method the endoderm is at first a solid mass, in which the enteric space
makes its appearance later by excavation. In the delaminate method the
enteric space is at first without a blastopore, and sometimes it never
acquires this opening, but a blastopore is frequently formed, and the
two-layered gastrula stage is reached, though by a very different route
from that taken in the formation of the invaginate gastrula. According to
the layer-theory, these two layers are homologous throughout the series
of Metazoa; their limits can always be accurately defined, they give rise
to the same organs in all cases, and the adult organs (excluding the
mesodermal organs) can be traced back to one or other of them with
absolute precision. Thus the ectoderm gives rise to the epidermis, to the
nervous system, and to the lining of the stomodaeum and proctodaeum, if
such parts of the alimentary canal are present. The endoderm, on the
other hand, gives rise to the lining of the enteron, and of the glands
which open into it.
Mesoderm.
So far as these two layers are concerned, and excluding the mesoderm, it
would appear that the layer-theory does apply in a very remarkable
manner to the whole of the Metazoa. But even here, when the actual facts
are closely scanned, there are found to be difficulties, which appear to
indicate that the theory may not perhaps be such an infallible guide as
it seems at first sight. Leaving out of consideration the case of the
Mammalia, in which the differentiation of the segmented ovum is not into
ectoderm and endoderm, and the case of the sponges, the most important
of these difficulties concern the stomodaeum and proctodaeum. The best
case to examine is that of _Peripatus capensis_, in which the blastopore
is at first a long slit, and gives rise to both the mouth and the anus
of the adult. Here there is always found at the lips of the blastopore,
and extending for a short distance inwards as enteric lining, a certain
amount of tissue, which by its characters must be regarded as ectoderm.
Now, in the closure of the blastopore between the mouth and anus, this
tissue, which at the mouth and anus develops into the lining of the
stomodaeum and proctodaeum, is left inside, and actually gives rise to
the median ventral epithelium of the alimentary canal. Hence the
development of _Peripatus capensis_ suggests the conclusion, if we
strictly apply the layer-theory, that a considerable portion of the true
mesenteron is lined by ectoderm, and is not homologous with the
corresponding portion of the mesenteron of other animals--a conclusion
which will on all hands be admitted to be absurd. The difficulties in
the application of the layer-theory become vastly greater when the
origin and fate of the mesoderm is considered. The mesoderm is, if we
may judge from the number of organs which are derived from it, much the
most important of the three layers. It generally arises later than the
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