(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).
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From Wiedersheim, Grundriss der vergleichenden Anatomie, by permission of Gustav Fischer. | ||||||||||||||||||||||||||||||
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Fig. 13.—Diagrammatic transverse sections to illustrate the morphology of the vertebral column. | ||||||||||||||||||||||||||||||
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A secondary modification of the centrum consists in the calcification of certain zones of the cartilaginous matrix. The precise arrangement of these calcified zones varies in different families and affords characters which are of taxonomic importance in palaeontology where only skeletal structures are available (see Selachians).
(b) Arcicentrous type. Already in the Selachians the vertebral column is to a certain extent strengthened by the broadening of the basis of the arcualia so as partially to surround the centra. In the Teleostomes, with the exceptions of those ganoids mentioned, the expanded bases of the arcualia undergo complete fusion to form cartilaginous centra which, unlike the chordacentrous centra, lie outside the primary sheath (figs. 13, D and E). In these forms no invasion of the secondary sheath by cartilage cells takes place. The composition of the groups of arcualia which give rise to the individual centrum is different in different groups. The end result is an amphicoelous or biconcave centrum in general appearance much like that of the Selachian.
In Lepidosteus the spaces between adjacent centra become filled by a secondary development of intervertebral cartilage which then splits in such a way that the definitive vertebrae are opisthocoelous, i.e. concave behind, convex in front.
Ribs.—In the Crossopterygians a double set of “ribs” is present on each side of the vertebral column, a ventral set lying immediately outside the splanchnocoelic lining and apparently serially homologous with the haemal arches of the caudal region, and a second set passing outwards in the thickness of the body wall at a more dorsal level. In the Teleostomes and Dipnoans only the first type is present; in the Selachians only the second. It would appear that it is the latter which is homologous with the ribs of vertebrates above fishes.
Median Fin Skeleton.—the foundation of the skeleton of the median fins consists of a series of rod-like elements, the radialia, each of which frequently is segmented into three portions. In a few cases the radialia correspond segmentally with the neural and haemal arches (living Dipnoans, Pleuracanthus tail region) and this suggests that they represent morphologically prolongations of the neural and haemal spines. That this is so is rendered probable by the fact that we must regard the evolution of the system of median fins as commencing with a simple flattening of the posterior part of the body. It is only natural to suppose that the edges of the flattened region would be at first supported merely by prolongations of the already existing spinous processes. In the Cyclostomes (where they are branched) and in the Selachians, the radialia form the main supports of the fin, though already in the latter they are reinforced by a new set of fin rays apparently related morphologically to the osseous or placoid skeleton (see below).
The series of radialia tends to undergo the same process of local concentration which characterizes the fin-fold as a whole. In its extreme form this leads to complete fusion of the basal portions of a number of radialia (dorsal fins of Holoptychius and various Selachians, and anal fin of Pleuracanthus). In view of the identity in function it is not surprising that a remarkable resemblance exists between the mechanical arrangements (of skeleton, muscles, &c.), of the paired and unpaired fins. The resemblance to paired fins becomes very striking in some of the cases where the basal fusion mentioned above takes place (Pleuracanthus).
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Fig. 14.—Chondrocranium of a young Lepidosiren, showing the suspension of the lower jaw by the upper portion of the mandibular arch. (After Agar.) | ||||||||||||
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The palatopterygoid bar (p.pt) is represented by a faint vestige which disappears before the stage figured. |
(B) Chondrocranium[1].—In front of the vertebral column lies the cartilaginous trough, the chondrocranium, which protects the brain. This consists of a praechordal portion—developed out of a pair of lateral cartilaginous rods—the trabeculae cranii—and a parachordal portion lying on either side of the anterior end of the notochord. This arises in development
- ↑ For development cf. Gaupp in Hertwig’s Handbuch der Entwickelungslehre.
