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

else than the structure of the body-cavity. The two layers that enclose it--the upper parietal layer (hpl) and the lower visceral layer (df)--are pressed together from without, but clearly distinguishable. This is even clearer a little later, when the medullary furrow is closed into the nerve-tube (Figure 1.93 mr).

Special importance attaches to the fact that here again the four secondary germinal layers are already sharply distinct, and easily separated from each other. There is only one very restricted area in which they are connected, and actually pass into each other; this is the region of the primitive mouth, which is contracted in the amniotes into a dorsal longitudinal cleft, the primitive groove. Its two lateral lip-borders form the primitive streak, which has long been recognised as the most important embryonic source and starting-point of further processes. Sections through this primitive streak (Figures 1.94 and 1.95) show that the two primary germinal layers grow at an early stage (in the discoid gastrula of the chick, a few hours after incubation) into the primitive streak (x), and that the two middle layers extend outward from this thickened axial plate (y) to the right and left between the former. The plates of the coelom-layers, the parietal skin-fibre-layer (m) and the visceral gut-fibre-layer (f), are seen to be still pressed close together, and only diverge later to form the body-cavity. Between the inner borders of the two flat coelom-pouches lies the chorda (Figure 1.95 x), which here again develops from the middle line of the dorsal wall of the primitive gut.

(FIGURES 1.94 AND 1.95. Transverse section of the primitive-streak (primitive mouth) of the chick. Figure 1.94 a few hours after the commencement of incubation, Figure 1.95 a little later. (From Waldeyer.) h horn-plate, n nerve-plate, m skin-fibre-layer, f gut-fibre-layer, d gut-gland-layer, y primitive streak or axial plate, in which all four germinal layers meet, x structure of the chorda, u region of the later primitive kidneys.)

Coelomation takes place in the vertebrates in just the same way as in the birds and reptiles. This was to be expected, as the characteristic gastrulation of the mammal has descended from that of the reptiles. In both cases a discoid gastrula with primitive streak arises from the segmented ovum, a two-layered germinal disk with long and small hinder primitive mouth. Here again the two primary germinal layers are only directly connected (Figure 1.96 pr) along the primitive streak (at the folding-point of the blastula), and from this spot (the border of the primitive mouth) the middle germinal layers (mk) grow out to right and left between the preceding. In the fine illustration of the coelomula of the rabbit which Van Beneden has given us (Figure 1.96) one can clearly see that each of the four secondary germinal layers consists of a single stratum of cells.

Finally, we must point out, as a fact of the utmost importance for our anthropogeny and of great general interest, that the four-layered coelomula of man has just the same construction as that of the rabbit (Figure 1.96). A vertical section that Count Spee made through the primitive mouth or streak of a very young human germinal disk (Figure 1.97) clearly shows that here again the four secondary germ-layers are inseparably connected only at the primitive streak, and that here also the two flattened coelom-pouches (mk) extend outwards to right and left from the primitive mouth between the outer and inner germinal layers. In this case, too, the middle germinal layer consists from the first of two separate strata of cells, the parietal (mp) and visceral (mv) mesoblasts.

(FIGURE 1.96. Transverse section of the primitive groove (or primitive mouth) of a rabbit. (From Van Beneden.) pr primitive mouth, ul lips of same (primitive lips), ak and ik outer and inner germinal layers, mk middle germinal layer, mp parietal layer, mv visceral layer of the mesoderm.

FIGURE 1.97. Transverse section of the primitive mouth (or groove) of a human embryo (at the coelomula stage). (From Count Spee.) pr primitive mouth, ul lips of same (primitive folds), ak and ik outer and inner germinal layers, mk middle layer, mp parietal layer, mv visceral layer of the mesoblasts.)

These concordant results of the best recent investigations (which have been confirmed by the observations of a number of scientists I have not enumerated) prove the unity of the vertebrate-stem in point of coelomation, no less than of gastrulation. In both respects the invaluable amphioxus--the sole survivor of the acrania--is found to be the original model that has preserved for us in palingenetic form by a tenacious heredity these most important embryonic processes. From this primary model of construction we can cenogenetically deduce all the embryonic forms of the other vertebrates, the craniota, by secondary modifications. My thesis of the universal formation of the gastrula by folding of the blastula has now been clearly proved for all the vertebrates; so also has been Hertwig's thesis of the origin of the middle germinal layers by the folding of a couple of coelom-pouches which appear at the border of the primitive mouth. Just as the gastraea-theory explains the origin and identity of the two primary layers, so the coelom-theory explains those of the four secondary layers. The point of origin is always the properistoma, the border of the original primitive mouth of the gastrula, at which the two primary layers pass directly into each other.

Moreover, the coelomula is important as the immediate source of the chordula, the embryonic reproduction of the ancient, typical, unarticulated, worm-like form, which has an axial chorda between the dorsal nerve-tube and the ventral gut-tube. This instructive chordula (Figures 1.83 to 1.86) provides a valuable support of our phylogeny; it indicates the important moment in our stem-history at which the stem of the chordonia (tunicates and vertebrates) parted for ever from the divergent stems of the other metazoa (articulates, echinoderms, and molluscs).

I may express here my opinion, in the form of a chordaea-theory, that the characteristic chordula-larva of the chordonia has in reality this great significance--it is the typical reproduction (preserved by heredity) of the ancient common stem-form of all the vertebrates and tunicates, the long-extinct Chordaea. We will return in

Chapter 2.

20 to these worm-like ancestors, which stand out as luminous points in the obscure stem-history of the invertebrate ancestors of our race.

 

CHAPTER XI(11. THE VERTEBRATE CHARACTER OF MAN.)

 

We have now secured a number of firm standing-places in the labyrinthian course of our individual development by our study of the important embryonic forms which we have called the cytula, morula, blastula, gastrula, coelomula, and chordula. But we have still in front of us the difficult task of deriving the complicated frame of the human body, with all its different parts, organs, members, etc., from the simple form of the chordula. We have previously considered the origin of this four-layered embryonic form from the two-layered gastrula. The two primary germinal layers, which form the entire body of the gastrula, and the two middle layers of the coelomula that develop between them, are the four simple cell-strata, or epithelia, which alone go to the formation of the complex body of man and the higher animals. It is so difficult to understand this construction that we will first seek a companion who may help us out of many difficulties.

This helpful associate is the science of comparative anatomy. Its task is, by comparing the fully-developed bodily forms in the various groups of animals, to learn the general laws of organisation according to which the body is constructed; at the same time, it has to determine the affinities of the various groups by critical appreciation of the degrees of difference between them. Formerly, this work was conceived in a teleological sense, and it was sought to find traces of the plan of the Creator in the actual purposive organisation of animals. But comparative anatomy has gone much deeper since the establishment of the theory of descent; its philosophic aim now is to explain the variety of organic forms by adaptation, and their similarity by heredity. At the same time, it has to recognise in the shades of difference in form the degree of blood-relationship, and make an effort to construct the ancestral tree of the animal world. In this way, comparative anatomy enters into the closest relations with comparative embryology on the one hand, and with the science of classification on the other.

Now, when we ask what position man occupies among the other organisms according to the latest teaching of comparative anatomy and classification, and how man's place in the zoological system is determined by comparison of the mature bodily forms, we get a very definite and significant reply; and this reply gives us extremely important conclusions that enable us to understand the embryonic development and its evolutionary purport. Since Cuvier and Baer, since the immense progress that was effected in the early decades of the nineteenth century by these two great zoologists, the opinion has generally prevailed that the whole animal kingdom may be distributed in a small number of great divisions or types. They are called types because a certain typical or characteristic structure is constantly preserved within each of these large sections. Since we applied the theory of descent to this doctrine of types, we have learned that this common type is an outcome of heredity; all the animals of one type are blood-relatives, or members of one stem, and can be traced to a common ancestral form. Cuvier and Baer set up four of these types: the vertebrates, articulates, molluscs, and radiates. The first three of these are still retained, and may be conceived as natural phylogenetic unities, as stems or phyla in the sense of the theory of descent. It is quite otherwise with the fourth type--the radiata. These animals, little known as yet at the beginning of the nineteenth century, were made to form a sort of lumber-room, into which were cast all the lower animals that did not belong to the other three types. As we obtained a closer acquaintance with them in the course of the last sixty years, it was found that we must distinguish among them from four to eight different types. In this way the total number of animal stems or phyla has been raised to eight or twelve (cf.

Chapter 2.

20).

These twelve stems of the animal kingdom are, however, by no means co-ordinate and independent types, but have definite relations, partly of subordination, to each other, and a very different phylogenetic meaning. Hence they must not be arranged simply in a row one after the other, as was generally done until thirty years ago, and is still done in some manuals. We must distribute them in three subordinate principal groups of very different value, and arrange the various stems phylogenetically on the principles which I laid down in my Monograph on the Sponges, and developed in the Study of the Gastraea Theory. We have first to distinguish the unicellular animals (protozoa) from the multicellular tissue-forming (metazoa). Only the latter exhibit the important processes of segmentation and gastrulation; and they alone have a primitive gut, and form germinal layers and tissues.

The metazoa, the tissue-animals or gut-animals, then sub-divide into two main sections, according as a body-cavity is or is not developed between the primary germinal layers. We may call these