The Evolution of Man, V.1., Ernst Haeckel [ebook pdf reader for pc .TXT] 📗

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 inheritance from earlier species, but are due to the adaptation of the foetus, or the infant-form, to certain conditions of its embryonic development. These cenogenetic phenomena are foreign or later additions; they allow us to draw no direct inference whatever as to corresponding processes in our ancestral history, but rather hinder us from doing so.

This careful discrimination between the primary or palingenetic processes and the secondary or cenogenetic is of great importance for the purposes of the scientific history of a species, which has to draw conclusions from the available facts of embryology, comparative anatomy, and paleontology, as to the processes in the formation of the species in the remote past. It is of the same importance to the student of evolution as the careful distinction between genuine and spurious texts in the works of an ancient writer, or the purging of the real text from interpolations and alterations, is for the student of philology. It is true that this distinction has not yet been fully appreciated by many scientists. For my part, I regard it as the first condition for forming any just idea of the evolutionary process, and I believe that we must, in accordance with it, divide embryology into two sections--palingenesis, or the science of recapitulated forms; and cenogenesis, or the science of supervening structures.

To give at once a few examples from the science of man's origin in illustration of this important distinction, I may instance the following processes in the embryology of man, and of all the higher vertebrates, as palingenetic: the formation of the two primary germinal layers and of the primitive gut, the undivided structure of the dorsal nerve-tube, the appearance of a simple axial rod between the medullary tube and the gut, the temporary formation of the gill-clefts and arches, the primitive kidneys, and so on.* (* All these, and the following structures, will be fully described in later chapters.--Translator.) All these, and many other important structures, have clearly been transmitted by a steady heredity from the early ancestors of the mammal, and are, therefore, direct indications of the presence of similar structures in the history of the stem. On the other hand, this is certainly not the case with the following embryonic forms, which we must describe as cenogenetic processes: the formation of the yelk-sac, the allantois, the placenta, the amnion, the serolemma, and the chorion--or, generally speaking, the various foetal membranes and the corresponding changes in the blood vessels. Further instances are: the dual structure of the heart cavity, the temporary division of the plates of the primitive vertebrae and lateral plates, the secondary closing of the ventral and intestinal walls, the formation of the navel, and so on. All these and many other phenomena are certainly not traceable to similar structures in any earlier and completely-developed ancestral form, but have arisen simply by adaptation to the peculiar conditions of embryonic life (within the foetal membranes). In view of these facts, we may now give the following more precise expression to our chief law of biogeny: The evolution of the foetus (or ontogenesis) is a condensed and abbreviated recapitulation of the evolution of the stem (or phylogenesis); and this recapitulation is the more complete in proportion as the original development (or palingenesis) is preserved by a constant heredity; on the other hand, it becomes less complete in proportion as a varying adaptation to new conditions increases the disturbing factors in the development (or cenogenesis).

The cenogenetic alterations or distortions of the original palingenetic course of development take the form, as a rule, of a gradual displacement of the phenomena, which is slowly effected by adaptation to the changed conditions of embryonic existence during the course of thousands of years. This displacement may take place as regards either the position or the time of a phenomenon.

The great importance and strict regularity of the time-variations in embryology have been carefully studied recently by Ernest Mehnert, in his Biomechanik (Jena, 1898). He contends that our biogenetic law has not been impaired by the attacks of its opponents, and goes on to say: "Scarcely any piece of knowledge has contributed so much to the advance of embryology as this; its formulation is one of the most signal services to general biology. It was not until this law passed into the flesh and blood of investigators, and they had accustomed themselves to see a reminiscence of ancestral history in embryonic structures, that we witnessed the great progress which embryological research has made in the last two decades." The best proof of the correctness of this opinion is that now the most fruitful work is done in all branches of embryology with the aid of this biogenetic law, and that it enables students to attain every year thousands of brilliant results that they would never have reached without it.

It is only when one appreciates the cenogenetic processes in relation to the palingenetic, and when one takes careful account of the changes which the latter may suffer from the former, that the radical importance of the biogenetic law is recognised, and it is felt to be the most illuminating principle in the science of evolution. In this task of discrimination it is the silver thread in relation to which we can arrange all the phenomena of this realm of marvels--the "Ariadne thread," which alone enables us to find our way through this labyrinth of forms. Hence the brothers Sarasin, the zoologists, could say with perfect justice, in their study of the evolution of the Ichthyophis, that "the great biogenetic law is just as important for the zoologist in tracing long-extinct processes as spectrum analyses is for the astronomer."

Even at an earlier period, when a correct acquaintance with the evolution of the human and animal frame was only just being obtained--and that is scarcely eighty years ago!--the greatest astonishment was felt at the remarkable similarity observed between the embryonic forms, or stages of foetal development, in very different animals; attention was called even then to their close resemblance to certain fully-developed animal forms belonging to some of the lower groups. The older scientists (Oken, Treviranus, and others) knew perfectly well that these lower forms in a sense illustrated and fixed, in the hierarchy of the animal world, a temporary stage in the evolution of higher forms. The famous anatomist Meckel spoke in 1821 of a "similarity between the development of the embryo and the series of animals." Baer raised the question in 1828 how far, within the vertebrate type, the embryonic forms of the higher animals assume the permanent shapes of members of lower groups. But it was impossible fully to understand and appreciate this remarkable resemblance at that time. We owe our capacity to do this to the theory of descent; it is this that puts in their true light the action of heredity on the one hand and adaptation on the other. It explains to us the vital importance of their constant reciprocal action in the production of organic forms. Darwin was the first to teach us the great part that was played in this by the ceaseless struggle for existence between living things, and to show how, under the influence of this (by natural selection), new species were produced and maintained solely by the interaction of heredity and adaptation. It was thus Darwinism that first opened our eyes to a true comprehension of the supremely important relations between the two parts of the science of organic evolution--Ontogeny and Phylogeny.

Heredity and adaptation are, in fact, the two constructive physiological functions of living things; unless we understand these properly we can make no headway in the study of evolution. Hence, until the time of Darwin no one had a clear idea of the real nature and causes of embryonic development. It was impossible to explain the curious series of forms through which the human embryo passed; it was quite unintelligible why this strange succession of animal-like forms appeared in the series at all. It had previously been generally assumed that the man was found complete in all his parts in the ovum, and that the development consisted only in an unfolding of the various parts, a simple process of growth. This is by no means the case. On the contrary, the whole process of the development of the individual presents to the observer a connected succession of different animal-forms; and