Not only is the coelom thus subdivided, but the enteron (gut, alimentary canal, digestive tube) itself shows indications of three main subsections in continuity with one another: - (1) proboscis-gut (Eicheldarm, stomochord, vide infra); (2) collar-gut (buccal cavity, throat); (3) truncal gut extending from the collar to the vent.
The proboscis-gut occurs as an outgrowth from the anterior dorsal wall of the collar-gut, and extends forward into the basal (posterior) region of the proboscis, through the neck into the proboscis-coelom, ending blindly in front. Although an integral portion of the gut, it has ceased to assist in alimentation, its epithelium undergoes vacuolar differentiation and hypertrophy, and its lumen becomes more or less vestigial. It has, in fact, become metamorphosed into a resistant supporting structure resembling in some respects the notochord of the true Chordata, but probably not directly comparable with the latter structure, being related to it solely by way of substitution. On account of the presence and mode of origin (from the gut-wall) of this organ Bateson introduced the term hemichorda as a phyletic name for the class Enteropneusta. As the proboscis-gut appears to have undoubtedly skeletal properties, and as it also has topographical relations with the mouth, it has been designated in English by the non-committal term stomochord.
It is not a simple diverticulum of the collar-gut, but a complex structure possessing paired lateral pouches and a ventral convexity (ventral caecum) which rests in a concavity at the front end of the body of the nuchal skeleton (fig. 3). In some species (Spengelidae) there is a long capillary vermiform extension of the stomochord in front. The nuchal skeleton is a non-cellular laminated thickening of basement-membrane underlying that portion of the stomochord which lies between the above-mentioned pouches and the orifice into the throat. At the point where the stomochord opens into the buccal cavity the nuchal skeleton bifurcates, and the two cornua thus produced pass obliquely backwards and downwards embedded in the wall of the throat, often giving rise to projecting ridges that bound a dorsal groove of the collar-gut which is in continuity with the wall of the stomochord (fig. 3).
At the base of the epidermis (which is in general ciliated) there is over the entire surface of the body a layer of nerve-fibres, occurring immediately outside the basement-membrane which separates the epidermis from the subjacent musculature. The nervous system is thus essentially epidermal in position and diffuse in distribution; but an interesting concentration of nerve-cells and fibres has taken place in the collar-region, where a medullary tube, closed in from the outside, opens in front and behind by anterior and posterior neuropores. This is the collar nerve-tube. Sometimes the central canal is wide and uninterrupted between the two neuropores; in other cases it becomes broken up into a large number of small closed medullary cavities, and in others again it is obsolete. In one family, the Ptychoderidae, the medullary tube of the collar is connected at intermediate points with the epidermis by means of a variable number of unpaired outgrowths from its dorsal wall, generally containing an axial lumen derived from and in continuity with the central canal.
These hollow roots terminate blindly in the dorsal epidermis of the collar, and place the nervous layer of the latter in direct connexion with the fibres of the nerve-tube. The exact significance of these roots is a matter for speculation, but it seems possible that they are epiphysial structures remotely comparable with the epiphysial (pineal) complex of the craniate vertebrates. In accordance with this view there would be also some probability in favour of regarding the collar nerve-tube of the Enteropneusta as the equivalent of the cerebral vesicle only of Amphioxus and the Ascidian tadpole, and also of the primary fore-brain of vertebrates.
Special thickenings of the diffuse nervous layer of the epidermis occur in certain regions and along certain lines. In the neck of the proboscis the fibrous layer is greatly thickened, and other intensifications of this layer occur in the dorsal and ventral middle lines of the trunk extending to the posterior end of the body. The dorsal epidermal nerve-tract is continued in front into the ventral wall of the collar nerve-tube, and at the point of junction there is a circular commissural thickening following the posterior rim of the collar and affording a special connexion between the dorsal and ventral nerve-tracts. From the ventral surface of the collar nerve-tube numerous motor fibres may be seen passing to the subjacent musculature. These fibres are not aggregated into roots.
Fig. 2. - Structure of branchial region.
dn, dorsal nerve.
vn, ventral nerve.