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

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 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 2.

29.)

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 1.

12). 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 Konigsberg (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 Muller, on the echinoderms. He was followed by Albert Kolliker, of Wurtzburg, 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 embryology, that their authors are too little acquainted with comparative anatomy and classification. Palaeontology 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 Muller 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 Kolliker 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 Kolliker. 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 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