Such are the chief structural features presented by these nuclei, which are present during the whole of the earlier periods of development and retain throughout the same appearance. There can be little doubt that their knobbed condition implies that they are undergoing a rapid division. The arguments for this view I have already insisted on, and, in spite of the observations of Dr Kleinenberg shewing that similar nuclei of Nephelis do not undergo division, the case for their doing so in the Elasmobranch eggs is to my mind a very strong one.
During this stage the distribution of these nuclei in the yolk becomes somewhat altered from that in the earlier stages. Although the nuclei are still scattered generally throughout the finer yolk-matter around the blastoderm, yet they are especially aggregated at one or two points. In the first place a special collection of them may be noticed immediately below the floor of the segmentation cavity. They here form a distinct row or even layer. If the presence of this layer is coupled with the fact that at this period cells are beginning to appear on the floor of the segmentation cavity, a strong argument is obtained for the supposition that around these nuclei cells are being produced, which pass into the blastoderm to form the floor. Of the actual formation of cells at this period I have not been able to obtain any satisfactory example, so that it remains a matter of deduction rather than of direct observation.
Another special aggregation of nuclei is generally present at the periphery of the blastoderm, and the same amount of doubt hangs over the fate of these as over that of the previously mentioned nuclei.
The next stage is the most important in the whole history of the formation of the layers. Not only does it serve to shew, that the process by which the layers are formed in Elasmobranchii can easily be derived from a simple gastrula type like that of Amphioxus, but it also serves as the key by which other meroblastic types of development may be explained. At the very commencement of this stage the embryonic swelling becomes more conspicuously visible than it was. It now projects above the level of the yolk in the form of a rim. At one point, which eventually forms the termination of the axis of the embryo, this projection is at its greatest; while on either side of this it gradually diminishes and finally vanishes. This projection I propose calling, as in my preliminary paper[129], the embryonic rim.
The segmentation cavity can still be seen from the surface, and a marked increase in the size of the blastoderm may be noticed. During the stage last described, the growth was but very slight; hence the rather sudden and rapid growth which now takes place becomes striking.
Longitudinal sections at this stage, as at the earlier stages, are the most instructive. Such a section on the same scale as Pl. 7, fig. 4, is represented in Pl. 7, fig. 5. It passes parallel to the long axis of the embryo, through the point of greatest development of the embryonic ring.
The three fresh features of the most striking kind are (1) the complete envelopment of the segmentation cavity within the lower layer cells, (2) the formation of the embryonic rim, (3) the increase in distance between the posterior end of the blastoderm and the segmentation cavity. The segmentation cavity has by no means relatively increased in size. The roof has precisely its earlier constitution, being composed of an internal lining of lower layer cells and an external one of epiblast. The thin lining of lower layer cells is, in the course of mounting the sections, very apt to fall off; but I am absolutely satisfied that it is never absent.
The floor of the cavity has undergone an important change, being now formed by a layer of cells instead of by the yolk. A precisely similar but more partial change in the constitution of the floor takes place in Osseous Fishes[130].
The mode in which the floor is formed is a question of some importance. The nuclei, which during the last stage formed a row beneath it, probably, as previously pointed out, take some share in its formation. An additional argument to those already brought forward in favour of this view may be derived from the fact that during this stage such a row of nuclei is no longer present.
This argument may be stated as follows: