Though it does not seem probable that a dorsal and ventral fin can have existed contemporaneously with lateral fins (at least not as continuous fins), yet, judging from such forms as the Rays, there is no reason why small balancing dorsal and caudal fins should not have co-existed with fully developed lateral fins.

Mesoblast. G-K.

The mesoblast in stage F forms two independent lateral plates, each with a splanchnic and somatic layer, and divided, as before explained, into a vertebral portion and a parietal portion. At their peripheral edge these plates are continuous with the general mesoblastic tissue of the non-embryonic part of the blastoderm; except in the free parts of the embryo, where they are necessarily separated from the mesoblast of the yolk-sac, and form completely independent lateral masses of cells.

During the stages G and H, the two layers of which the mesoblast is composed cease to be in contact, and leave between them a space which constitutes the commencement of the body-cavity (Pl. 10, fig. 1). From the very first this cavity is more or less clearly divided into two distinct parts; one of them in the vertebral portion of the plates of mesoblast, the other in the parietal. The cavity in the parietal part of the plates alone becomes the true body-cavity. It extends uninterruptedly through the anterior parts of the embryo, but does not appear in the caudal region, being there indicated only by the presence of two layers in the mesoblast plates. Though fairly wide below, it narrows dorsally before becoming continuous with the cavity in the vertebral plates. The line of junction of the vertebral and parietal plates is a little ventral to the dorsal summit of the alimentary canal (Pl. 10, fig. 5). Owing to the fact that the vertebral plates are split up into a series of segments (protovertebræ), the section of the body-cavity they enclose is necessarily also divided into a series of segments, one for each protovertebra.

Thus the whole body-cavity consists of a continuous parietal space which communicates by a series of apertures with a number of separate cavities enclosed in the protovertebræ. The cavity in each of the protovertebræ is formed of a narrowed dorsal and a dilated ventral segment, the latter on the level of the dorsal aorta (Pl. 11, fig. 5). Cavities are present in all the vertebral plates with the exception of a few far back in the tail; and exist in part of the caudal region posterior to that in which a cavity in the parietal plate is present.

Protovertebræ. Each protovertebra[203] or vertebral segment of the mesoblast plate forms a flattened rectangular body, ventrally continuous with the parietal plate of mesoblast. During stage G the dorsal edge of the protovertebræ is throughout on about a level with the ventral third of the spinal cord. Each vertebral plate is composed of two layers, a somatic and a splanchnic, and encloses the already-mentioned section of the body-cavity. The cells of both layers of the plate are columnar, and each consists of a very large nucleus, invested by a delicate layer of protoplasm.

Before the end of stage H the inner or splanchnic wall of the protovertebra loses its simple constitution, owing to the middle part of it, opposite the dorsal two-thirds of the notochord, undergoing peculiar changes. These changes are indicated in transverse sections (Pl. 11, figs. 5 and 6, mp´), by the cells in the part we are speaking of acquiring a peculiar angular appearance, and becoming one or two deep; and the meaning of the changes is at once shewn by longitudinal horizontal sections. These prove (Pl. 12, fig. 10) that the cells in this situation have become elongated in a longitudinal direction, and, in fact, form typical spindle-shaped embryonic muscle-cells, each with a large nucleus. Every muscle-cell extends for the whole length of a protovertebra, and in the present stage, or at any rate in stage I, acquires a peculiar granulation, which clearly foreshadows transverse striation (Pl. 12, figs. 11-13).

Thus by stage H a small portion of the splanchnopleure which forms the inner layer of each protovertebra, becomes differentiated into a distinct band of longitudinal striated muscles; these almost at once become functional, and produce the peculiar serpentine movements of the embryo, spoken of in a previous chapter, p. [291].

It may be well to say at once that these muscles form but a very small part of the muscles which eventually appear; which latter are developed at a very much later period from the remaining cells of the protovertebræ. The band developed at this stage appears to be a special formation, which has arisen through the action of natural selection, to enable the embryo to meet its respiratory requirements, by continually moving about, and so subjecting its body to fresh oxydizing influences; and as such affords an interesting example of an important structure acquired during and for embryonic life.

Though the cavities in the protovertebræ are at first perfectly continuous with the general body-cavity, of which indeed they merely form a specialized part, yet by the close of stage H they begin to be constricted off from the general body-cavity, and this process is continued rapidly, and completed shortly after stage I, and considerably before the commencement of stage K (Pl. 11, figs. 6 and 8). While this is taking place, part of the splanchnic layer of each protovertebra, immediately below the muscle-band just described, begins to proliferate, and produce a number of cells, which at once grow in between the muscles and the notochord. These cells are very easily seen both in transverse and longitudinal sections, and form the commencing vertebral bodies (Pl. 11, fig. 6, and Pl. 12, figs. 10 and 11, Vr).