Again, Kupffer, in his recent papers on the embryology of Ammocoetes,
asserts that especial information as to the number of primitive segments is
afforded by the appearance in the early stages of a series of epibranchial
ganglia in connection with the cranial nerves, which remain permanently in
the case of the vagus nerves, but disappear in the case of pro-otic nerves.
He considers that the evidence points to the number of segments in the mid-
and hind-brain region as being primitively fifteen, viz. six segments
belonging to the trigeminal and abducens group, three segments belonging
respectively to the facial, auditory, and glossopharyngeal, and six to the
vagus.
From this sketch we see that the modern tendency is to make six segments at
least out of the region of the trigeminal nerves rather than two. In this
region, as already mentioned, the evidence of segmentation is based more
clearly on the somatic than on the splanchnic segments. We ought,
therefore, in the first place, to consider the teaching of the eye-muscles
and their nerves and the coelomic cavities in connection with them, and see
whether the hypothesis that such muscles represent the original
dorso-ventral somatic muscles of the palæostracan ancestor is in harmony
with and explains the facts of modern research.
EYE-MUSCLES AND THEIR NERVES.
The only universally recognized somatic nerves belonging to these segments
which exist in the adult are the nerves to the eye-muscles, of which,
according to van Wijhe, the oculomotor is the nerve of the 1st segment, the
trochlearis of the 2nd, and the abducens of the 3rd; while the nerves and
muscles belonging to the 4th and 5th segments, _i.e._ the 2nd facial and
glossopharyngeal segments respectively, show only the merest rudiments, and
do not exist in the adult. One significant fact appears in this statement
of van Wijhe, and is accepted by all those who follow him, viz. that the
oculomotor nerve has equal segmental value with the trochlearis and the
abducens, although it supplies a number of muscles, each of which, on the
face of it, has the same anatomical value as the superior oblique or
external rectus. Dohrn alone, as far as I know, as already pointed out,
insists upon the multiple character of the oculomotor nerve.
{264}As far as the anatomist is concerned, the evidence is becoming clearer
and clearer that the nucleus of the IIIrd nerve is a composite ganglion
composed of a number of nuclei, each similar to that of the trochlearis, so
that if the trochlearis nucleus is a segmental motor nucleus, then the
oculomotor nucleus is a combined nucleus belonging to at least four
segmental nerves, each of which has the same value as that of the
trochlearis.
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