Anatomy, Comparative; Embryology, Human; Evolution; Human beings -- Origin
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.
Fig.63. Ovum of the opossum (Didelphys) divided into four. Fig. 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.
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
rightly observes: firstly, formation of the entoderm by cleavage at the
centre and further growth at the edge; secondly, invagination. In the
monotremes more primitive conditions have been retained better than in
the reptiles and birds. In the latter, before the commencement of the
gastrula-folding, we have, at least at the periphery, a two-layered
embryo forming from the cleavage. But in the monotremes the formation
of the cenogenetic entoderm does not precede the invagination; hence in
this case the construction of the germinal layers is less modified than
in the other amniota.
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