Fig. 197. Longitudinal section through an advanced embryo of Bombinator.
m. mouth; an. anus; l. liver; ne. neurenteric canal; mc. medullary canal; ch. notochord; pn. pineal gland.
If the above position be admitted, it is not permissible to shirk the conclusions which seem necessarily to follow, however great the difficulties may be which are involved in their acceptance. These conclusions have in part already been dealt with by Dohrn in his suggestive tract (No. [250]). In the first place the alimentary canal must primitively have been continued to the end of the tail; and if so, it is hardly credible that the existing anus can have been the original one. Although, therefore, it is far from easy, on the physiological principles involved in the Darwinian theory, to understand the formation of a new anus[112]; it is nevertheless necessary to believe that the present vertebrate anus is a formation acquired within the group of the Chordata, and not inherited from some older group. This involves a series of further consequences. The opening of the urinogenital ducts into the cloaca must also be secondary, and it is probable that the segmental tubes were primitively continued along the whole postanal region of the vertebrate tail, opening into the body cavity which embryology proves to have been originally present there. They are in fact continued in many existing forms for some distance behind the present anus. If the present anus is secondary, there must have been a primitive anus, which was probably situated behind the postanal vesicle; and therefore in the region of the neurenteric canal. The neurenteric canal is, however, the remnant of the blastopore (vide p. [277]). It follows, therefore, that the vertebrate blastopore is probably almost, if not exactly identical in position with the primitive anus. This consideration may assist in explaining the remarkable phenomenon of the existence of the neurenteric canal. The attempt has already been made to shew that the central canal of the nervous system is really a groove converted into a tube and lined by the external epidermis. This tube (as may be concluded from embryological considerations) was probably at first open posteriorly, and no doubt terminated at the primitive anus. On the closure of the primitive anal opening, the terminal portions of the postanal gut and the neural tube, may conceivably have been so placed that both of them opened into a common cavity, which previously had communication with the exterior by the anus. Such an arrangement would necessarily result in the formation of a neurenteric canal. It seems not impossible that a dilated vesicle, often present at the end of the postanal gut (vide [fig. 28*], p. [58]), may have been the common cavity into which both neural and alimentary tubes opened[113]. Till further light is thrown by fresh discoveries upon the primitive condition of the posterior continuation of the vertebrate alimentary tract, it is perhaps fruitless to attempt to work out more in detail the above speculation.
Body cavity and mesoblastic somites. The Chordata, or at least the most primitive existing members of the group, are characterized by the fact that the body cavity arises as a pair of outgrowths of the archenteric cavity. This feature[114] in the development is a nearly certain indication that the Chordata are a very primitive stock. The most remarkable point with reference to the development of the two outgrowths is, however, the fact that the dorsal part of each outgrowth becomes separated from the ventral. Its walls become segmented and form the mesoblastic somites, which eventually, on the obliteration of their cavity, give rise to the muscle-plates and to the tissue surrounding the notochord. It is not easy to understand the full significance of the processes concerned in the formation of the mesoblastic somites (vide p. [296]). The mesoblastic somites have no doubt a striking resemblance to the mesoblastic somites of the Chætopods, and most probably the segmentation of the mesoblast in the two groups is a phenomenon of the same nature; but the difference in origin between the two types of mesoblastic somites is so striking, and the development of the muscular system from them is so dissimilar in the two groups, as to render a direct descent of the Chordata from the Chætopoda very improbable. The ventral parts of the original outgrowth give rise to the permanent body cavity, which appears originally to have been divided into two parts by a dorsal and a ventral mesentery.
The notochord. The most characteristic organ of the Chordata is without doubt the notochord. The ontogenetic development of this organ probably indicates that it arose as a differentiation of the dorsal wall of the archenteron; at the same time it is not perhaps safe to lay too much stress upon its mode of development. Embryological and anatomical evidence demonstrate, however, in the clearest manner that the early Chordata were provided with this organ as their sole axial skeleton; and no invertebrate group can fairly be regarded as genetically related to the Chordata till it can be shewn to possess some organ either derived from a notochord, or capable of having become developed into a notochord. No such organ has as yet been recognized in any invertebrate group[115].
Gill-clefts. The gill-clefts, which are essentially pouches of the throat opening externally, constitute extremely characteristic organs of the Chordata, and have always been taken into consideration in any comparison between the Chordata and the Invertebrata.
Amongst the Invertebrata organs of undoubtedly the same nature are, so far as I know, only found in Balanoglossus, where they were discovered by Kowalevsky. The resemblance in this case is very striking; but although it is quite possible that the gill-clefts in Balanoglossus are genetically connected with those of the Chordata, yet the organization of Balanoglossus is as a whole so different from that of the Chordata that no comparison can be instituted between the two groups in the present state of our knowledge.
Other organs of the Invertebrata have some resemblance to the gill-clefts. The lateral pits of the Nemertines, which appear to grow out as a pair of œsophageal diverticula, which are eventually placed in communication with the exterior by a pair of ciliated canals (vide Vol. II. pp. 200 and 202), are such organs.
Semper (No. [256]) has made the interesting discovery that in the budding of Nais and Chætogaster two lateral masses of cells, in each of which a lumen may be formed, unite with the oral invagination and primitive alimentary canal to form the permanent cephalic gut. The lateral masses of cells are regarded by him as branchial passages homologous in some way with those in the Chordata. The somewhat scanty observations on this subject which he has recorded do not appear to me to lend much support to this interpretation.
It is probable that the part of the alimentary tract in which gill-clefts are present was originally a simple unperforated tube provided with highly vascular walls; and that respiration was carried on in it by the alternate introduction and expulsion of sea water. A more or less similar mode of respiration has been recently shewn by Eisig[116] to take place in the fore part of the alimentary tract of many Chætopods. This part of the alimentary tract was probably provided with paired cæcal pouches with their blind ends in contiguity with the skin.
Perforations placing these pouches in communication with the exterior must be supposed to have been formed; and the existence of openings into the alimentary tract at the end of the tentacles of many Actiniæ and of the hepatic diverticula of some nudibranchiate Molluscs (Eolis, &c.[117]) shews that such perforations may easily be made. On the formation of such perforations the water taken in at the mouth would pass out by them; and the respiration would be localized in the walls of the pouches leading to them, and thus the typical mode of respiration of the Chordata would be established.