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- The Evolution of Man, V.1. - 6/54 -


and only at a late stage a form that has a resemblance to the ape; until at last the definite human form emerges and closes the series of transformations. These suggestive facts are, as I said, still almost unknown to the general public--so completely unknown that, if one casually mentions them, they are called in question or denied outright as fairy-tales. Everybody knows that the butterfly emerges from the pupa, and the pupa from a quite different thing called a larva, and the larva from the butterfly's egg. But few besides medical men are aware that MAN, in the course of his individual formation, passes through a series of transformations which are not less surprising and wonderful than the familiar metamorphoses of the butterfly.

The mere description of these remarkable changes through which man passes during his embryonic life should arouse considerable interest. But the mind will experience a far keener satisfaction when we trace these curious facts to their causes, and when we learn to behold in them natural phenomena which are of the highest importance throughout the whole field of human knowledge. They throw light first of all on the "natural history of creation," then on psychology, or "the science of the soul," and through this on the whole of philosophy. And as the general results of every branch of inquiry are summed up in philosophy, all the sciences come in turn to be touched and influenced more or less by the study of the evolution of man.

But when I say that I propose to present here the most important features of these phenomena and trace them to their causes, I take the term, and I interpret my task, in a very much wider sense than is usual. The lectures which have been delivered on this subject in the universities during the last half-century are almost exclusively adapted to medical men. Certainly, the medical man has the greatest interest in studying the origin of the human body, with which he is daily occupied. But I must not give here this special description of the embryonic processes such as it has hitherto been given, as most of my readers have not studied anatomy, and are not likely to be entrusted with the care of the adult organism. I must content myself with giving some parts of the subject only in general outline, and must not enter upon all the marvellous, but very intricate and not easily described, details that are found in the story of the development of the human frame. To understand these fully a knowledge of anatomy is needed. I will endeavour to be as plain as possible in dealing with this branch of science. Indeed, a sufficient general idea of the course of the embryonic development of man can be obtained without going too closely into the anatomic details. I trust we may be able to arouse the same interest in this delicate field of inquiry as has been excited already in other branches of science; though we shall meet more obstacles here than elsewhere.

The story of the evolution of man, as it has hitherto been expounded to medical students, has usually been confined to embryology--more correctly, ontogeny--or the science of the development of the individual human organism. But this is really only the first part of our task, the first half of the story of the evolution of man in that wider sense in which we understand it here. We must add as the second half--as another and not less important and interesting branch of the science of the evolution of the human stem--phylogeny: this may be described as the science of the evolution of the various animal forms from which the human organism has been developed in the course of countless ages. Everybody now knows of the great scientific activity that was occasioned by the publication of Darwin's Origin of Species in 1859. The chief direct consequence of this publication was to provoke a fresh inquiry into the origin of the human race, and this has proved beyond question our gradual evolution from the lower species. We give the name of "Phylogeny" to the science which describes this ascent of man from the lower ranks of the animal world. The chief source that it draws upon for facts is "Ontogeny," or embryology, the science of the development of the individual organism. Moreover, it derives a good deal of support from paleontology, or the science of fossil remains, and even more from comparative anatomy, or morphology.

These two branches of our science--on the one side ontogeny or embryology, and on the other phylogeny, or the science of race-evolution--are most vitally connected. The one cannot be understood without the other. It is only when the two branches fully co-operate and supplement each other that "Biogeny" (or the science of the genesis of life in the widest sense) attains to the rank of a philosophic science. The connection between them is not external and superficial, but profound, intrinsic, and causal. This is a discovery made by recent research, and it is most clearly and correctly expressed in the comprehensive law which I have called "the fundamental law of organic evolution," or "the fundamental law of biogeny." This general law, to which we shall find ourselves constantly recurring, and on the recognition of which depends one's whole insight into the story of evolution, may be briefly expressed in the phrase: "The history of the foetus is a recapitulation of the history of the race"; or, in other words, "Ontogeny is a recapitulation of phylogeny." It may be more fully stated as follows: The series of forms through which the individual organism passes during its development from the ovum to the complete bodily structure is a brief, condensed repetition of the long series of forms which the animal ancestors of the said organism, or the ancestral forms of the species, have passed through from the earliest period of organic life down to the present day.

The causal character of the relation which connects embryology with stem-history is due to the action of heredity and adaptation. When we have rightly understood these, and recognised their great importance in the formation of organisms, we can go a step further and say: Phylogenesis is the mechanical cause of ontogenesis.* (* The term "genesis," which occurs throughout, means, of course, "birth" or origin. From this we get: Biogeny = the origin of life (bios); Anthropogeny = the origin of man (anthropos); Ontogeny = the origin of the individual (on); Phylogeny = the origin of the species (phulon); and so on. In each case the term may refer to the process itself, or to the science describing the process.--Translator.) In other words, the development of the stem, or race, is, in accordance with the laws of heredity and adaptation, the cause of all the changes which appear in a condensed form in the evolution of the foetus.

The chain of manifold animal forms which represent the ancestry of each higher organism, or even of man, according to the theory of descent, always form a connected whole. We may designate this uninterrupted series of forms with the letters of the alphabet: A, B, C, D, E, etc., to Z. In apparent contradiction to what I have said, the story of the development of the individual, or the ontogeny of most organisms, only offers to the observer a part of these forms; so that the defective series of embryonic forms would run: A, B, D, F, H, K, M, etc.; or, in other cases, B, D, H, L, M, N, etc. Here, then, as a rule, several of the evolutionary forms of the original series have fallen out. Moreover, we often find--to continue with our illustration from the alphabet--one or other of the original letters of the ancestral series represented by corresponding letters from a different alphabet. Thus, instead of the Roman B and D, we often have the Greek Beta and Delta. In this case the text of the biogenetic law has been corrupted, just as it had been abbreviated in the preceding case. But, in spite of all this, the series of ancestral forms remains the same, and we are in a position to discover its original complexion.

In reality, there is always a certain parallel between the two evolutionary series. But it is obscured from the fact that in the embryonic succession much is wanting that certainly existed in the earlier ancestral succession. If the parallel of the two series were complete, and if this great fundamental law affirming the causal connection between ontogeny and phylogeny in the proper sense of the word were directly demonstrable, we should only have to determine, by means of the microscope and the dissecting knife, the series of forms through which the fertilised ovum passes in its development; we should then have before us a complete picture of the remarkable series of forms which our animal ancestors have successively assumed from the dawn of organic life down to the appearance of man. But such a repetition of the ancestral history by the individual in its embryonic life is very rarely complete. We do not often find our full alphabet. In most cases the correspondence is very imperfect, being greatly distorted and falsified by causes which we will consider later. We are thus, for the most part, unable to determine in detail, from the study of its embryology, all the different shapes which an organism's ancestors have assumed; we usually--and especially in the case of the human foetus--encounter many gaps. It is true that we can fill up most of these gaps satisfactorily with the help of comparative anatomy, but we cannot do so from direct embryological observation. Hence it is important that we find a large number of lower animal forms to be still represented in the course of man's embryonic development. In these cases we may draw our conclusions with the utmost security as to the nature of the ancestral form from the features of the form which the embryo momentarily assumes.

To give a few examples, we can infer from the fact that the human ovum is a simple cell that the first ancestor of our species was a tiny unicellular being, something like the amoeba. In the same way, we know, from the fact that the human foetus consists, at the first, of two simple cell-layers (the gastrula), that the gastraea, a form with two such layers, was certainly in the line of our ancestry. A later human embryonic form (the chordula) points just as clearly to a worm-like ancestor (the prochordonia), the nearest living relation of which is found among the actual ascidiae. To this succeeds a most important embryonic stage (acrania), in which our headless foetus presents, in the main, the structure of the lancelet. But we can only indirectly and approximately, with the aid of comparative anatomy and ontogeny, conjecture what lower forms enter into the chain of our ancestry between the gastraea and the chordula, and between this and the lancelet. In the course of the historical development many intermediate structures have gradually fallen out, which must certainly have been represented in our ancestry. But, in spite of these many, and sometimes very appreciable, gaps, there is no contradiction between the two successions. In fact, it is the chief purpose of this work to prove the real harmony and the original parallelism of the two. I hope to show, on a substantial basis of facts, that we can draw most important conclusions as to our genealogical tree from the actual and easily-demonstrable series of embryonic changes. We shall then be in a position to form a general idea of the wealth of animal forms which have figured in the direct line of our ancestry in the lengthy history of organic life.

In this evolutionary appreciation of the facts of embryology we must, of course, take particular care to distinguish sharply and clearly between the primitive, palingenetic (or ancestral) evolutionary processes and those due to cenogenesis.* (* Palingenesis = new birth, or re-incarnation (palin = again, genesis or genea = development); hence its application to the phenomena which are recapitulated by heredity from earlier ancestral forms. Cenogenesis = foreign or negligible development (kenos and genea); hence, those phenomena which come later in the story of life to disturb the inherited structure, by a fresh adaptation to environment.--Translator.) By palingenetic processes, or embryonic recapitulations, we understand all those phenomena in the development of the individual which are transmitted from one generation to another by heredity, and which, on that account, allow us to draw direct inferences as to corresponding structures in the development of the species. On the other hand, we give the name of cenogenetic processes, or embryonic variations, to all those phenomena in the foetal development that cannot be traced to


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