Behind the pharynx comes the main portion of the alimentary canal
concerned with the digestion and absorption of the food. This forms a
tube varying greatly in length, more elongated and coiled in the higher
Teleostomes, shorter and straighter in the Selachians, Dipnoans and
lower Teleostomes. The oesophagus or gullet, usually forming a short,
wide tube, leads into the glandular, more or less dilated stomach. This
is frequently in the form of a letter J, the longer limb being
continuous with the gullet, the shorter with the intestine. The curve of
the J may be as in _Polypterus_ and the perch produced backwards into a
large pocket. The intestine is usually marked off from the stomach by a
ring-like sphincter muscle forming the pyloric valve. In the lower
gnathostomatous fishes (Selachians, Crossopterygians, Dipnoans,
sturgeons) the intestine possesses the highly characteristic spiral
valve, a shelf-like projection into its lumen which pursues a spiral
course, and along the turns of which the food passes during the course
of digestion. From its universal occurrence in the groups mentioned we
conclude that it is a structure of a very archaic type, once
characteristic of ancestral Gnathostomata; a hint as to its
morphological significance is given by its method of development.[15] In
an early stage of development the intestinal rudiment is coiled into a
spiral and it is by the fusion together of the turns that the spiral
valve arises. The only feasible explanation of this peculiar method of
development seems to lie in the assumption that the ancestral
gnathostome possessed an elongated, coiled intestine which subsequently
became shortened with a fusion of its coils. In the higher fishes the
spiral valve has disappeared--being still found, however, in a reduced
condition in _Amia_ and _Lepidosteus_, and possibly as a faint vestige
in one or two Teleosts (certain _Clupeidae_[16] and _Salmonidae_[17]).
In the majority of the Teleosts the absence of spiral valves is coupled
with a secondary elongation of the intestinal region, which in extreme
cases (_Loricariidae_) may be accompanied by a secondary spiral coiling.
The terminal part of the alimentary canal--the cloaca--is characterized
by the fact that into it open the two kidney ducts. In Teleostomes the
cloaca is commonly flattened out, so that the kidney ducts and the
alimentary canal come to open independently on the outer surface.
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