical conditions. But though they are not physical in their nature, and though it is difficult or impossible to conceive that they are physical in their origin, they are, for Darwin and his followers, factors in the evolutionary process in its physical or organic aspect. By the physiologist within his special and well-defined universe of discourse they may be properly regarded as epiphenomena; but by the naturalist in his more catholic survey of nature they cannot be so regarded, and were not so regarded by Darwin. Intelligence has contributed to evolution of which it is in a sense a product.