Fig. 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.
Fig. 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 Kölliker, 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 Lieberkühn’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.
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.