The cranial flexure commences coincidently with the closing in of the neural canal in the region of the brain, and the division into fore, mid, and hind brain becomes visible at the same time as or even before the closing of the canal occurs. The embryo has now become more or less transparent, and protovertebræ, of which about twenty are present, can now be seen in the fresh specimens. The heart, however, is not yet formed.
Up to this period, a period at which the embryo becomes very similar in external appearance to any other vertebrate embryo, I have followed in my description a chronological order. I shall now cease to do so, since it would be too long for a preliminary notice of this kind, but shall confine myself to the history of a few organs whose development is either more important or more peculiar than that of the others.
The Protovertebræ.
I have thought it worth while to give a short history of the development of the protovertebræ, firstly, because it is very easy to follow this in the Dog-fish, and, secondly, because I believe that the Dog-fish have more nearly retained the primitive condition of the protovertebræ than any other vertebrate whose embryology has hitherto been described with sufficient detail.
I intend to describe, at the same time, the development of the spinal nerves.
I left each lateral mass of mesoblast in my last stage as a plate which had not yet become split into a somatic and a splanchnic sheet (Pl. 3, fig. 8a, vp), but which had become cut by transverse lines (not, indeed, extending to the outer limit of the sheet, but as yet not cut off by longitudinal lines of cleavage) into segments, which I called protovertebræ.
This sheet of mesoblast is fairly thick at its proximal (upper) end, but thins off laterally to a sheet two cells deep, and its cells are so arranged as to foreshadow its subsequent splitting into somatic and splanchnic sheets. Its upper (proximal) end is at this stage level with the bottom of the neural canal, but soon begins to grow upwards, and at the same time the splitting into somatopleure and splanchnopleure commences (Pl. 3, fig. 10, so and sp).
The separation between the two sheets is first visible in its uppermost part, and thence extends outwards. By this means each of the protovertebræ becomes divided into two sheets, which are only connected at their upper ends and outside the region of the body. I speak of the whole lateral sheet as being composed of protovertebræ, because at this time no separation into vertebral and lateral plates can be seen; but I may anticipate matters by saying that only the upper portion of the sheet from the level of the top of the digestive canal, becomes subsequently the true protovertebræ. From this it is clear that the pleuro-peritoneal cavity extends primitively quite up to the top of the protovertebræ; and that thus a portion of a sheet of mesoblast, at first perfectly continuous with the splanchnic sheet from which is derived the muscular wall of the alimentary canal, is converted into a part of the voluntary muscular system of the body, having no connection whatever with the involuntary muscular system of the digestive tract.
The pleuro-peritoneal cavity is first distinctly formed at a time when only two visceral clefts are present. Before the appearance of a third visceral cleft in a part of the innermost layer of each protovertebræ (which may be called the splanchnic layer, from its being continuous with the mesoblast of the splanchnopleure), opposite the bottom of the neural tube, some of the cells commence to become distinguishable from the rest, and to form a separate mass. This mass becomes much more distinct a little later, its cells being characterised by being spindle-shaped, and having an elongated nucleus which becomes deeply stained by reagents (Pl. 4, fig. 11, mp´). Coincidently with its appearance the young Dog-fish commences spontaneously to move rapidly from side to side with a kind of serpentine motion, so that, even if I had not traced the development of this differentiated mass of cells till it becomes a band of muscles close to the notochord, I should have had little doubt of its muscular nature. It is indicated in figs. 11, 12, 13, by the letters mp´. Its early appearance is most probably to be looked upon as an adaptation consequent upon the respiratory requirements of the young Dog-fish necessitating movements within the egg.
Shortly after this date, at a period when three visceral clefts are present, I have detected the first traces of the spinal nerves.