Fig. 194. Transverse section through the trunk of an embryo slightly older than fig. 28 e.
nc. neural canal; pr. posterior root of spinal nerve; x. subnotochordal rod; ao. aorta; sc. somatic mesoblast; sp. splanchnic mesoblast; mp. muscle-plate; mp´. portion of muscle-plate converted into muscle; Vv. portion of the vertebral plate which will give rise to the vertebral bodies; al. alimentary tract.
The origin and nature of the mouth. The most obvious point connected with the development of the mouth is the fact that in all vertebrate embryos it is placed ventrally, at some little distance from the front end of the body. This feature is retained in the adult stage in Elasmobranchii, the Myxinoids, and some Ganoids, but is lost in other vertebrate forms. A mouth, situated as is the embryonic vertebrate mouth, is very ill adapted for biting; and though it acquires in this position a distinctly biting character in the Elasmobranchii, yet it is almost certain that it had not such a character in the ancestral Chordata, and that its terminal position in higher types indicates a step in advance of the Elasmobranchii.
On the structure of the primitive mouth there appears to me to be some interesting embryological evidence, to which attention has already been called in the preceding chapters. In a large number of the larvæ or embryos of the lower Vertebrates the mouth has a more or less distinctly suctorial character, and is connected with suctorial organs which may be placed either in front of or behind it. The more important instances of this kind are (1) the Tadpoles of the Anura, with their posteriorly placed suctorial disc, (2) Lepidosteus larva ([fig. 195]) with its anteriorly placed suctorial disc, (3) the adhesive papillæ of the larvæ of the Tunicata. To these may be added the suctorial mouth of the Myxinoid fishes[107].
All these considerations point to the conclusion that in the ancestral Chordata the mouth had a more or less definitely suctorial character[108], and was placed on the ventral surface immediately behind the præoral lobe; and that this mouth has become in the higher types gradually modified for biting purposes, and has been carried to the front end of the head.
The mouth in Elasmobranchii and other Vertebrates is originally a wide somewhat rhomboidal cavity ([fig. 28] G); on the development of the mandibular and its maxillary (pterygo-quadrate) process the opening of the mouth becomes narrowed to a slit. The wide condition of the mouth may not improbably be interpreted as a remnant of the suctorial state. The fact that no more definite remnants of the suctorial mouth are found in so primitive a group as the Elasmobranchii is probably to be explained by the fact that the members of this group undergo an abbreviated development within the egg.
While the embryological data appear to me to point to the existence of a primitive suctorial mouth, very different conclusions have been put forward by other embryologists, more especially by Dohrn, which are sufficiently striking and suggestive to merit a further discussion.
As mentioned above, both Dohrn and Semper hold that the Vertebrata are descended from Chætopod-like forms, in which the ventral surface has become the dorsal. In consequence of this view Dohrn has arrived at the following conclusions: (1) that primitively the alimentary canal perforated the nervous system in the region of the original œsophageal nerve-ring; (2) that there was therefore an original dorsal mouth (the present ventral mouth of the Chætopoda); and (3) that the present mouth was secondary and derived from two visceral clefts which have ventrally coalesced.