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

and also the hard chalky shell, are only formed within the oviduct round the impregnated ovum.

When the fertilisation of the bird's ovum has taken place within the mother's body, we find in the lens-shaped stem-cell the progress of flat, discoid segmentation (Figure 1.57). First two equal segmentation-cells (A) are formed from the ovum. These divide into four (B), then into eight, sixteen (C), thirty-two, sixty-four, and so on. The cleavage of the cells is always preceded by a division of their nuclei. The cleavage surfaces between the segmentation-cells appear at the free surface of the tread as clefts. The first two divisions are vertical to each other, in the form of a cross (B). Then there are two more divisions, which cut the former at an angle of forty-five degrees. The tread, which thus becomes the germinal disk, now has the appearance of an eight-rayed star. A circular cleavage next taking place round the middle, the eight triangular cells divide into sixteen, of which eight are in the middle and eight distributed around (C). Afterwards circular clefts and radial clefts, directed towards the centre, alternate more or less irregularly (D, E). In most of the amniotes the formation of concentric and radial clefts is irregular from the very first; and so also in the hen's egg. But the final outcome of the cleavage-process is once more the formation of a large number of small cells of a similar nature. As in the case of the fish-ovum, these segmentation-cells form a round, lens-shaped disk, which corresponds to the morula, and is embedded in a small depression of the white yelk. Between the lens-shaped disk of the morula-cells and the underlying white yelk a small cavity is now formed by the accumulation of fluid, as in the fishes. Thus we get the peculiar and not easily recognisable blastula of the bird (Figure 1.58). The small segmentation-cavity (fh) is very flat and much compressed. The upper or dorsal wall (dw) is formed of a single layer of clear, distinctly separated cells; this corresponds to the upper or animal hemisphere of the triton-blastula (Figure 1.45). The lower or ventral wall of the flat dividing space (vw) is made up of larger and darker segmentation-cells; it corresponds to the lower or vegetal hemisphere of the blastula of the water-salamander (Figure 1.45 dz). The nuclei of the yelk-cells, which are in this case especially numerous at the edge of the lens-shaped blastula, travel into the white yelk, increase by cleavage, and contribute even to the further growth of the germinal disk by furnishing it with food-stuff.

(FIGURE 1.57. Diagram of discoid segmentation in the bird's ovum (magnified about ten times). Only the formative yelk (the tread) is shown in these six figures (A to F), because cleavage only takes place in this. The much larger food-yelk, which does not share in the cleavage, is left out and merely indicated by the dark ring without.)

The invagination or the folding inwards of the bird-blastula takes place in this case also at the hinder pole of the subsequent chief axis, in the middle of the hind border of the round germinal disk (Figure 1.59 s). At this spot we have the most brisk cleavage of the cells; hence the cells are more numerous and smaller here than in the fore-half of the germinal disk. The border-swelling or thick edge of the disk is less clear but whiter behind, and is more sharply separated from contiguous parts. In the middle of its hind border there is a white, crescent-shaped groove--Koller's sickle-groove (Fig 1.59 s); a small projecting process in the centre of it is called the sickle-knob (sk). This important cleft is the primitive mouth, which was described for a long time as the "primitive groove." If we make a vertical section through this part, we see that a flat and broad cleft stretches under the germinal disk forwards from the primitive mouth; this is the primitive gut (Figure 1.60 ud). Its roof or dorsal wall is formed by the folded upper part of the blastula, and its floor or ventral wall by the white yelk (wd), in which a number of yelk-nuclei (dk) are distributed. There is a brisk multiplication of these at the edge of the germinal disk, especially in the neighbourhood of the sickle-shaped primitive mouth.

We learn from sections through later stages of this discoid bird-gastrula that the primitive gut-cavity, extending forward from the primitive mouth as a flat pouch, undermines the whole region of the round flat lens-shaped blastula (Figure 1.61 ud). At the same time, the segmentation-cavity gradually disappears altogether, the folded inner germinal layer (ik) placing itself from underneath on the overlying outer germinal layer (ak). The typical process of invagination, though greatly disguised, can thus be clearly seen in this case, as Goette and Rauber, and more recently Duval (Figure 1.61), have shown.

(FIGURE 1.58. Vertical section of the blastula of a hen (discoblastula). fh segmentation-cavity, dw dorsal wall of same, vw ventral wall, passing directly into the white yelk (wd) (From Duval.)

FIGURE 1.59. The germinal disk of the hen's ovum at the beginning of gastrulation; A before incubation, B in the first hour of incubation. (From Koller.) ks germinal-disk, V its fore and H its hind border; es embryonic shield, s sickle-groove, sk sickle knob, d yelk.

FIGURE 1.60. Longitudinal section of the germinal disk of a siskin (discogastrula). (From Duval.) ud primitive gut, vl, hl fore and hind lips of the primitive mouth (or sickle-edge); ak outer germinal layer, ik inner germinal layer, dk yelk-nuclei, wd white yelk.

FIGURE 1.61. Longitudinal section of the discoid gastrula of the nightingale. (From Duval.) ud primitive gut, vl, hl fore and hind lips of the primitive mouth; ak, ik outer and inner germinal layers; vr fore-border of the discogastrula.)

The older embryologists (Pander, Baer, Remak), and, in recent times especially, His, Kolliker, and others, said that the two primary germinal layers of the hen's ovum--the oldest and most frequent subject of observation!--arose by horizontal cleavage of a simple germinal disk. In opposition to this accepted view, I affirmed in my Gastraea Theory (1873) that the discoid bird-gastrula, like that of all other vertebrates, is formed by folding (or invagination), and that this typical process is merely altered in a peculiar way and disguised by the immense accumulation of food-yelk and the flat spreading of the discoid blastula at one part of its surface. I endeavoured to establish this view by the derivation of the vertebrates from one source, and especially by proving that the birds descend from the reptiles, and these from the amphibia. If this is correct, the discoid gastrula of the amniotes must have been formed by the folding-in of a hollow blastula, as has been shown by Remak and Rusconi of the discoid gastrula of the amphibia, their direct ancestors. The accurate and extremely careful observations of the authors I have mentioned (Goette, Rauber, and Duval) have decisively proved this recently for the birds; and the same has been done for the reptiles by the fine studies of Kupffer, Beneke, Wenkebach, and others. In the shield-shaped germinal disk of the lizard (Figure 1.62), the crocodile, the tortoise, and other reptiles, we find in the middle of the hind border (at the same spot as the sickle groove in the bird) a transverse furrow (u), which leads into a flat, pouch-like, blind sac, the primitive gut. The fore (dorsal) and hind (ventral) lips of the transverse furrow correspond exactly to the lips of the primitive mouth (or sickle-groove) in the birds.

(FIGURE 1.62. Germinal disk of the lizard (Lacerta agilis). (From Kupffer.) u primitive mouth, s sickle, es embryonic shield, hf and df light and dark germinative area.)

The gastrulation of the mammals must be derived from this special embryonic development of the reptiles and birds. This latest and most advanced class of the vertebrates has, as we shall see afterwards, evolved at a comparatively recent date from an older group of reptiles; and all these amniotes must have come originally from a common stem-form. Hence the distinctive embryonic process of the mammal must have arisen by cenogenetic modifications from the older form of gastrulation of the reptiles and birds. Until we admit this thesis we cannot understand the formation of the germinal layers in the mammal, and therefore in man.

I first advanced this fundamental principle in my essay On the Gastrulation of Mammals (1877), and sought to show in this way that I assumed a gradual degeneration of the food-yelk and the yelk-sac on the way from the proreptiles to the mammals. "The cenogenetic process of adaptation," I said, "which has occasioned the atrophy of the rudimentary yelk-sac of the mammal, is perfectly clear. It is due to the fact that the young of the mammal, whose ancestors were certainly oviparous, now remain a long time in the womb. As the great store of food-yelk, which the oviparous ancestors gave to the egg, became superfluous in their descendants owing to the long carrying in the womb, and the maternal blood in the wall of the uterus made itself the chief source of nourishment, the now useless yelk-sac was bound to atrophy by embryonic adaptation."

My opinion met with little approval at the time; it was vehemently attacked by Kolliker, Hensen, and His in particular. However, it has been gradually accepted, and has recently been firmly established by a large number of excellent studies of mammal gastrulation, especially by Edward Van Beneden's studies of the rabbit and bat, Selenka's on the marsupials and rodents, Heape's and Lieberkuhn's on the mole, Kupffer and Keibel's on the rodents, Bonnet's on the ruminants, etc. From the general comparative point of view, Carl Rabl in his theory of the mesoderm, Oscar Hertwig in the latest edition of his Manual (1902), and Hubrecht in his Studies in Mammalian Embryology (1891), have supported the opinion, and sought to derive the peculiarly modified gastrulation of the mammal from that of the reptile.

(FIGURE 1.63. Ovum of the opossum (Didelphys) divided into four. (From Selenka.) b the four segmentation-cells, r directive body, c unnucleated coagulated matter, p, albumin-membrane.)

In the meantime (1884) the studies of Wilhelm Haacke and Caldwell provided a proof of the long-suspected and very interesting fact, that the lowest mammals, the monotremes, LAY EGGS, like the birds and reptiles, and are not viviparous like the other mammals. Although the gastrulation of the monotremes was not really known until studied by Richard Semon in 1894, there could be little doubt, in view of the great size of their food-yelk, that their ovum-segmentation was discoid, and led to the formation of a sickle-mouthed discogastrula, as in the case of the reptiles and birds. Hence I had, in 1875 (in my essay on The Gastrula and Ovum-segmentation of Animals), counted the monotremes among the discoblastic vertebrates. This hypothesis was established as a fact nineteen years afterwards by the careful observations of Semon; he gave in the second volume of his great work, Zoological Journeys in Australia (1894), the first description and correct explanation of the discoid gastrulation of the monotremes. The fertilised ova of the two living monotremes (Echidna and Ornithorhynchus) are balls of one-fifth of an inch in diameter, enclosed in a stiff shell; but they grow considerably during development, so that when laid the egg is three times as large. The structure of the plentiful yelk, and especially the relation of the yellow and the white yelk, are just the same as in the reptiles and birds. As with these, partial cleavage takes place at a spot on the surface at which the small formative yelk and the nucleus it encloses are found. First is formed a lens-shaped circular germinal disk. This is made up of several strata of cells, but it spreads over the yelk-ball, and thus becomes a one-layered blastula. If we then imagine the yelk it contains to be dissolved and replaced by a clear liquid, we have the characteristic blastula of the higher mammals. In these the gastrulation proceeds in two phases, as Semon