THE EVOLUTION OF MAN
Volume I

CHAPTER III
MODERN EMBRYOLOGY

We may distinguish three chief periods in the growth of our science of human embryology. The first has been considered in the preceding chapter; it embraces the whole of the preparatory period of research, and extends from Aristotle to Caspar Friedrich Wolff, or to the year 1759, in which the epoch-making Theoria generationis was published. The second period, with which we have now to deal, lasts about a century—that is to say, until the appearance of Darwin’s Origin of Species, which brought about a change in the very foundations of biology, and, in particular, of embryology. The third period begins with Darwin. When we say that the second period lasted a full century, we must remember that Wolff’s work had remained almost unnoticed during half the time—namely, until the year 1812. During the whole of these fifty-three years not a single book that appeared followed up the path that Wolff had opened, or extended his theory of embryonic development. We merely find his views—perfectly correct views, based on extensive observations of fact—mentioned here and there as erroneous; their opponents, who adhered to the dominant theory of preformation, did not even deign to reply to them. This unjust treatment was chiefly due to the extraordinary authority of Albrecht von Haller; it is one of the most astonishing instances of a great authority, as such, preventing for a long time the recognition of established facts.

The general ignorance of Wolff’s work was so great that at the beginning of the nineteenth century two scientists of Jena, Oken (1806) and Kieser (1810), began independent research into the development of the alimentary canal of the chick, and hit upon the right clue to the embryonic puzzle, without knowing a word about Wolff’s important treatise on the same subject. They were treading in his very footsteps without suspecting it. This can be easily proved from the fact that they did not travel as far as Wolff. It was not until Meckel translated into German Wolff’s book on the alimentary system, and pointed out its great importance, that the eyes of anatomists and physiologists were suddenly opened. At once a number of biologists instituted fresh embryological inquiries, and began to confirm Wolff’s theory of epigenesis.

This resuscitation of embryology and development of the epigenesis-theory was chiefly connected with the university of Würtzburg. One of the professors there at that time was Döllinger, an eminent biologist, and father of the famous Catholic historian who later distinguished himself by his opposition to the new dogma of papal infallibility. Döllinger was both a profound thinker and an accurate observer. He took the keenest interest in embryology, and worked at it a good deal. However, he is not himself responsible for any important result in this field. In 1816 a young medical doctor, whom we may at once designate as Wolff’s chief successor, Karl Ernst von Baer, came to Würtzburg. Baer’s conversations with Döllinger on embryology led to a fresh series of most extensive investigations. Döllinger had expressed a wish that some young scientist should begin again under his guidance an independent inquiry into the development of the chick during the hatching of the egg. As neither he nor Baer had money enough to pay for an incubator and the proper control of the experiments, and for a competent artist to illustrate the various stages observed, the lead of the enterprise was given to Christian Pander, a wealthy friend of Baer’s who had been induced by Baer to come to Würtzburg. An able engraver, Dalton, was engaged to do the copper-plates. In a short time the embryology of the chick, in which Baer was taking the greatest indirect interest, was so far advanced that Pander was able to sketch the main features of it on the ground of Wolff’s theory in the dissertation he published in 1817. He clearly enunciated the theory of germinal layers which Wolff

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had anticipated, and established the truth of Wolff’s idea of a development of the complicated systems of organs out of simple leaf-shaped primitive structures. According to Pander, the leaf-shaped object in the hen’s egg divides, before the incubation has proceeded twelve hours, into two different layers, an external serous layer and an internal mucous layer; between the two there develops later a third layer, the vascular (blood-vessel) layer.1

Karl Ernst von Baer, who had set afoot Pander’s investigation, and had shown the liveliest interest in it after Pander’s departure from Würtzburg, began his own much more comprehensive research in 1819. He published the mature result nine years afterwards in his famous work, Animal Embryology: Observation and Reflection (not translated). This classic work still remains a model of careful observation united to profound philosophic speculation. The first part appeared in 1828, the second in 1837. The book proved to be the foundation on which the whole science of embryology has built down to our own day. It so far surpassed its predecessors, and Pander in particular, that it has become, after Wolff’s work, the chief base of modern embryology.

Baer was one of the greatest scientists of the nineteenth century, and exercised considerable influence on other branches of biology as well. He built up the theory of germinal layers, as a whole and in detail, so clearly and solidly that it has been the starting-point of embryological research ever since. He taught that in all the vertebrates first two and then four of these germinal layers are formed; and that the earliest rudimentary organs of the body arise by the conversion of these layers into tubes. He described the first appearance of the vertebrate embryo, as it may be seen in the globular yelk of the fertilised egg, as an oval disk which first divides into two layers. From the upper or animal layer are developed all the organs which accomplish the phenomena of animal life—the functions of sensation and motion, and the covering of the body. From the lower or vegetative layer come the organs which effect the vegetative life of the organism—nutrition, digestion, blood-formation, respiration, secretion, reproduction, etc.

Each of these original layers divides, according to Baer, into two thinner and superimposed layers or plates. He calls the two plates of the animal layer, the skin-stratum and muscle-stratum. From the upper of these plates, the skin-stratum, the external skin, or outer covering of the body, the central nervous system, and the sense-organs, are formed. From the lower, or muscle-stratum, the muscles, or fleshy parts and the bony skeleton—in a word, the motor organs—are evolved. In the same way, Baer said, the lower or vegetative layer splits into two plates, which he calls the vascular-stratum and the mucous-stratum. From the outer of the two (the vascular) the heart, blood-vessels, spleen, and the other vascular glands, the kidneys, and sexual glands, are formed. From the fourth or mucous layer, in fine, we get the internal and digestive lining of the alimentary canal and all its dependencies, the liver, lungs, salivary glands, etc. Baer had, in the main, correctly judged the significance of these four secondary embryonic layers, and he followed the conversion of them into the tube-shaped primitive organs with great perspicacity. He first solved the difficult problem of the transformation of this four-fold, flat, leaf-shaped, embryonic disk into the complete vertebrate body, through the conversion of the layers or plates into tubes. The flat leaves bend themselves in obedience to certain laws of growth; the borders of the curling plates approach nearer and nearer; until at last they come into actual contact. Thus out of the flat gut-plate is formed a hollow gut-tube, out of the flat spinal plate a hollow nerve-tube, from the skin-plate a skin-tube, and so on.

Among the many great services which Baer rendered to embryology, especially vertebrate embryology, we must not forget his discovery of the human ovum. Earlier scientists had, as a rule, of course, assumed that man developed out of an egg, like the other animals. In fact, the preformation theory held that the germs of the whole of humanity were stored already in Eve’s ova. But the real ovum escaped detection until the year 1827. This ovum is extremely small, being a tiny round vesicle about the 1/120 of an inch in diameter; it can be seen under very favourable circumstances with the naked eye as a tiny particle, but is otherwise quite invisible. This particle is formed in the ovary inside a much larger

1. The technical terms which are bound to creep into this chapter will be fully understood later on.—Translator.

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globule, which takes the name of the Graafian follicle, from its discoverer, Graaf, and had previously been regarded as the true ovum. However, in 1827 Baer proved that it was not the real ovum, which is much smaller, and is contained within the follicle. (Compare the end of Chapter XXIX.)

Baer was also the first to observe what is known as the segmentation sphere of the vertebrate; that is to say, the round vesicle which first develops out of the impregnated ovum, and the thin wall of which is made up of a single layer of regular, polygonal (many-cornered) cells (see the illustration in Chapter XII). Another discovery of his that was of great importance in constructing the vertebrate stem and the characteristic organisation of this extensive group (to which man belongs) was the detection of the axial rod, or the chorda dorsalis. There is a long, round, cylindrical rod of cartilage which runs down the longer axis of the vertebrate embryo; it appears at an early stage, and is the first sketch of the spinal column, the solid skeletal axis of the vertebrate. In the lowest of the vertebrates, the amphioxus, the internal skeleton consists only of this cord throughout life. But even in the case of man and all the higher vertebrates it is round this cord that the spinal column and the brain are afterwards formed.

However, important as these and many other discoveries of Baer’s were in vertebrate embryology, his researches were even more influential, from the circumstance that he was the first to employ the comparative method in studying the development of the animal frame. Baer occupied himself chiefly with the embryology of vertebrates (especially the birds and fishes). But he by no means confined his attention to these, gradually taking the various groups of the invertebrates into his sphere of study. As the general result of his comparative embryological research, Baer distinguished four different modes of development and four corresponding groups in the animal world. These chief groups or types are: 1, the vertebrata; 2, the articulata; 3, the mollusca; and 4, all the lower groups which were then wrongly comprehended under the general name of the radiata. Georges Cuvier had been the first to formulate this distinction, in 1812. He showed that these groups present specific differences in their whole internal structure, and the connection and disposal of their systems of organs; and that, on the other hand, all the animals of the same type—say, the vertebrates—essentially agreed in their inner structure, in spite of the greatest superficial differences. But Baer proved that these four groups are also quite differently developed from the ovum; and that the series of embryonic forms is the same throughout for animals of the same type, but different in the case of other animals. Up to that time the chief aim in the classification of the animal kingdom was to arrange all the animals from lowest to highest, from the infusorium to man, in one long and continuous series. The erroneous idea prevailed nearly everywhere that there was one uninterrupted chain of evolution from the lowest animal to the highest. Cuvier and Baer proved that this view was false, and that we must distinguish four totally different types of animals, on the ground of anatomic structure and embryonic development.

Baer’s epoch-making works aroused an extraordinary and widespread interest in embryological research. Immediately afterwards we find a great number of observers at work in the newly opened field, enlarging it in a very short time with great energy by their various discoveries in detail. Next to Baer’s comes the admirable work of Heinrich Rathke, of Königsberg (died 1860); he made an extensive study of the embryology, not only of the invertebrates (crustaceans, insects, molluscs), but also, and particularly, of the vertebrates (fishes, tortoises, serpents, crocodiles, etc.). We owe the first comprehensive studies of mammal embryology to the careful research of Wilhelm Bischoff, of Munich; his embryology of the rabbit (1840), the dog (1842), the guinea-pig (1852), and the doe (1854), still form classical studies. About the same time a great impetus was given to the embryology of the invertebrates. The way was opened through this obscure province by the studies of the famous Berlin zoologist, Johannes Müller, on the echinoderms. He was followed by Albert Kölliker, of Würtzburg, writing on the cuttlefish (or the cephalopods), Siebold and Huxley on worms and zoophytes, Fritz Muller (Desterro) on the crustacea, Weismann on insects, and so on. The number of workers in this field has greatly increased of late, and a quantity of new and astonishing discoveries have been made. One notices, in several of these recent works on

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embryology, that their authors are too little acquainted with comparative anatomy and classification. Paleontology is, unfortunately, altogether neglected by many of these new workers, although this interesting science furnishes most important facts for phylogeny, and thus often proves of very great service in ontogeny.

A very important advance was made in our science in 1839, when the cellular theory was established, and a new field of inquiry bearing on embryology was suddenly opened. When the famous botanist, M. Schleiden, of Jena, showed in 1838, with the aid of the microscope, that every plant was made up of innumerable elementary parts, which we call cells, a pupil of Johannes Müller at Berlin, Theodor Schwann, applied the discovery at once to the animal organism. He showed that in the animal body as well, when we examine its tissues in the microscope, we find these cells everywhere to be the elementary units. All the different tissues of the organism, especially the very dissimilar tissues of the nerves, muscles, bones, external skin, mucous lining, etc., are originally formed out of cells; and this is also true of all the tissues of the plant. These cells are separate living beings; they are the citizens of the State which the entire multicellular organism seems to be. This important discovery was bound to be of service to embryology, as it raised a number of new questions. What is the relation of the cells to the germinal layers? Are the germinal layers composed of cells, and what is their relation to the cells of the tissues that form later? How does the ovum stand in the cellular theory? Is the ovum itself a cell, or is it composed of cells? These important questions were now imposed on the embryologist by the cellular theory.

The most notable effort to answer these questions—which were attacked on all sides by different students—is contained in the famous work, Inquiries into the Development of the Vertebrates (not translated) of Robert Remak, of Berlin (1851). This gifted scientist succeeded in mastering, by a complete reform of the science, the great difficulties which the cellular theory had at first put in the way of embryology. A Berlin anatomist, Carl Boguslaus Reichert, had already attempted to explain the origin of the tissues. But this attempt was bound to miscarry, since its not very clear-headed author lacked a sound acquaintance with embryology and the cell theory, and even with the structure and development of the tissue in particular. Remak at length brought order into the dreadful confusion that Reichert had caused; he gave a perfectly simple explanation of the origin of the tissues. In his opinion the animal ovum is always a simple cell: the germinal layers which develop out of it are always composed of cells; and these cells that constitute the germinal layers arise simply from the continuous and repeated cleaving (segmentation) of the original solitary cell. It first divides into two and then into four cells; out of these four cells are born eight, then sixteen, thirty-two, and so on. Thus, in the embryonic development of every animal and plant there is formed first of all out of the simple egg cell, by a repeated subdivision, a cluster of cells, as Kölliker had already stated in connection with the cephalopods in 1844. The cells of this group spread themselves out flat and form leaves or plates; each of these leaves is formed exclusively out of cells. The cells of different layers assume different shapes, increase, and differentiate; and in the end there is a further cleavage (differentiation) and division of work of the cells within the layers, and from these all the different tissues of the body proceed.

These are the simple foundations of histogeny, or the science that treats of the development of the tissues (hista), as it was established by Remak and Kölliker. Remak, in determining more closely the part which the different germinal layers play in the formation of the various tissues and organs, and in applying the theory of evolution to the cells and the tissues they compose, raised the theory of germinal layers, at least as far as it regards the vertebrates, to a high degree of perfection.

Remak showed that three layers are formed out of the two germinal layers which compose the first simple leaf-shaped structure of the vertebrate body (or the “germinal disk”), as the lower layer splits into two plates. These three layers have a very definite relation to the various tissues. First of all, the cells which form the outer skin of the body (the epidermis), with its various dependencies (hairs, nails, etc.)—that is to say, the entire outer envelope of the body—are developed out of the outer or upper layer; but there are also developed in a curious way out of the same layer the cells which form the central nervous system, the

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brain and the spinal cord. In the second place, the inner or lower germinal layer gives rise only to the cells which form the epithelium (the whole inner lining) of the alimentary canal and all that depends on it (the lungs, liver, pancreas, etc.), or the tissues that receive and prepare the nourishment of the body. Finally, the middle layer gives rise to all the other tissues of the body, the muscles, blood, bones, cartilage, etc. Remak further proved that this middle layer, which he calls “the motor-germinative layer,” proceeds to subdivide into two secondary layers. Thus we find once more the four layers which Baer had indicated. Remak calls the outer secondary leaf of the middle layer (Baer’s “muscular layer”) the “skin layer” (it would be better to say, skin-fibre layer); it forms the outer wall of the body (the true skin, the muscles, etc.). To the inner secondary leaf (Baer’s “vascular layer”) he gave the name of the “alimentary-fibre layer”; this forms the outer envelope of the alimentary canal, with the mesentery, the heart, the blood-vessels, etc.

On this firm foundation provided by Remak for histogeny, or the science of the formation of the tissues, our knowledge has been gradually built up and enlarged in detail. There have been several attempts to restrict and even destroy Remak’s principles. The two anatomists, Reichert (of Berlin) and Wilhelm His (of Leipzic), especially, have endeavoured in their works to introduce a new conception of the embryonic development of the vertebrate, according to which the two primary germinal layers would not be the sole sources of formation. But these efforts were so seriously marred by ignorance of comparative anatomy, an imperfect acquaintance with ontogenesis, and a complete neglect of phylogenesis, that they could not have more than a passing success. We can only explain how these curious attacks of Reichert and His came to be regarded for a time as advances by the general lack of discrimination and of grasp of the true object of embryology.

Wilhelm His published, in 1868, his extensive Researches into the Earliest Form of the Vertebrate Body,1 one of the curiosities of embryological literature. The author imagines that he can build a “mechanical theory of embryonic development” by merely giving an exact description of the embryology of the chick, without any regard to comparative anatomy and phylogeny, and thus falls into an error that is almost without parallel in the history of biological literature. As the final result of his laborious investigations, His tells us “that a comparatively simple law of growth is the one essential thing in the first development. Every formation, whether it consist in cleavage of layers, or folding, or complete division, is a consequence of this fundamental law.” Unfortunately, he does not explain what this “law of growth” is; just as other opponents of the theory of selection, who would put in its place a great “law of evolution,” omit to tell us anything about the nature of this. Nevertheless, it is quite clear from His’s works that he imagines constructive Nature to be a sort of skilful tailor. The ingenious operator succeeds in bringing into existence, by “evolution,” all the various forms of living things by cutting up in different ways the germinal layers, bending and folding, tugging and splitting, and so on.

His’s embryological theories excited a good deal of interest at the time of publication, and have evoked a fair amount of literature in the last few decades. He professed to explain the most complicated parts of organic construction (such as the development of the brain) in the simplest way on mechanical principles, and to derive them immediately from simple physical processes (such as unequal distribution of strain in an elastic plate). It is quite true that a mechanical or monistic explanation (or a reduction of natural processes) is the ideal of modern science, and this ideal would be realised if we could succeed in expressing these formative processes in mathematical formulæ. His has, therefore, inserted plenty of numbers and measurements in his embryological works, and given them an air of “exact” scholarship by putting in a quantity of mathematical tables. Unfortunately, they are of no value, and do not help us in the least in forming an “exact” acquaintance with the embryonic phenomena. Indeed, they wander from the true path altogether by neglecting the phylogenetic method; this, he thinks, is “a mere by-path,” and is “not necessary at all for the explanation of the facts of

1. None of His’s works have been translated into English.

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embryology,” which are the direct consequence of physiological principles. What His takes to be a simple physical process—for instance, the folding of the germinal layers (in the formation of the medullary tube, alimentary tube, etc.)—is, as a matter of fact, the direct result of the growth of the various cells which form those organic structures; but these growth-motions have themselves been transmitted by heredity from parents and ancestors, and are only the hereditary repetition of countless phylogenetic changes which have taken place for thousands of years in the race-history of the said ancestors. Each of these historical changes was, of course, originally due to adaptation; it was, in other words, physiological, and reducible to mechanical causes. But we have, naturally, no means of observing them now. It is only by the hypotheses of the science of evolution that we can form an approximate idea of the organic links in this historic chain.

All the best recent research in animal embryology has led to the confirmation and development of Baer and Remak’s theory of the germinal layers. One of the most important advances in this direction of late was the discovery that the two primary layers out of which is built the body of all vertebrates (including man) are also present in all the invertebrates, with the sole exception of the lowest group, the unicellular protozoa. Huxley had detected them in the medusa in 1849. He showed that the two layers of cells from which the body of this zoophyte is developed correspond, both morphologically and physiologically, to the two original germinal layers of the vertebrate. The outer layer, from which come the external skin and the muscles, was then called by Allman (1853) the “ectoderm” (outer layer, or skin); the inner layer, which forms the alimentary and reproductory organs, was called the “entoderm” (= inner layer). In 1867 and the following years the discovery of the germinal layers was extended to other groups of the invertebrates. In particular, the indefatigable Russian zoologist, Kowalevsky, found them in all the most diverse sections of the invertebrates—the worms, tunicates, echinoderms, molluscs, articulates, etc.

In my monograph on the sponges (1872) I proved that these two primary germinal layers are also found in that group, and that they may be traced from it right up to man, through all the various classes, in identical form. This “homology of the two primary germinal layers” extends through the whole of the metazoa, or tissue-forming animals; that is to say, through the whole animal kingdom, with the one exception of its lowest section, the unicellular beings, or protozoa. These lowly organised animals do not form germinal layers, and therefore do not succeed in forming true tissue. Their whole body consists of a single cell (as is the case with the amœbæ and infusoria), or of a loose aggregation of only slightly differentiated cells, though it may not even reach the full structure of a single cell (as with the monera). But in all other animals the ovum first grows into two primary layers, the outer or animal layer (the ectoderm, epiblast, or ectoblast), and the inner or vegetal layer (the entoderm, hypoblast, or endoblast); and from these the tissues and organs are formed. The first and oldest organ of all these metazoa is the primitive gut (or progaster) and its opening, the primitive mouth (prostoma). The typical embryonic form of the metazoa, as it is presented for a time by this simple structure of the two-layered body, is called the gastrula; it is to be conceived as the hereditary reproduction of some primitive common ancestor of the metazoa, which we call the gastræa. This applies to the sponges and other zoophyta, and to the worms, the mollusca, echinoderma, articulata, and vertebrata. All these animals may be comprised under the general heading of “gut animals,” or metazoa, in contradistinction to the gutless protozoa.

I have pointed out in my Study of the Gastræa Theory [not translated] (1873) the important consequences of this conception in the morphology and classification of the animal world. I also divided the realm of metazoa into two great groups, the lower and higher metazoa. In the first are comprised the cœlenterata (also called zoophytes, or plant-animals). In the lower forms of this group the body consists throughout life merely of the primary germinal layers, with the cells sometimes more and sometimes less differentiated. But with the higher forms of the cœlentarata (the corals, higher medusæ, ctenophoræ, and platodes) a middle layer, or mesoderm, often of considerable size, is developed between the

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other two layers; but blood and an internal cavity are still lacking.

To the second great group of the metazoa I gave the name of the cœlomaria, or bilaterata (or the bilateral higher forms). They all have a cavity within the body (cœloma), and most of them have blood and blood-vessels. In this are comprised the six higher stems of the animal kingdom, the annulata and their descendants, the mollusca, echinoderma, articulata, tunicata, and vertebrata. In all these bilateral organisms the two-sided body is formed out of four secondary germinal layers, of which the inner two construct the wall of the alimentary canal, and the outer two the wall of the body. Between the two pairs of layers lies the cavity (cœloma).

Although I laid special stress on the great morphological importance of this cavity in my Study of the Gastræa Theory, and endeavoured to prove the significance of the four secondary germinal layers in the organisation of the cœlomaria, I was unable to deal satisfactorily with the difficult question of the mode of their origin. This was done eight years afterwards by the brothers Oscar and Richard Hertwig in their careful and extensive comparative studies. In their masterly Cœlum Theory: An Attempt to Explain the Middle Germinal Layer [not translated] (1881) they showed that in most of the metazoa, especially in all the vertebrates, the body-cavity arises in the same way, by the outgrowth of two sacs from the inner layer. These two cœlom-pouches proceed from the rudimentary mouth of the gastrula, between the two primary layers. The inner plate of the two-layered cœlom-pouch (the visceral layer) joins itself to the entoderm; the outer plate (parietal layer) unites with the ectoderm. Thus are formed the double-layered gut-wall within and the double-layered body-wall without; and between the two is formed the cavity of the cœlom, by the blending of the right and left cœlom-sacs. We shall see this more fully in Chapter X.

The many new points of view and fresh ideas suggested by my gastræa theory and Hertwig’s cœlom theory led to the publication of a number of writings on the theory of germinal layers. Most of them set out to oppose it at first, but in the end the majority supported it. Of late years both theories are accepted in their essential features by nearly every competent man of science, and light and order have been introduced into this once dark and contradictory field of research. A further cause of congratulation for this solution of the great embryological controversy is that it brought with it a recognition of the need for phylogenetic study and explanation.

Interest and practice in embryological research have been remarkably stimulated during the past thirty years by this appreciation of phylogenetic methods. Hundreds of assiduous and able observers are now engaged in the development of comparative embryology and its establishment on a basis of evolution, whereas they numbered only a few dozen not many decades ago. It would take too long to enumerate even the most important of the countless valuable works which have enriched embryological literature since that time. References to them will be found in the latest manuals of embryology of Kölliker, Balfour, Hertwig, Kollman, Korschelt, and Heider.

Kölliker’s Entwickelungsgeschichte des Menschen und der höherer Thiere, the first edition of which appeared forty-two years ago, had the rare merit at that time of gathering into presentable form the scattered attainments of the science, and expounding them in some sort of unity on the basis of the cellular theory and the theory of germinal layers. Unfortunately, the distinguished Würtzburg anatomist, to whom comparative anatomy, histology, and ontogeny owe so much, is opposed to the theory of descent generally and to Darwinism in particular. All the other manuals I have mentioned take a decided stand on evolution. Francis Balfour has carefully collected and presented with discrimination, in his Manual of Comparative Embryology (1880), the very scattered and extensive literature of the subject; he has also widened the basis of the gastræa theory by a comparative description of the rise of the organs from the germinal layers in all the chief groups of the animal kingdom, and has given a most thorough empirical support to the principles I have formulated. A comparison of his work with the excellent Text-book of the Embryology of the Vertebrates (1890) [translation 1895] of Korschelt and Heider shows what astonishing progress has been made in the science in the course of ten years. I would especially recommend the manuals of Julius Kollmann and Oscar Hertwig to those readers who are stimulated to further study by these chapters on human

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embryology. Kollmann’s work is commendable for its clear treatment of the subject and very fine original illustrations; its author adheres firmly to the biogenetic law, and uses it throughout with considerable profit. That is not the case in Oscar Hertwig’s recent Text-book of the Embryology of Man and the Mammals [translations 1892 and 1899] (seventh edition 1902). This able anatomist has of late often been quoted as an opponent of the biogenetic law, although he himself had demonstrated its great value thirty years ago. His recent vacillation is partly due to the timidity which our “exact” scientists have with regard to hypotheses; though it is impossible to make any headway in the explanation of facts without them. However, the purely descriptive part of embryology in Hertwig’s Text-book is very thorough and reliable.

A new branch of embryological research has been studied very assiduously in the last decade of the nineteenth century—namely, “experimental embryology.” The great importance which has been attached to the application of physical experiments to the living organism for the last hundred years, and the valuable results that it has given to physiology in the study of the vital phenomena, have led to its extension to embryology. I was the first to make experiments of this kind during a stay of four months on the Canary Island, Lanzerote, in 1866. I there made a thorough investigation of the almost unknown embryology of the siphonophoræ. I cut a number of the embryos of these animals (which develop freely in the water, and pass through a very curious transformation), at an early stage, into several pieces, and found that a fresh organism (more or less complete, according to the size of the piece) was developed from each particle. More recently some of my pupils have made similar experiments with the embryos of vertebrates (especially the frog) and some of the invertebrates. Wilhelm Roux, in particular, has made extensive experiments, and based on them a special “mechanical embryology,” which has given rise to a good deal of discussion and controversy. Roux has published a special journal for these subjects since 1895, the Archiv für Entwickelungsmechanik. The contributions to it are very varied in value. Many of them are valuable papers on the physiology and pathology of the embryo. Pathological experiments—the placing of the embryo in abnormal conditions—have yielded many interesting results; just as the physiology of the normal body has for a long time derived assistance from the pathology of the diseased organism. Other of these mechanical-embryological articles return to the erroneous methods of His, and are only misleading. This must be said of the many contributions of mechanical embryology which take up a position of hostility to the theory of descent and its chief embryological foundation—the biogenetic law. This law, however, when rightly understood, is not opposed to, but is the best and most solid support of, a sound mechanical embryology. Impartial reflection and a due attention to paleontology and comparative anatomy should convince these one-sided mechanicists that the facts they have discovered—and, indeed, the whole embryological process—cannot be fully understood without the theory of descent and the biogenetic law.

Title and Contents
Glossary
Chapter II
Chapter IV
Figs. 1–209
Figs. 210–408