Braem, who has made a critical examination of the germ-layer
theory,[103] has found it impossible to give a morphological definition
of a germ-layer, and has adopted a physiological criterion. He thinks
that in whatever way a germ-layer arises, whether by folding, or by
delamination, etc., it exists independently of its method or place of
origin. A layer is not endodermal because it forms the inner wall of a
gastrula, but it is endodermal because it develops into the digestive
tract. The germ-layers of different forms are only similarly placed, but
whether they are homologous will depend on other things. On this view
the inner tube of the ascidian bud that gives rise to both digestive
tract and to the nervous system is simply an indifferent layer until it
gives rise to these structures. Its cells may be looked upon as
indifferent, as are those of the blastula. Thus the difficulty of the
morphologist is not solved, but the knot is cut. For Braem the
germ-layers are convenient terms, since he rejects any historical
significance that they may have, and it is just this side of the
question that the morphologist has attempted to work out. While the
evidence shows that the germ-layers cannot have any such final
attributes as embryologists have attempted to assign to them, and that
Braem has called attention to the real and important problems connected
with the study of development, yet it may still be admitted without
endangering the newer point of view, that there may be also an
historical question in connection with the germ-layers, if not in the
sense of a repetition of an ancestral adult gastræa, yet in the sense
that similarity in embryonic development may in some cases find its
historical explanation in a common descent.
If in the light of this discussion we turn to the phenomena of
regeneration, we again find evidence showing that the germ-layer theory
fails to apply in all cases. It has been pointed out that in
lumbriculus, and in the naids, the new mesoderm is derived from the
ectoderm, and does not come from the old mesodermal tissues. The
mesoderm of the embryo in annelids is derived from one, and later from
two, superficial cells of the blastula,[104] that push in about the time
of gastrulation. They cannot, at this time, be referred to one layer
rather than to the other. It cannot be affirmed, therefore, that in
regeneration, the mesoderm arises from a different layer from that in
the embryo, but neither can this be denied. The most important point in
this connection is that the new mesoderm comes from the ectoderm that is
already differentiated, and not from the mesodermal tissues. It is
clear, however, that while the lining of the pharynx in the embryo is
ectodermal, it is endodermal in the regenerated part.
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