According to the statements of Bambeke and Götte, the Amphibians present rather remarkable peculiarities in the development of their muscular system. Each side-plate of mesoblast is divided into a somatic and a splanchnic layer, continuous throughout the vertebral and parietal portions of the plate. The vertebral portions (protovertebræ) of the plates soon become separated from the parietal, and form an independent mass of cells constituted of two layers, which were originally continuous with the somatic and splanchnic layers of the parietal plates. The outer or somatic layer of the vertebral plates is formed of a single row of cells, but the inner or splanchnic layer is made up of a central kernel of cells and an inner single layer. This central kernel is the first portion of the vertebral body to undergo any change, and it becomes converted into the main dorso-lateral muscles of the body, which apparently correspond with the muscles derived from the whole muscle-plate of the Elasmobranchii. From the inner layer of the splanchnic division there are next formed the main internal ventral muscles, rectus abdominis, &c., as well as the chief connective-tissue elements of the parts surrounding the spinal cord. The outer layer of the vertebral plates forms the dermis and subcutaneous connective tissue, as well as some of the superficial muscles of the trunk and the muscles of the limbs.
Dr Götte appears to think that the vertebral plates in Amphibians present a perfectly normal development very similar to that of other Vertebrates. The divergences between Amphibians and other Vertebrates appear, however, to myself, to be very great, and although the very careful account given by Dr Götte is probably to be relied on, yet some further explanation than he has offered of the development of these parts amongst the Amphibians would seem to be required.
A primary stage in which the two layers of the vertebral plates are continuous with the somatic and splanchnic layers of the body-wall is equally characteristic of Amphibians, Elasmobranchii and Mammals. In the subsequent development, however, a great difference between the types becomes apparent, for whereas in Elasmobranchii both layers of the vertebral plates combine to form the muscle-plates, out of which the great dorso-lateral muscles are formed, in Amphibians what appear to be the equivalent muscles are derived from a few of the cells (the kernel) of the inner layer of the vertebral plates only. The cells which form the lateral muscles in Amphibians might be thought to correspond in position with the cells which become, in Elasmobranchii, converted into the special early formed band of muscles (m.p´.), rather than, as their development seems to indicate, with the whole Elasmobranch muscle-plates[209].
Osseous Fishes are stated to agree with Amphibians in the development of their protovertebræ and muscular system[210], but further observations on this point are required.
Though the development of the general muscular system and muscle-plates does not, according to existing statements, take place on quite the same type throughout the Vertebrate subkingdom, yet the comparison which has been instituted between Elasmobranchii and other Vertebrates appears to prove that there are one or two common features in their development, which may be regarded as primitive, and as having been inherited from the ancestors of Vertebrates. These features are (1) The extension of the body-cavity into the vertebral plates, and subsequent enclosure of this cavity between the two layers of the muscle-plates; (2) The primitive division of the vertebral plate into a somatic and a splanchnic layer, and the formation of a large part of the voluntary muscular system out of the splanchnic layer.
* * * * *
The ultimate derivation of the mesoblast forms one of the numerous burning questions of modern embryology, and there are advocates to be found for almost every one of the possible views the question admits of.
All who accept the doctrine of descent are agreed that primitively only two embryonic layers were present—the epiblast and the hypoblast—and that the mesoblast subsequently appeared as a distinct layer, after a certain complexity of organization had been attained.
The general agreement stops, however, at this point, and the greatest divergence of opinion exists with reference to all further questions which bear on the development of the mesoblast. There appear to be four possibilities as to the origin of this layer.
It may be derived: