Traces of what appear to be pre-spiracular clefts exist in the embryos
of various forms. Perhaps the most remarkable of these is to be found in
the larval Crossopterygian,[12] and apparently also in _Amia_[13] at
least, amongst the other ganoids, where a pair of entodermal pouches
become cut off from the main entoderm and, establishing an opening to
the exterior, give rise to the lining of the cement organs of the larva.
Posteriorily there is evidence that the extension backwards of the
series of gill clefts was much greater in the primitive fishes. In the
surviving sharks (_Chlamydoselachus_ and _Notidanus cinereus_), there
still exist in the adult respectively six and seven branchial clefts,
while in embryonic Selachians there are frequently to be seen pouch-like
outgrowths of entoderm apparently representing rudimentary gill pouches
but which never develop. Further evidence of the progressive reduction
in the series of clefts is seen in the reduction of their functional
activity at the two ends of the series. The spiracle, even where
persisting in the adult, has lost its gill lamellae either entirely or
excepting a few vestigial lamellae forming a "pseudobranch" on its
anterior wall (Selachians, sturgeons). A similar reduction affects the
lamellae on the anterior wall of the hyobranchial cleft (except in
Selachians) and on the posterior wall of the last branchial cleft.
A pseudobranch is frequently present in Teleostomes on the anterior
wall of the hyobranchial cleft, i.e. on the inner or posterior face of
the operculum. It is believed by some morphologists to belong really
to the cleft in front.[14]
_Phylogeny._--The phylogeny of the gill clefts or pouches is
uncertain. The only organs of vertebrates comparable with them
morphologically are the enterocoelic pouches of the entoderm which
give rise to the mesoderm. It is possible that the respiratory
significance of the wall of the gill cleft has been secondarily
acquired. This is indicated by the fact that they appear in some cases
to be lined by an ingrowth of ectoderm. This suggests that there may
have been a spreading inwards of respiratory surface from the external
gills. It is conceivable that before their walls became directly
respiratory the gill clefts served for the pumping of fresh water over
the external gills at the bases of which they lie.
_Lung._--As in the higher vertebrates, there develops in all the main
groups of gnathostomatous fishes, except the Selachians, an outgrowth of
the pharyngeal wall intimately associated with gaseous interchange. In
the Crossopterygians and Dipnoans this pharyngeal outgrowth agrees
exactly in its mid-ventral origin and in its blood-supply with the lungs
of the higher vertebrates, and there can be no question about its being
morphologically the same structure as it is also in function.
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