The Evolution of Man — Volume 2

by Ernst Haeckel

The line of argument we followed in this explanation of the ontogenetic facts was simply a consistent application of the biogenetic law. In this we have throughout taken strict account of the distinction between palingenetic and cenogenetic phenomena. Palingenesis (or "synoptic development") alone enables us to draw conclusions from the observed embryonic form to the stem-form preserved by heredity. Such inference becomes more or less precarious when there has been cenogenesis, or disturbance of development, owing to fresh adaptations. We cannot understand embryonic development unless we appreciate this very important distinction. Here we stand at the very limit that separates the older and the new science or philosophy of nature. The whole of the results of recent morphological research compel us irresistibly to recognise the biogenetic law and its far-reaching consequences. These are, it is true, irreconcilable with the legends and doctrines of former days, that have been impressed on us by religious education. But without the biogenetic law, without the distinction between palingenesis and cenogenesis, and without the theory of evolution on which we base it, it is quite impossible to understand the facts of organic development; without them we cannot cast the faintest gleam of explanation over this marvellous field of phenomena. But when we recognise the causal correlation of ontogeny and phylogeny expressed in this law, the wonderful facts of embryology are susceptible of a very simple explanation; they are found to be the necessary mechanical effects of the evolution of the stem, determined by the laws of heredity and adaptation. The correlative action of these laws under the universal influence of the struggle for existence, or—as we may say in a word, with Darwin—"natural selection," is entirely adequate to explain the whole process of embryology in the light of phylogeny. It is the chief merit of Darwin that he explained by his theory of selection the correlation of the laws of heredity and adaptation that Lamarck had recognised, and pointed out the true way to reach a causal interpretation of evolution.

The phenomenon that it is most imperative to recognise in this connection is the inheritance of functional variations. Jean Lamarck was the first to appreciate its fundamental importance in 1809, and we may therefore justly give the name of Lamarckism to the theory of descent he based on it. Hence the radical opponents of the latter have very properly directed their attacks chiefly against the former. One of the most distinguished and most narrow-minded of these opponents, Wilhelm His, affirms very positively that "characteristics acquired in the life of the individual are not inherited."

The inheritance of acquired characters is denied, not only by thorough opponents of evolution, but even by scientists who admit it and have contributed a good deal to its establishment, especially Weismann, Galton, Ray Lankester, etc. Since 1884 the chief opponent has been August Weismann, who has rendered the greatest service in the development of Darwin's theory of selection. In his work on The Continuity of the Germ-plasm, and in his recent excellent Lectures on the Theory of Descent (1902), he has with great success advanced the opinion that "only those characters can be transmitted to subsequent generations that were contained in rudimentary form in the embryo." However, this germ-plasm theory, with its attempt to explain heredity, is merely a "provisional molecular hypothesis"; it is one of those metaphysical speculations that attribute the evolutionary phenomena exclusively to internal causes, and regard the influence of the environment as insignificant. Herbert Spencer, Theodor Eimer, Lester Ward, Hering, and Zehnder have pointed out the untenable consequences of this position. I have given my view of it in the tenth edition of the History of Creation (pages 192 and 203). I hold, with Lamarck and Darwin, that the hereditary transmission of acquired characters is one of the most important phenomena in biology, and is proved by thousands of morphological and physiological experiences. It is an indispensable foundation of the theory of evolution.

Of the many and weighty arguments for the truth of this conception of evolution I will for the moment merely point to the invaluable evidence of dysteleology, the science of rudimentary organs. We cannot insist too often or too strongly on the great morphological significance of these remarkable organs, which are completely useless from the physiological point of view. We find some of these useless parts, inherited from our lower vertebrate ancestors, in every system of organs in man and the higher Vertebrates. Thus we find at once on the skin a scanty and rudimentary coat of hair, only fully developed on the head, under the shoulders, and at a few other parts of the body. The short hairs on the greater part of the body are quite useless and devoid of physiological value; they are the last relic of the thicker hairy coat of our simian ancestors. The sensory apparatus presents a series of most remarkable rudimentary organs. We have seen that the whole of the shell of the external ear, with its cartilages, muscles, and skin, is in man a useless appendage, and has not the physiological importance that was formerly ascribed to it. It is the degenerate remainder of the pointed, freely moving, and more advanced mammal ear, the muscles of which we still have, but cannot work them. We found at the inner corner of our eye a small, curious, semi-lunar fold that is of no use whatever to us, and is only interesting as the last relic of the nictitating membrane, the third, inner eye-lid that had a distinct physiological purpose in the ancient sharks, and still has in many of the Amniotes.

The motor apparatus, in both the skeleton and muscular systems, provides a number of interesting dysteleological arguments. I need only recall the projecting tail of the human embryo, with its rudimentary caudal vertebrae and muscles; this is totally useless in man, but very interesting as the degenerate relic of the long tail of our simian ancestors. From these we have also inherited various bony processes and muscles, which were very useful to them in climbing trees, but are useless to us. At various points of the skin we have cutaneous muscles which we never use—remnants of a strongly-developed cutaneous muscle in our lower mammal ancestors. This "panniculus carnosus" had the function of contracting and creasing the skin to chase away the flies, as we see every day in the horse. Another relic in us of this large cutaneous muscle is the frontal muscle, by which we knit our forehead and raise our eye-brows; but there is another considerable relic of it, the large cutaneous muscle in the neck (platysma myoides), over which we have no voluntary control.

Not only in the systems of animal organs, but also in the vegetal apparatus, we find a number of rudimentary organs, many of which we have already noticed. In the alimentary apparatus there are the thymus-gland and the thyroid gland, the seat of goitre and the relic of a ciliated groove that the Tunicates and Acrania still have in the gill-pannier; there is also the vermiform appendix to the caecum. In the vascular system we have a number of useless cords which represent relics of atrophied vessels that were once active as blood-canals—the ductus Botalli between the pulmonary artery and the aorta, the ductus venosus Arantii between the portal vein and the vena cava, and many others. The many rudimentary organs in the urinary and sexual apparatus are particularly interesting. These are generally developed in one sex and rudimentary in the other. Thus the spermaducts are formed from the Wolffian ducts in the male, whereas in the female we have merely rudimentary traces of them in Gaertner's canals. On the other hand, in the female the oviducts and womb are developed from the Mullerian ducts, while in the male only the lowest ends of them remain as the "male womb" (vesicula prostatica). Again, the male has in his nipples and mammary glands the rudiments of organs that are usually active only in the female.

A careful anatomic study of the human frame would disclose to us numbers of other rudimentary organs, and these can only be explained on the theory of evolution. Robert Wiedersheim has collected a large number of them in his work on The Human Frame as a Witness to its Past. They are some of the weightiest proofs of the truth of the mechanical conception and the strongest disproofs of the teleological view. If, as the latter demands, man or any other organism had been designed and fitted for his life-purposes from the start and brought into being by a creative act, the existence of these rudimentary organs would be an insoluble enigma; it would be impossible to understand why the Creator had put this useless burden on his creatures to walk a path that is in itself by no means easy. But the theory of evolution gives the simplest possible explanation of them. It says: The rudimentary organs are parts of the body that have fallen into disuse in the course of centuries; they had definite functions in our animal ancestors, but have lost their physiological significance. On account of fresh adaptations they have become superfluous, but are transmitted from generation to generation by heredity, and gradually atrophy.

We have inherited not only these rudimentary parts, but all the organs of our body, from the mammals—proximately from the apes. The human body does not contain a single organ that has not been inherited from the apes. In fact, with the aid of our biogenetic law we can trace the origin of our various systems of organs much further, down to the lowest stages of our ancestry. We can say, for instance, that we have inherited the oldest organs of the body, the external skin and the internal coat of the alimentary system, from the Gastraeads; the nervous and muscular systems from the Platodes; the vascular system, the body-cavity, and the blood from the Vermalia; the chorda and the branchial gut from the Prochordonia; the articulation of the body from the Acrania; the primitive skull and the higher sense-organs from the Cyclostomes; the limbs and jaws from the Selachii; the five-toed foot from the Amphibia; the palate from the Reptiles; the hairy coat, the mammary glands, and the external sexual organs from the Pro-mammals. When we formulated "the law of the ontogenetic connection of systematically related forms," and determined the relative age of organs, we saw how it was possible to draw phylogenetic conclusions from the ontogenetic succession of systems of organs.

With the aid of this important law and of comparative anatomy we were also enabled to determine "man's place in nature," or, as we put it, assign to man his position in the classification of the animal kingdom. In recent zoological classification the animal world is divided into twelve stems or phyla, and these are broadly sub-divided into about sixty classes, and these classes into at least 300 orders. In his whole organisation man is most certainly, in the first place, a member of one of these stems, the vertebrate stem; secondly, a member of one particular class in this stem, the Mammals; and thirdly, of one particular order, the order of Primates. He has all the characteristics that distinguish the Vertebrates from the other eleven animal stems, the Mammals from the other sixty classes, and the Primates from the 300 other orders of the animal kingdom. We may turn and twist as we like, but we cannot get over this fact of anatomy and classification. Of late years this fact has given rise to a good deal of discussion, and especially of controversy as to the particular anatomic relationship of man to the apes. The most curious opinions have been advanced on this "ape-question," or "pithecoid-theory." It is as well, therefore, to go into it once more and distinguish the essential from the unessential. (Cf. Chapter 2.23.)

We start from the undisputed fact that man is in any case—whether we accept or reject his special blood-relationship to the apes—a true mammal; in fact, a placental mammal. This fundamental fact can be proved so easily at any moment from comparative anatomy that it has been universally admitted since the separation of the Placentals from the lower mammals (Marsupials and Monotremes). But for every consistent subscriber to the theory of evolution it must follow at once that man descends from a common stem-form with all the other Placentals, the stem-ancestor of the Placentals, just as we must admit a common mesozoic ancestor of all the mammals. This is, however, to settle decisively the great and burning question of man's place in nature, whether or no we go on to admit a nearer or more distant relationship to the apes. Whether man is or is not a member of the ape-order (or, if you prefer, the primate-order.) in the phylogenetic sense, in any case his direct blood-relationship to the rest of the mammals, and especially the Placentals, is established. It is possible that the affinities of the various orders of mammals to each other are different from what we hypothetically assume to-day. But, in any case, the common descent of man and all the other mammals from one stem-form is beyond question. This long-extinct Promammal was probably evolved from Proreptiles during the Triassic period, and must certainly be regarded as the monotreme and oviparous ancestor of ALL the mammals.