[180] Vide Preliminary Account, etc. Q. Jl. Micros. Science, Oct. 1874, Pl. 14, 8a. [This Edition, No. V. Pl. 3, 8a.] This and the other section from the same embryo (stage F) may be referred to. I have not thought it worth while repeating them here.
[181] I underestimated the distinctness of this formation in my earlier paper, loc. cit., although I recognised the fact that the mesoblast cells became arranged in two distinct layers.
[182] Zeitschrift f. wiss. Zoologie, 1873.
[183] Nearly simultaneously with Chapter III. of the present monograph on the Development of Elasmobranchii, which dealt in a fairly complete manner with the genesis of cells outside the blastoderm, there appeared two important papers dealing with the same subject for Teleostei. One of these, by Professor Bambeke, “Embryologie des Poissons Osseux,” Mém. Cour. Acad. Belgique, 1875, which appeared some little time before my paper, and a second by Dr Klein, Quart. Jour. of Micr. Sci. April, 1876. In both of these papers a development of nuclei and of cells is described as occurring outside the blastoderm in a manner which accords fairly well with my own observations.
The conclusions of both these investigators differ however from my own. They regard the finely granular matter, in which the nuclei appear, as pertaining to the blastoderm, and morphologically quite distinct from the yolk. From their observations we can clearly recognise that the material in which the nuclei appear is far more sharply separated off from the yolk in Osseous Fish than in Elasmobranchii, and this sharp separation forms the main argument for the view of these authors. Dr Klein admits, however, that this granular matter (which he calls parablast) graduates into the typical food-yolk, though he explains this by supposing that the parablast takes up part of the yolk for the purpose of growth.
It is clear that the argument from a sharp separation of yolk and parablast cannot have much importance, when it is admitted (1) that in Osseous Fish there is a gradation between the two substances, while (2) in Elasmobranchii the one merges slowly and insensibly into the other.
The only other argument used by these authors is stated by Dr Klein in the following way. “The fact that the parablast has, at the outset, been forming one unit with what represents the archiblast, and, while increasing has spread i.e. grown over the yolk which underlies the segmentation-cavity, is, I think, the most absolute proof that the yolk is as much different from the parablast as it is from the archiblast.” This argument to me merely demonstrates that certain of the nutritive elements of the yolk become in the course of development converted into protoplasm, a phenomenon which must necessarily be supposed to take place on my own as well as on Dr Klein's view of the nature of the yolk. My own views on the subject have already been fully stated. I regard the so-called yolk as composed of a larger or smaller amount of food-material imbedded in protoplasm, and the meroblastic ovum as a body constituted of the same essential parts as a holoblastic ovum, though divided into regions which differ in the proportion of protoplasm they contain. I do not propose to repeat the positive arguments used by me in favour of this view, but content myself with alluding to the protoplasmic network found by Schultz and myself extending through the whole yolk, and to the similar network described by Bambeke as being present in the eggs of Osseous Fish after deposition but before impregnation. The existence of these networks is to me a conclusive proof of the correctness of my views. I admit that in Teleostei the 'parablast' contains more protoplasm than the homologous material in the Elasmobranch ovum, while it is probable that after impregnation the true yolk of Teleostei contains little or no protoplasm; but these facts do not appear to me to militate against my views.
I agree with Prof. Bambeke in regarding the cells derived from the sub-germinal matter as homologous with the so-called yolk-cells of the Amphibian embryo.
I have recently, in some of the later stages of development, met with very peculiar nuclei of the yolk immediately beneath the blastoderm at some little distance from the embryo, Pl. 10, fig. 8. They were situated not in finely sub-germinal matter, but amongst large yolk-spherules. They were very large, and presented still more peculiar forms than those already described by me, being produced into numerous long filiform processes. The processes from the various nuclei were sometimes united together, forming a regular network of nuclei quite unlike anything that I have previously seen described.
The sub-germinal matter, in which the nuclei are usually formed, becomes during the later stages of development far richer in protoplasm than during the earlier. It continually arises at fresh points, and often attains to considerable dimensions, no doubt by feeding on yolk-spherules. Its development appears to be determined by the necessities of growth in the blastoderm or embryo.