2. The pneumatophore or air-bladder (fig. 68, n), for passive
locomotion, forming a float which keeps the cormus at or near the
surface of the water. The pneumatophore arises from the ectoderm as a
pit or invagination, part of which forms a gas-secreting gland, while
the rest gives rise to an air-sack lined by a chitinous cuticle. The
orifice of invagination forms a pore which may be closed up or may
form a protruding duct or funnel. As in the analogous swim-bladder of
fishes, the gas in the pneumatophore can be secreted or absorbed,
whereby the specific gravity of the body can be diminished or
increased, so as to cause it to float nearer the surface or at a
deeper level. Never more than one pneumatophore is found in a cormus,
and when present it is always situated at the highest point above the
swimming bells, if these are present also. In _Velella_ the
pneumatophore becomes of complex structure and sends air-tubes, lined
by a chitin and resembling tracheae, down into the compact coenosarc,
thus evidently serving a respiratory as well as a hydrostatic
function.
Divergent views have been held as to the morphological significance of
the pneumatophore. E. Haeckel regarded the whole structure as a
glandular ectodermal pit formed on the ex-umbral surface of a
medusa-person. C. Chun and, more recently, R. Woltereck [59], on the
other hand, have shown that the ectodermal pit which gives rise to the
pneumatophore represents an entocodon. Hence the cavity of the
air-sack is equivalent to a sub-umbral cavity in which no manubrium is
formed, and the pore or orifice of invagination would represent the
margin of the umbrella. In the wall of the sack is a double layer of
endoderm, the space between which is a continuation of the
coelenteron. By coalescence of the endoderm-layers, the coelenteron
may be reduced to vessels, usually eight in number, opening into a
ring-sinus surrounding the pore. Thus the disposition of the
endoderm-cavities is roughly comparable to the gastrovascular system
of a medusa.
The difference between the theories of Haeckel and Chun is connected
with a further divergence in the interpretation of the stem or axis of
the cormus. Haeckel regards it as the equivalent of the manubrium, and
as it is implanted on the blind end of the pneumatophore, such a view
leads necessarily to the air-sack and gland being a development on the
ex-umbral surface of the medusa-person. Chun and Woltereck, on the
other hand, regard the stem as a _stolo prolifer_ arising from the
aboral pole, that is to say, from the ex-umbrella, similar to that
which grows out from the ex-umbral surface of the embryo of the
Narcomedusae and produces buds, a view which is certainly supported by
the embryological evidence to be adduced shortly.
In the siphosome the following types of appendages occur:--
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