2. _Cartilaginous or Chondral Skeleton._--(A) Vertebral column.[27] In
the embryonic connective tissue or mesenchyme lying just outside the
primary sheath of the notochord there are developed a dorsal and a
ventral series of paired nodules of cartilage known as _arcualia_ (fig.
13, d.a, v.a). The dorsal arcualia are commonly prolonged upwards by
supradorsal cartilages which complete the _neural arches_ and serve to
protect the spinal cord. The ventral arcualia become, in the tail region
only, also incorporated in complete arches--the _haemal arches_. In
correlation with the flattening of the body of the fish from side to
side the arches are commonly prolonged into elongated neural or haemal
spines.
The relations of the arcualia to the segmentation of the body, as
shown by myotomes and spinal nerves, is somewhat obscure. The
mesenchyme in which they arise is segmental in origin (sclerotom),
which suggests that they too may have been primitively segmental, but
in existing fishes there are commonly two sets of arcualia to each
body segment.
In gnathostomatous fishes the arcualia play a most important part in
that cartilaginous tissue derived from them comes into special
relationships with the notochord and gives rise to the vertebral column
which functionally replaces this notochord in most of the fishes. This
replacement occurs according to two different methods, giving rise to
the different types of vertebral column known as chordacentrous and
arcicentrous.
(a) Chordacentrous type. An incipient stage in the evolution of a
chordacentrous vertebral column occurs in the Dipneusti, where cartilage
cells from the arcualia become amoeboid and migrate into the substance
of the secondary sheath, boring their way through the primary sheath
(fig. 13, C). They wander throughout the whole extent of the secondary
sheath, colonizing it as it were, and settle down as typical stationary
cartilage cells. The secondary sheath is thus converted into a cylinder
of cartilage. In Selachians exactly the same thing takes place, but in
recent forms development goes a step further, as the cartilage cylinder
becomes broken into a series of segments, known as vertebral centra. The
wall of each segment becomes much thickened in the middle so that the
notochord becomes constricted within each centrum and the space occupied
by it is shaped like the cavity of a dice-box. When free from notochord
and surrounding tissues such a cartilaginous centrum presents a deep
conical cavity at each end (_amphicoelous_).
[Illustration: From Wiedersheim, _Grundriss der vergleichenden
Anatomie_, by permission of Gustav Fischer.
FIG. 13.--Diagrammatic transverse sections to illustrate the morphology
of the vertebral column.
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