Anatomy, Comparative; Embryology, Human; Evolution; Human beings -- Origin
All these amphiblastic vertebrates, Petromyzon and Cestracion,
Accipenser and Ceratodus, and also the salamanders and batrachia,
belong to the old, conservative groups of our stem. Their unequal
ovum-segmentation and gastrulation have many peculiarities in detail,
but can always be reduced with comparative ease to the original
cleavage and gastrulation of the lowest vertebrate, the amphioxus; and
this is little removed, as we have seen, from the very simple
archigastrula of the Sagitta and Monoxenia (see Figures 1.29 to 1.36).
All these and many other classes of animals generally agree in the
circumstance that in segmentation their ovum divides into a large
number of cells by repeated cleavage. All such ova have been called,
after Remak, "whole-cleaving" (holoblasta), because their division
into cells is complete or total.
(FIGURE 1.51. Gastrulation of ceratodus (from Semon). A and C stage
with four cells, B and D with sixteen cells. A and B are seen from
above, C and D sideways. E stage with thirty-two
cells; F blastula; G gastrula in longitudinal section. fh
segmentation-cavity. gh primitive gut or gastric cavity.)
In a great many other classes of animals this is not the case, as we
find (in the vertebrate stem) among the birds, reptiles, and most of
the fishes; among the insects and most of the spiders and crabs (of
the articulates); and the cephalopods (of the molluscs). In all these
animals the mature ovum, and the stem-cell that arises from it in
fertilisation, consist of two different and separate parts, which we
have called formative yelk and nutritive yelk. The formative yelk
alone consists of living protoplasm, and is the active, evolutionary,
and nucleated part of the ovum; this alone divides in segmentation,
and produces the numerous cells which make up the embryo. On the other
hand, the nutritive yelk is merely a passive part of the contents of
the ovum, a subordinate element which contains nutritive material
(albumin, fat, etc.), and so represents in a sense the provision-store
of the developing embryo. The latter takes a quantity of food out of
this store, and finally consumes it all. Hence the nutritive yelk is
of great indirect importance in embryonic development, though it has
no direct share in it. It either does not divide at all, or only later
on, and does not generally consist of cells. It is sometimes large and
sometimes small, but generally many times larger than the formative
yelk; and hence it is that it was formerly thought the more important
of the two. As the respective significance of these two parts of the
ovum is often wrongly described, it must be borne in mind that the
nutritive yelk is only a secondary addition to the primary cell, it is
an inner enclosure, not an external appendage. All ova that have this
independent nutritive yelk are called, after Remak,
"partially-cleaving" (meroblasta). Their segmentation is incomplete or
partial.
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