Since this paper of Froriep's, it has been generally recognized, and
Gegenbaur has accepted Froriep's view, that the three hindmost metameres,
which distinctly show the characteristics of vertebræ, belong to the spinal
and not to the cranial region, so that the metameric segmentation of the
cranial region proper has become {262}more and more associated with the
branchial segmentation. Froriep's discovery of the rudimentary branchial
sense-organs as a factor in the segmentation question has led Beard to the
conclusion that the olfactory and auditory organs represent in a permanent
form two of these rudimentary branchial sense-organs. He therefore includes
both the olfactory and auditory nerves in his list of cranial segmental
nerves, and makes eleven cranial branchial segments in front of the spinal
segments represented by the hypoglossal.
A still larger number of cranial segments is supposed to exist, according
to the researches of Dohrn and Killian, in the embryos of _Torpedo
ocellata_. The former, holding to the view that vertebrates arose from
annelids, considered that the head was formed of a series of metameres, to
each one of which a mesoderm-segment, a gill-arch, a gill-cleft, a
segmental nerve and vessel belonged. He found in the front head-region of a
Torpedo embryo, corresponding to van Wijhe's first four somites, no less
than twelve to fifteen mesoderm segments, and concluded, therefore, that
the eye-muscle nerves, especially the oculomotor, represented many
segmental nerves, and were not the nerves of single segments; so, also,
that the inferior maxillary part of the trigeminal and the hyoid nerve of
the facial are probably not single nerves, but a fusion of several. Killian
comes to much the same conclusion as Dohrn, for he finds seventeen to
eighteen separate mesoderm segments in the head, of which twelve belong to
the trigeminal and facial region.
Since Rabl's paper, a number of papers have appeared, especially from
America, dealing with yet another criterion of the original segmentation of
the head, viz. a series of divisions of the central nervous system itself,
which are seen at a very early stage of development, and are called
neuromeres; the divisions in the cranial region being known as
encephalomeres, and those of the spinal region as myomeres. Locy's paper
has especially brought these divisions into prominence as a factor in the
question of segmentation. They are essentially segments of the epiblast and
not of the mesoblast; they are conspicuous in very early stages, and appear
to be in relation with the cranial nerves, according to Locy. He recognizes
in _Squalus acanthias_, in front of the spino-occipital region, fourteen
pairs of such encephalomeres and a median unsegmented termination, which
may represent one more pair fused in the middle line, making at least
fifteen. He distributes these fifteen segments as follows: {263}fore-brain
three and unsegmented termination, mid-brain two, and hind-brain nine.
Public-domain text, read in full here on John Shaqi.
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