Anatomy, Comparative; Embryology, Human; Evolution; Human beings -- Origin
The four secondary germinal layers are so distributed in the structure
of the body in all the coelomaria (or all metazoa that have a
body-cavity) that the outer two, joined fast together, constitute the
body-wall, and the inner two the ventral wall; the two walls are
separated by the cavity of the coelom. Each of the walls is made up of
a limiting layer and a middle layer. The two limiting layers chiefly
give rise to epithelia, or covering-tissues, and glands and nerves,
while the middle layers form the great bulk of the fibrous tissue,
muscles, and connective matter. Hence the latter have also been called
fibrous or muscular layers. The outer middle layer, which lies on the
inner side of the skin-sense-layer, is the skin fibre-layer; the inner
middle layer, which attaches from without to the ventral glandular
layer, is the ventral fibre layer. The former is usually called
briefly the parietal, and the latter the visceral layer or mesoderm.
Of the many different names that have been given to the four secondary
germinal layers, the following are those most in use to-day:--
1. Skin-sense-layer (outer limiting layer) and 2. Skin-fibre-layer
(outer middle layer).
I. Neural layer (neuroblast) and II. Parietal layer (myoblast). The
two secondary germinal layers of the body-wall: 1. Epithelial. 2.
Fibrous.
3. Gut-fibre-layer (inner middle layer) and 4. Gut-gland-layer (inner
limiting layer).
III. Visceral layer (gonoblast) and IV. Enteral layer (enteroblast).
The two secondary germinal layers of the gut-wall: 3. Fibrous. 4.
Epithelial.
The first scientist to recognise and clearly distinguish the four
secondary germinal layers was Baer. It is true that he was not quite
clear as to their origin and further significance, and made several
mistakes in detail in explaining them. But, on the whole, their great
importance did not escape him. However, in later years his view had to
be given up in consequence of more accurate observations. Remak then
propounded a three-layer theory, which was generally accepted. These
theories of cleavage, however, began to give way thirty years ago,
when Kowalevsky (1871) showed that in the case of Sagitta (a very
clear and typical subject of gastrulation) the two middle germinal
layers and the two limiting layers arise not by cleavage, but by
folding--by a secondary invagination of the primary inner germ-layer.
This invagination or folding proceeds from the primitive mouth, at the
two sides of which (right and left) a couple of pouches are formed. As
these coelom-pouches or coelom-sacs detach themselves from the
primitive gut, a double body-cavity is formed (Figures 1.74 to 1.76).
(FIGURE 1.77. Coelomula of sagitta, in section. (From Hertwig.) D
dorsal side, V ventral side, ik inner germinal layer, mv visceral
mesoblast, lh body-cavity, mp parietal mesoblast, ak outer germinal
layer.)
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