The entocodon is to be regarded, therefore, not as primarily an
ingrowth of ectoderm, but rather as an upgrowth of both body-layers,
in the form of a circular rim (IVa), representing the umbrellar
margin; it is comparable to the bulging that forms the umbrella in the
direct method of budding, but takes place before a manubrium is
formed, and is greatly reduced in size, so as to become a little pit.
By a simple modification, the open pit becomes a solid ectodermal
ingrowth, just as in Teleostean fishes the hollow medullary tube, or
the auditory pit of other vertebrate embryos, is formed at first as a
solid cord of cells, which acquires a cavity secondarily. Moreover,
the entocodon, however developed, gives rise at first to a closed
cavity, representing a closing over of the umbrella, temporary in the
bud destined to be a free medusa, but usually permanent in the sessile
gonophore. As has been shown above, the closing up of the sub-umbral
cavity is one of the earliest degenerative changes in the evolution of
the gonophore, and we may regard it as the umbrellar fold taking on a
protective function, either temporarily for the bud or permanently for
the gonophore.
To sum up, the entocodon is a precocious formation of the umbrella,
closing over to protect the organs in the umbrellar cavity. The
possession of an entocodon proves the medusa-nature of the bud, and
can only be explained on the theory that gonophores are degenerate
medusae, and is inexplicable on the opposed view that medusae are
derived from gonophores secondarily set free. In the sporosac,
however, the medusa-individual has become so degenerate that even the
documentary proof, so to speak, of its medusoid nature may have been
destroyed, and only circumstantial evidence of its nature can be
produced.
4. _Germinal Budding._--This method of budding is commonly described as
budding from a single body-layer, instead of from both layers. The layer
that produces the bud is invariably the ectoderm, i.e. the layer in
which, in Hydromedusae, the generative cells are lodged; and in some
cases the buds are produced in the exact spot in which later the gonads
appear. From these facts, and from those of the sporogony, to be
described below, we may regard budding to this type as taking place from
the germinal epithelium rather than from ordinary ectoderm.
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