I did not deal with them in that logical sequence because it was necessary
for their understanding to introduce first the conception of modified
appendages as important factors in any consideration of vertebrate
segments; a conception which followed naturally after the evidence afforded
by the skeleton in Chapter III., and by the branchial segments in Chapter
IV. So, too, now, although the discussion of the prosomatic segmentation
ought logically to follow immediately on that of the mesosomatic
segmentation, I have determined to devote this chapter to the evidence of
the olfactory organs, because the arguments as to the segments belonging to
the trigeminal nerve-group are so much easier to understand if the position
of the olfactory apparatus is first made clear.
In all vertebrates the nose is double and opens into the pharynx, until we
descend to the fishes, where the whole group Pisces has been divided into
two subsidiary groups, Monorhinæ and Amphirhinæ, according as they possess
a median unpaired olfactory opening, or a paired opening. The Monorhinæ
include only the Cyclostomata--the lampreys and hag-fishes.
In the lampreys the single olfactory tube ends blindly, while in the
hag-fishes it opens into the pharynx. In the lamprey, both in Petromyzon
and Ammocoetes, the opening of this nasal tube is a conspicuous object on
the dorsal surface of the head in front of the transparent spot which
indicates the position of the right median eye. It is especially
significant, as showing the primitive nature of this median olfactory
passage, that a perfectly similar opening in the {220}same position is
always found in the dorsal head-shields of all the Cephalaspidæ and
Tremataspidæ, as will be explained more fully in Chapter X.
All the evidence points to the conclusion that the olfactory apparatus of
the vertebrate originated as a single median tube, containing the special
olfactory sense-epithelium, which, although median and single, was
innervated by the olfactory nerve of each side. The external opening of
this tube in the lamprey is dorsal. How does it terminate ventrally?
The ventral termination of this tube is most instructive and suggestive. It
terminates blindly at the very spot where the infundibular tube terminates
blindly and the notochord ends. After transformation, when the Ammocoete
becomes the Petromyzon, the tube still ends blindly, and does not open into
the pharynx as in Myxine; it, however, no longer terminates at the
infundibulum, but extends beyond it towards the pharynx.
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