(4) Part of the peritoneal epithelium.
From the splanchnic layer are formed
(1) A great part of the voluntary muscular system.
(2) Part of the intermuscular connective tissue (?).
(3) The axial skeleton.
(4) The muscular and connective-tissue wall of the alimentary tract.
(5) A great part of the peritoneal epithelium.
General Considerations. In the history which has just been given of the development of the mesoblast, there are several points which appear to me to throw light upon the primitive origin of that layer. Before entering into these it is however necessary to institute a comparison between the history of the mesoblast in Elasmobranchii and in other Vertebrates, in order to distinguish as far as possible the primitive and the secondary characters present in the various groups.
Though the Mammals are to be looked on as the most differentiated group amongst the Vertebrates, yet in their embryonic history they retain many very primitive features, and, as has been recently shewn by Hensen[204], present numerous remarkable approximations to the Elasmobranchii. We find accordingly[205] that the primitive lateral plates of mesoblast undergo nearly the same changes in these two groups. In Mammals there is at first a continuous cavity extending through both the parietal and vertebral portions of each plate, and dividing the plates into a somatic and a splanchnic layer: this cavity is the primitive body-cavity. The vertebral portion of each plate with its contained cavity then becomes divided off from the parietal. The later development of these parts is not accurately known, but it seems that the outer portion of each vertebral plate, composed of two layers (somatic and splanchnic) enclosing between them a remnant of the primitive body-cavity, becomes separated off as a muscle-plate. The remainder forms a vertebral rudiment, &c. Thus the extension of the body-cavity into the vertebral portion of the mesoblast, and the constriction of the vertebral portion of the cavity from the remainder, are as distinctive features of Mammals as they are of the Elasmobranchii.
In Birds[206] the horizontal splitting of the mesoblast into somatic and splanchnic layers appears, as in Mammals, to extend at first to the summit of the protovertebræ, but these bodies become so early separated from the parietal plates that this fact has usually been overlooked or denied; but even on the second day of incubation the outer layer of the protovertebræ is continuous with the somatic layer of the lateral plates, and the inner layer and kernel of the protovertebræ with the splanchnic layer of the lateral plates[207]. After the isolation of the protovertebræ the primitive position of the split which separated their somatic and splanchnic layers becomes obscured, but when on the third day the muscle-plates are formed they are found to be constituted of two layers, an inner and an outer, which enclose between them a central cavity. This remarkable fact, which has not received much attention, though noticeable in most figures, receives a simple explanation as a surviving rudiment on Darwinian principles. The central cavity of the muscle-plate is, in fact, a remnant of the vertebral extension of the body-cavity, and is the same cavity as that found in the muscle-plates of Elasmobranchii. The two layers of the muscle-plate also correspond with the two layers present in Elasmobranchii, the one belonging to the somatic, the other to the splanchnic layer of mesoblast. The remainder of the protovertebræ internal to the muscle-plates is very large in Birds, and is the equivalent of that portion of the protovertebræ which in Elasmobranchii is split off to form the vertebral bodies[208] (Pl. 11, figs. 6, 7, 8, Vr). Thus, though the history of the development of the mesoblast is not precisely the same for Birds as for Elasmobranchii, yet the differences between the two groups are of such a character as to prove in a striking manner that the Avian development is a derivation from a more primary form, like that of the Elasmobranchii.