Such are the main features presented by the mesoblast in Elasmobranchii, which favour the view of its having originally formed the walls of the alimentary diverticula. Against this view of its nature are the facts (1) of the mesoblast plates being at first solid, and (2), as a consequence of this, of the body-cavity never communicating with the alimentary canal. These points, in view of our knowledge of embryological modifications, cannot be regarded as great difficulties to my view. We have many examples of organs, which, though in most cases arising as involutions, yet appear in other cases as solid ingrowths. Such examples are afforded by the optic vesicle, auditory vesicle, and probably also by the central nervous system, of Osseous Fish. In most Vertebrates these organs are formed as hollow involutions from the exterior; in Osseous Fish, however, as solid involutions, in which a cavity secondarily appears.
The segmental duct of Elasmobranchii or the Wolffian duct (segmental duct) of Birds are cases of a similar kind, being organs which must originally have been formed as hollow involutions, but which now arise as solid bodies.
Only one more instance of this kind need be cited, taken from the Echinoderms.
The body-cavity and the mesoblast investing it arise in the case of most Echinoderms as hollow involutions of the alimentary tract, but in some exceptional groups, Ophiura and Amphiura, are stated to be solid at first and only subsequently to become hollow. Should the accuracy of Metschnikoff's account of this point be confirmed, an almost exact parallel to what has been supposed by me to have occurred with the mesoblast in Elasmobranchii, and other groups, will be supplied.
The tendency of our present knowledge appears to be in favour of regarding the body-cavity in Vertebrates as having been primitively the cavity of alimentary diverticula, and the mesoblast as having formed the walls of the diverticula.
This view, to say the least of it, suits the facts which we know far better than any other theory which has been proposed, and though no doubt the à priori difficulties in its way are very great, yet it appears to me to be sufficiently strongly supported to deserve the attention of investigators. In the meantime, however, our knowledge of invertebrate embryology is so new and imperfect that no certainty on a question like that which has just been discussed can be obtained; and any generalizations made at present are not unlikely to be upset by the discovery of fresh facts.
The only other point in connection with the mesoblast which I would call attention to is the formation of the vertebral bodies.
My observations confirm those of Remak and Gegenbaur, shewing that there is a primary segmentation of the vertebral bodies corresponding to that of the muscle-plates, followed by a secondary segmentation in which the central lines of the vertebral bodies are opposite the partitions between the muscle-plates.
The explanation of these changes is not difficult to find. The primary segmentation of the body is that of the muscle-plates, which must have been present at a time when the vertebral bodies had no existence. As soon however as the notochordal sheath was required to be strong as well as flexible, it necessarily became divided into a series of segments.
The conditions under which the lateral muscles can cause the flexure of the vertebral column are clearly that each muscle-segment shall be capable of acting on two vertebræ; and this condition can only be fulfilled when the muscle-segments are opposite the intervals between the vertebræ. Owing to this necessity, when the vertebral segments became formed, their centres corresponded, not with the centres of the muscle-plates, but with the inter-muscular septa.