THE EVOLUTION OF MAN
Volume I

CHAPTER II
THE OLDER EMBRYOLOGY

It is in many ways useful, on entering upon the study of any science, to cast a glance at its historical development. The saying that “everything is best understood in its growth” has a distinct application to science. While we follow its gradual development we get a clearer insight into its aims and objects. Moreover, we shall see that the present condition of the science of human evolution, with all its characteristics, can only be rightly understood when we examine its historical growth. This task will, however, not detain us long. The study of man’s evolution is one of the latest branches of natural science, whether you consider the embryological or the phylogenetic section of it.

Apart from the few germs of our science which we find in classical antiquity, and which we shall notice presently, we may say that it takes its definite rise, as a science, in the year 1759, when one of the greatest German scientists, Caspar Friedrich Wolff, published his Theoria generationis. That was the foundation-stone of the science of animal embryology. It was not until fifty years later, in 1809, that Jean Lamarck published his Philosophie Zoologique—the first effort to provide a base for the theory of evolution; and it was another half-century before Darwin’s work appeared (in 1859), which we may regard as the first scientific attainment of this aim. But before we go further into this solid establishment of evolution, we must cast a brief glance at that famous philosopher and scientist of antiquity, who stood alone in this, as in many other branches of science, for more than 2000 years: the “father of Natural History,” Aristotle.

The extant scientific works of Aristotle deal with many different sides of biological research; the most comprehensive of them is his famous History of Animals. But not less interesting is the smaller work, On the Generation of Animals (Peri zoon geneseos). This work treats especially of embryonic development, and it is of great interest as being the earliest of its kind and the only one that has come down to us in any completeness from classical antiquity.

Aristotle studied embryological questions in various classes of animals, and among the lower groups he learned many most remarkable facts which we only rediscovered between 1830 and 1860. It is certain, for instance, that he was acquainted with the very peculiar mode of propagation of the cuttlefishes, or cephalopods, in which a yelk-sac hangs out of the mouth of the fœtus. He knew, also, that embryos come from the eggs of the bee even when they have not been fertilised. This “parthenogenesis” (or virgin-birth) of the bees has only been established in our time by the distinguished zoologist of Munich, Siebold. He discovered that male bees come from the unfertilised, and female bees only from the fertilised, eggs. Aristotle further states that some kinds of fishes (of the genus serranus) are hermaphrodites, each individual having both male and female organs and being able to fertilise itself; this, also, has been recently confirmed. He knew that the embryo of many fishes of the shark family is attached to the mother’s body by a sort of placenta, or nutritive organ very rich in blood; apart from these, such an arrangement is only found among the higher mammals and

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man. This placenta of the shark was looked upon as legendary for a long time, until Johannes Müller proved it to be a fact in 1839. Thus a number of remarkable discoveries were found in Aristotle’s embryological work, proving a very good acquaintance of the great scientist—possibly helped by his predecessors—with the facts of ontogeny, and a great advance upon succeeding generations in this respect.

In the case of most of these discoveries he did not merely describe the fact, but added a number of observations on its significance. Some of these theoretical remarks are of particular interest, because they show a correct appreciation of the nature of the embryonic processes. He conceives the development of the individual as a new formation, in the course of which the various parts of the body take shape successively. When the human or animal frame is developed in the mother’s body, or separately in an egg, the heart—which he regards as the starting-point and centre of the organism—must appear first. Once the heart is formed the other organs arise, the internal ones before the external, the upper (those above the diaphragm) before the lower (or those beneath the diaphragm). The brain is formed at an early stage, and the eyes grow out of it. These observations are quite correct. And, if we try to form some idea from these data of Aristotle’s general conception of the embryonic process, we find a dim prevision of the theory which Wolff showed 2000 years afterwards to be the correct view. It is significant, for instance, that Aristotle denied the eternity of the individual in any respect. He said that the species or genus, the group of similar individuals, might be eternal, but the individual itself is temporary. It comes into being in the act of procreation, and passes away at death.

During the 2000 years after Aristotle no progress whatever was made in general zoology, or in embryology in particular. People were content to read, copy, translate, and comment on Aristotle. Scarcely a single independent effort at research was made in the whole of the period. During the Middle Ages the spread of strong religious beliefs put formidable obstacles in the way of independent scientific investigation. There was no question of resuming the advance of biology. Even when human anatomy began to stir itself once more in the sixteenth century, and independent research was resumed into the structure of the developed body, anatomists did not dare to extend their inquiries to the unformed body, the embryo, and its development. There were many reasons for the prevailing horror of such studies. It is natural enough, when we remember that a Bull of Boniface VIII excommunicated every man who ventured to dissect a human corpse. If the dissection of a developed body were a crime to be thus punished, how much more dreadful must it have seemed to deal with the embryonic body still enclosed in the womb, which the Creator himself had decently veiled from the curiosity of the scientist! The Christian Church, then putting many thousands to death for unbelief, had a shrewd presentiment of the menace that science contained against its authority. It was powerful enough to see that its rival did not grow too quickly.

It was not until the Reformation broke the power of the Church, and a refreshing breath of the spirit dissolved the icy chains that bound science, that anatomy and embryology, and all the other branches of research, could begin to advance once more. However, embryology lagged far behind anatomy. The first works on embryology appear at the beginning of the sixteenth century. The Italian anatomist, Fabricius ab Aquapendente, a professor at Padua, opened the advance. In his two books (De formato fœtu, 1600, and De formatione fœtus, 1604) he published the older illustrations and descriptions of the embryos of man and other mammals, and of the hen. Similar imperfect illustrations were given by Spigelius (De formato fœtu, 1631), and by Needham (1667) and his more famous compatriot, Harvey (1652), who discovered the circulation of the blood in the animal body and formulated the important principle, Omne vivum ex vivo (all life comes from pre-existing life). The Dutch scientist, Swammerdam, published in his Bible of Nature the earliest observations on the embryology of the frog and the division of its egg-yelk. But the most important embryological studies in the sixteenth century were those of the famous Italian, Marcello Malpighi, of Bologna, who led the way both in zoology and botany. His treatises, De formatione pulli and De ovo incubato (1687), contain the first consistent description of the development of the chick in the fertilised egg.

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Here I ought to say a word about the important part played by the chick in the growth of our science. The development of the chick, like that of the young of all other birds, agrees in all its main features with that of the other chief vertebrates, and even of man. The three highest classes of vertebrates—mammals, birds, and reptiles (lizards, serpents, tortoises, etc.)—have from the beginning of their embryonic development so striking a resemblance in all the chief points of structure, and especially in their first forms, that for a long time it is impossible to distinguish between them. We have known now for some time that we need only examine the embryo of a bird, which is the easiest to get at, in order to learn the typical mode of development of a mammal (and therefore of man). As soon as scientists began to study the human embryo, or the mammal-embryo generally, in its earlier stages about the middle and end of the seventeenth century, this important fact was very quickly discovered. It is both theoretically and practically of great value. As regards the theory of evolution, we can draw the most weighty inferences from this similarity between the embryos of widely different classes of animals. But for the practical purposes of embryological research the discovery is invaluable, because we can fill up the gaps in our imperfect knowledge of the embryology of the mammals from the more thoroughly studied embryology of the bird. Hens’ eggs are easily to be had in any quantity, and the development of the chick may be followed step by step in artificial incubation. The development of the mammal is much more difficult to follow, because here the embryo is not detached and enclosed in a large egg, but the tiny ovum remains in the womb until the growth is completed. Hence, it is very difficult to keep up sustained observation of the various stages in any great extent, quite apart from such extrinsic considerations as the cost, the technical difficulties, and many other obstacles which we encounter when we would make an extensive study of the fertilised mammal. The chicken has, therefore, always been the chief object of study in this connection. The excellent incubators we now have enable us to observe it in any quantity and at any stage of development, and so follow the whole course of its formation step by step.

By the end of the seventeenth century Malpighi had advanced as far as it was possible to do with the imperfect microscope of his time in the embryological study of the chick. Further progress was arrested until the instrument and the technical methods should be improved. The vertebrate embryos are so small and delicate in their earlier stages that you cannot go very far into the study of them without a good microscope and other technical aid. But this substantial improvement of the microscope and the other apparatus did not take place until the beginning of the nineteenth century.

Embryology made scarcely any advance in the first half of the eighteenth century, when the systematic natural history of plants and animals received so great an impulse through the publication of Linné’s famous Systema Naturæ. Not until 1759 did the genius arise who was to give it an entirely new character, Caspar Friedrich Wolff. Until then embryology had been occupied almost exclusively in unfortunate and misleading efforts to build up theories on the imperfect empirical material then available.

The theory which then prevailed, and remained in favour throughout nearly the whole of the eighteenth century, was commonly called at that time “the evolution theory”; it is better to describe it as “the preformation theory.”1 Its chief point is this: There is no new formation of structures in the embryonic development of any organism, animal or plant, or even of man; there is only a growth, or unfolding, of parts which have been constructed or pre-formed from all eternity, though on a very small scale and closely packed together. Hence, every living germ contains all the organs and parts of the body, in the form and arrangement they will present later, already within it, and thus the whole embryological process is merely an evolution in the literal sense of the word, or an unfolding, of parts that were pre-formed and folded up in it. So, for instance, we find in the hen’s egg not merely a simple cell, that divides and subdivides and forms germinal layers, and at last, after all kinds of variation and cleavage and reconstruction, brings forth

1. This theory is usually known as the “evolution theory” in Germany, in contradistinction to the “epigenesis theory.” But as it is the latter that is called the “evolution theory” in England, France, and Italy, and “evolution” and “epigenesis” are taken to be synonymous, it seems better to call the first the “pre-formation theory.”

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the body of the chick; but there is in every egg from the first a complete chicken, with all its parts made and neatly packed. These parts are so small or so transparent that the microscope cannot detect them. In the hatching, these parts merely grow larger, and spread out in the normal way.

When this theory is consistently developed it becomes a “scatulation theory.”1 According to its teaching, there was made in the beginning one couple or one individual of each species of animal or plant; but this one individual contained the germs of all the other individuals of the same species who should ever come to life. As the age of the earth was generally believed at that time to be fixed by the Bible at 5000 or 6000 years, it seemed possible to calculate how many individuals of each species had lived in the period, and so had been packed inside the first being that was created. The theory was consistently extended to man, and it was affirmed that our common parent Eve had had stored in her ovary the germs of all the children of men.

The theory at first took the form of a belief that it was the females who were thus encased in the first being. One couple of each species was created, but the female contained in her ovary all the future individuals of the species, of either sex. However, this had to be altered when the Dutch microscopist, Leeuwenhoek, discovered the male spermatozoa in 1690, and showed that an immense number of these extremely fine and mobile thread-like beings exist in the male sperm (this will be explained in Chapter VII). This astonishing discovery was further advanced when it was proved that these living bodies, swimming about in the seminal fluid, were real animalcules, and, in fact, were the pre-formed germs of the future generation. When the male and female procreative elements came together at conception, these thread-like spermatozoa (“seed-animals”) were supposed to penetrate into the fertile body of the ovum and begin to develop there, as the plant seed does in the fruitful earth. Hence, every spermatozoon was regarded as a homunculus, a tiny complete man; all the parts were believed to be pre-formed in it, and merely grew larger when it reached its proper medium in the female ovum. This theory, also, was consistently developed in the sense that in each of these thread-like bodies the whole of its posterity was supposed to be present in the minutest form. Adam’s sexual glands were thought to have contained the germs of the whole of humanity.

This “theory of male scatulation” found itself at once in keen opposition to the prevailing “female” theory. The two rival theories at once opened a very lively campaign, and the physiologists of the eighteenth century were divided into two great camps—the Animalculists and the Ovulists—which fought vigorously. The animalculists held that the spermatozoa were the true germs, and appealed to the lively movements and the structure of these bodies. The opposing party of the Ovulists, who clung to the older “evolution theory,” affirmed that the ovum is the real germ, and that the spermatozoa merely stimulate it at conception to begin its growth; all the future generations are stored in the ovum. This view was held by the great majority of the biologists of the eighteenth century, in spite of the fact that Wolff proved it in 1759 to be without foundation. It owed its prestige chiefly to the circumstance that the most weighty authorities in the biology and philosophy of the day decided in favour of it, especially Haller, Bonnet, and Leibnitz.

Albrecht Haller, professor at Göttingen, who is often called the father of physiology, was a man of wide and varied learning, but he does not occupy a very high position in regard to insight into natural phenomena. He made a vigorous defence of the “evolutionary theory” in his famous work, Elementa physiologiae, affirming: “There is no such thing as formation (nulla est epigenesis). No part of the animal frame is made before another; all were made together.” He thus denied that there was any evolution in the proper sense of the word, and even went so far as to say that the beard existed in the new-born child and the antlers in the hornless fawn; all the parts were there in advance, and were merely hidden from the eye of man for the time being. Haller even calculated the number of human beings that God must have created on the sixth day and stored away in Eve’s ovary. He put the number at 200,000 millions, assuming the age of the world to be 6000 years, the average age of a human being to be thirty years, and the population of the world at

1. “Packing theory” would be the literal translation. Scatula is the Latin for a case or box.—Translator.

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that time to be 1000 millions. And the famous Haller maintained all this nonsense, in spite of its ridiculous consequences, even after Wolff had discovered the real course of embryonic development and established it by direct observation!

Among the philosophers of the time the distinguished Leibnitz was the chief defender of the “preformation theory,” and by his authority and literary prestige won many adherents to it. Supported by his system of monads, according to which body and soul are united in inseparable association and by their union form the individual, or the “monad,” Leibnitz consistently extended the “scatulation theory” to the soul, and held that this was no more evolved than the body. He says, for instance, in his Théodicée: “I mean that these souls, which one day are to be the souls of men, are present in the seed, like those of other species; in such wise that they existed in our ancestors as far back as Adam, or from the beginning of the world, in the forms of organised bodies.”

The theory seemed to receive considerable support from the observations of one of its most zealous supporters, Bonnet. In 1745 he discovered, in the plant-louse, a case of parthenogenesis, or virgin-birth, an interesting form of reproduction that has lately been found by Siebold and others among various classes of the articulata, especially crustacea and insects. Among these and other animals of certain lower species the female may reproduce for several generations without having been fertilised by the male. These ova that do not need fertilisation are called “false ova,” pseudova or spores. Bonnet saw that a female plant-louse, which he had kept in cloistral isolation, and rigidly removed from contact with males, had on the eleventh day (after forming a new skin for the fourth time) a living daughter, and during the next twenty days ninety-four other daughters; and that all of them went on to reproduce in the same way without any contact with males. It seemed as if this furnished an irrefutable proof of the truth of the scatulation theory, as it was held by the Ovulists; it is not surprising to find that the theory then secured general acceptance.

This was the condition of things when suddenly, in 1759, Caspar Friedrich Wolff appeared, and dealt a fatal blow at the whole preformation theory with his new theory of epigenesis. Wolff, the son of a Berlin tailor, was born in 1733, and went through his scientific and medical studies, first at Berlin under the famous anatomist Meckel, and afterwards at Halle. Here he secured his doctorate in his twenty-sixth year, and in his academic dissertation (November 28th, 1759), the Theoria generationis, expounded the new theory of a real development on a basis of epigenesis. This treatise is, in spite of its smallness and its obscure phraseology, one of the most valuable in the whole range of biological literature. It is equally distinguished for the mass of new and careful observations it contains, and the far-reaching and pregnant ideas which the author everywhere extracts from his observations and builds into a luminous and accurate theory of generation. Nevertheless, it met with no success at the time. Although scientific studies were then assiduously cultivated owing to the impulse given by Linné—although botanists and zoologists were no longer counted by dozens, but by hundreds, hardly any notice was taken of Wolff’s theory. Even when he established the truth of epigenesis by the most rigorous observations, and demolished the airy structure of the preformation theory, the “exact” scientist Haller proved one of the most strenuous supporters of the old theory, and rejected Wolff’s correct view with a dictatorial “There is no such thing as evolution.” He even went on to say that religion was menaced by the new theory! It is not surprising that the whole of the physiologists of the second half of the eighteenth century submitted to the ruling of this physiological pontiff, and attacked the theory of epigenesis as a dangerous innovation. It was not until more than fifty years afterwards that Wolff’s work was appreciated. Only when Meckel translated into German in 1812 another valuable work of Wolff’s on The Formation of the Alimentary Canal (written in 1768), and called attention to its great importance, did people begin to think of him once more; yet this obscure writer had evinced a profounder insight into the nature of the living organism than any other scientist of the eighteenth century.

Wolff’s idea led to an appreciable advance over the whole field of biology. There is such a vast number of new and important observations and pregnant thoughts in his writings that we have only gradually learned to appreciate them rightly in the course of the nineteenth

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century. He opened up the true path for research in many directions. In the first place, his theory of epigenesis gave us our first real insight into the nature of embryonic development. He showed convincingly that the development of every organism consists of a series of new formations, and that there is no trace whatever of the complete form either in the ovum or the spermatozoon. On the contrary, these are quite simple bodies, with a very different purport. The embryo which is developed from them is also quite different, in its internal arrangement and outer configuration, from the complete organism. There is no trace whatever of preformation or in-folding of organs. To-day we can scarcely call epigenesis a theory, because we are convinced it is a fact, and can demonstrate it at any moment with the aid of the microscope.

Wolff furnished the conclusive empirical proof of his theory in his classic dissertation on The Formation of the Alimentary Canal (1768). In its complete state the alimentary canal of the hen is a long and complex tube, with which the lungs, liver, salivary glands, and many other small glands, are connected. Wolff showed that in the early stages of the embryonic chick there is no trace whatever of this complicated tube with all its dependencies, but instead of it only a flat, leaf-shaped body; that, in fact, the whole embryo has at first the appearance of a flat, oval-shaped leaf. When we remember how difficult the exact observation of so fine and delicate a structure as the early leaf-shaped body of the chick must have been with the poor microscopes then in use, we must admire the rare faculty for observation which enabled Wolff to make the most important discoveries in this most difficult part of embryology. By this laborious research he reached the correct opinion that the embryonic body of all the higher animals, such as the birds, is for some time merely a flat, thin, leaf-shaped disk—consisting at first of one layer, but afterwards of several. The lowest of these layers is the alimentary canal, and Wolff followed its development from its commencement to its completion. He showed how this leaf-shaped structure first turns into a groove, then the margins of this groove fold together and form a closed canal, and at length the two external openings of the tube (the mouth and anus) appear.

Moreover, the important fact that the other systems of organs are developed in the same way, from tubes formed out of simple layers, did not escape Wolff. The nerveless system, muscular system, and vascular (blood-vessel) system, with all the organs appertaining thereto, are, like the alimentary system, developed out of simple leaf-shaped structures. Hence, Wolff came to the view by 1768 which Pander developed in the Theory of Germinal Layers fifty years afterwards. His principles are not literally correct; but he comes as near to the truth in them as was possible at that time, and could be expected of him.

Our admiration of this gifted genius increases when we find that he was also the precursor of Goethe in regard to the metamorphosis of plants and of the famous cellular theory. Wolff had, as Huxley showed, a clear presentiment of this cardinal theory, since he recognised small microscopic globules as the elementary parts out of which the germinal layers arose.

Finally, I must invite special attention to the mechanical character of the profound philosophic reflections which Wolff always added to his remarkable observations. He was a great monistic philosopher, in the best meaning of the word. It is unfortunate that his philosophic discoveries were ignored as completely as his observations for more than half a century. We must be all the more careful to emphasise the fact of their clear monistic tendency.

Title and Contents
Glossary
Chapter I
Chapter III
Figs. 1–209
Figs. 210–408