The evidence at our disposal appears to me to indicate that the third nerve belongs to the cranio-spinal series of segmental nerves, while the optic and olfactory nerves appear to me equally clearly not to belong to this series[104]. The mid-brain, as giving origin to the third nerve, would appear not to have been part of the ganglion of the præoral lobe.
These considerations indicate with fair probability that the part of the head containing the fore-brain is the equivalent of the præoral lobe of many Invertebrate forms; and the primitive position of the Vertebrate mouth on the ventral side of the head affords a distinct support for this view. It must however be admitted that this part of the head is not sharply separated in development from that behind; and, though the fore-brain is usually differentiated very early as a distinct lobe of the primitive nervous tube, yet that such differentiation is hardly more marked than in the other parts of the brain. The termination of the notochord immediately behind the fore-brain is, however, an argument in favour of the morphological distinctness of the latter structure.
The evidence at our disposal appears to indicate that the posterior part of the head was not differentiated from the trunk in lower Chordata; but that, as the Chordata rose in the scale of development, more and more centralizing work became thrown on the anterior part of the nervous cord, and pari passu this part became differentiated into the mid- and hind-brain. An analogy for such a differentiation is supplied in the compound subœsophageal ganglion of many Arthropods; and, as will be shewn in the chapter on the nervous system, there is strong embryological evidence that the mid- and hind-brains had primitively the same structure as the spinal cord. The head appears however to have suffered in the course of its differentiation a great concentration in its posterior part, which becomes progressively more marked, even within the limits of the surviving Vertebrata. This concentration is especially shewn in the structure of the vagus nerve, which, as first pointed out by Gegenbaur, bears evidence of having been originally composed of a great series of nerves, each supplying a visceral cleft. Rudiments of the posterior nerves still remain as the branches to the œsophagus and stomach[105].
The atrophy of the posterior visceral clefts seems to have taken place simultaneously with the concentration of the neural part of the head; but the former process did not proceed so rapidly as the latter, so that the visceral region of the head is longer in the lower Vertebrata than the neural region, and is dorsally overlapped by the anterior part of the spinal cord and the anterior muscle-plates (vide[fig. 47]).
On the above view the posterior part of the head must have been originally composed of a series of somites like those of the trunk, but in existing Vertebrata all trace of these, except in so far as they are indicated by the visceral clefts, has vanished in the adult. The cranial nerves however, especially in the embryo, still indicate the number of anterior somites; and an embryonic segmentation of the mesoblast has also been found in many lower forms in the region of the head, giving rise to a series of cavities known as head-cavities, enclosed by mesoblastic walls which afterwards break up into muscles. These cavities correspond with the nerves, and it appears that there is a præmandibular cavity corresponding with the third nerve ([fig. 193], 1pp) and a mandibular cavity (2pp) and a cavity in each of the succeeding visceral arches. The fifth nerve, the seventh nerve, the glossopharyngeal nerve, and the successive elements of the vagus nerve correspond with the posterior head-cavities.
Fig. 193. Transverse section through the front part of the head of a young Pristiurus embryo.
The section, owing to the cranial flexure, cuts both the fore- and the hind-brain. It shews the præmandibular and mandibular head-cavities 1pp and 2pp, etc.
fb. fore-brain; l. lens of eye; m. mouth; pt. upper end of mouth, forming pituitary involution; 1ao. mandibular aortic arch; 1pp. and 2pp. first and second head-cavities; 1vc. first visceral cleft; V. fifth nerve; aun. ganglion of auditory nerve; VII. seventh nerve; aa. dorsal aorta; acv. anterior cardinal vein; ch. notochord.
The medullary canal. The general history of the medullary plate seems to point to the conclusion that the central canal of the nervous system has been formed by a groove having appeared in the ancestor of the Chordata along the median dorsal line, which caused the sides of the nervous plate, which was placed immediately below the skin, or may perhaps at that stage not have been distinctly differentiated from the skin, to be bent upwards; and that this groove subsequently became converted into a canal. This view is not only supported by the actual development of the central canal of the nervous system (the types of Teleostei, Lepidosteus and Petromyzon being undoubtedly secondary), but also (1) by the presence of cilia in the epithelium lining the canal, probably inherited from cilia coating the external skin, and (2) by the posterior roots arising from the extreme dorsal line ([fig. 194]), a position which can most easily be explained on the supposition that the two sides of the plate, from which the nerves originally proceeded have been folded up so as to meet each other in the median dorsal line[106].
The medullary plate, before becoming folded to form the medullary groove, is (except in Amphibia) without any indication of being composed of two halves. In both the embryo and adult the walls of the tube have however a structure which points to their having arisen from the coalescence of two lateral, and most probably at one time independent, cords; and as already indicated this is the view I am myself inclined to adopt; vide pp. [303] and [304].