The works of Francis Maitland Balfour, Volume 2 (of 4)

by Francis M. Balfour

The formation of the alimentary cavity commences shortly after the complete investiture of the embryo by the epiblast cells. At the oral pole an invagination of epiblast cells takes place ([fig. 83] B), which makes its way towards the opposite pole. More especially from the figures given by Agassiz, and from the explanation of his plates, it would seem that a large chamber is formed in the hypoblast at the end of the invaginated tube, into which this tube soon opens ([fig. 83] C). The invaginated tube would seem to give rise to the so-called stomach, while the chamber at its aboral extremity is no doubt the infundibulum, which as may be gathered from Kowalevsky’s statements, is lined by a flattened epithelium. At a later period the gastrovascular canals grow out from the infundibulum as four pouches, which are surrounded by, and grow at the expense of, the large central cells, which have in the meantime arranged themselves in four masses, and appear to serve as a kind of yolk. The nuclei of these large cells according to Kowalevsky disappear, and the cells themselves break up into continually smaller masses.

The main difficulty in the above description of Agassiz is the origin of the infundibulum. In the absence of definite statements on this head it seems reasonable to conclude that it arises as a space hollowed out in the central cells, and that its walls are formed of elements derived from the yolk cells[84]. On this interpretation the alimentary canal of the Ctenophora would consist, as in the Acraspedote Medusæ and Actinozoa, of two sections: (1) A true hypoblastic section consisting of the infundibulum and the gastrovascular canals derived from it; and (2) an epiblastic section—the stomodæum—forming the stomach.

The observations of Kowalevsky on the alimentary system do not wholly tally with those of Agassiz. He finds that the oral side of the embryo becomes hollowed out, and that the hollow, lined by flattened cells, becomes constricted off as the infundibulum, from which the radial canals subsequently grow out. To the infundibulum there leads a narrow canal lined by a columnar epithelium which becomes the gastric cavity.

While the alimentary canal is becoming formed a series of important changes takes place in other parts of the embryo. The rows of locomotive paddles first appear as four longitudinal equidistant linear thickenings of the epiblast near the aboral pole ([fig. 83] D). On the projecting surface of these ridges stiff cilia appear which coalesce together to form the paddles. While the embryo is still within the egg the rows of paddles are quite short and also double. There are in Pleurobrachia about eight or nine pairs of paddles in each row. Each double row eventually separates into two.

In all the forms except the Eurostomata (Beroe) two tentacles grow out as thickenings of the epiblast ([fig. 84] B, t.). They are placed at the opposite poles of the long transverse axis of the embryo.

A process of the contractile gelatinous tissue of the body, the origin of which is described below, makes its way, according to Kowalevsky, into the tentacles.

The central apparatus of the nervous system and the otoliths are formed at the aboral pole from a thickening of the epiblast, but the full details of their formation have not been elucidated. It may be well to preface my account of their development with a short statement of their adult structure.

They consist in the adult of a vesicle with a ciliated lining situated at the bifurcation of the two anal tubes, and of certain structures connected with this vesicle. From the floor of the vesicle is suspended a mass of otoliths by four leaf-like bodies known as suspenders. The roof is very delicate and has the form of a four-sided pyramid. Six openings lead into the vesicle. Through four of these, placed at the four corners, there pass out four ciliated grooves continuous with the suspenders. These grooves, after leaving the otolithic vesicle, bifurcate and pass to the eight rows of paddles. At the two sides the walls of the vesicle are continuous with two thickened ciliated plates with swollen edges, opposite the centres of which are two lateral openings into the vesicle, completing the six openings. Through the lateral openings the sea-water is driven by the action of the cilia of the plates.

The development of these parts is as follows—In the aboral thickening of epiblast a cavity makes its appearance, the walls of which constitute the rudiment of the otolithic vesicle ([fig. 83] B and C, s.c.). The roof of the cavity is extremely delicate. On each side of it a thickening of cells becomes established, regarded by Kowalevsky as the rudiment of the nervous ganglia. These thickenings appear to give origin to the lateral ciliated plates. The otoliths arise from cells at four separate points at the corners of the ciliated plates opposite the rows of paddles ([fig. 84] A, ot.).