Figs. 89 and 90—Transverse section of cœlomula embryos of triton. (From Hertwig.) Fig. 89, section through the primitive mouth. Fig. 90, section in front of the primitive mouth, u primitive mouth. dh gut-cavity, dz yelk-cells, dp yelk-stopper, ak outer and ik inner germinal layer, pb parietal and vb visceral middle layer, m medullary plate, ch chorda.
Fig. 91. A, B, C.—Vertical section of the dorsal part of three triton-embryos. (From Hertwig.) In Fig. A the medullary swellings (the parallel borders of the medullary plate) begin to rise; in Fig. B they grow towards each other; in Fig. C they join and form the medullary tube. mp medullary plate, mf medullary folds, n nerve-tube, ch chorda, lh body-cavity, mk1 and mk2 parietal and visceral mesoblasts, uv primitive-segment cavities, ak ectoderm, ik entoderm, dz yelk-cells, dh gut-cavity.
From the evolutionary point of view the cœlom-pouches are, in any case, older than the chorda; since they also develop in the same way as in the chordonia in a number of invertebrates which have no chorda (for instance, Sagitta, Figs. 76–78). Moreover, in the amphioxus the first outline of the chorda appears later than that of the cœlom-sacs. Hence we must, according to the biogenetic law, postulate a special intermediate form between the gastrula and the chordula, which we will call cœlomula, an unarticulated, worm-like body with primitive gut, primitive mouth, and a double body-cavity, but no chorda. This embryonic form, the bilateral cœlomula (Fig. 81), may in turn be regarded as the ontogenetic reproduction (maintained by heredity) of an ancient ancestral form of the cœlomaria, the Cœlomæa (cf. Chapter XX).
In Sagitta and other worm-like animals the two cœlom-pouches (presumably gonads or sex-glands) are separated by a complete median partition, the dorsal and ventral mesentery (Fig. 78 dm, vm); but in the vertebrates only the upper part of this vertical partition is maintained, and forms the dorsal mesentery. This mesentery afterwards takes the form of a thin membrane, which fastens the visceral tube to the chorda (or the vertebral column). At the under side of the visceral tube the cœlom-sacs blend together, their inner or median walls breaking down and disappearing. The body-cavity then forms a single simple hollow, in which the gut is quite free, or only attached to the dorsal wall by means of the mesentery.
The development of the body-cavity and the formation of the chordula in the higher vertebrates is, like that of the gastrula, chiefly modified by the pressure of the food-yelk on the embryonic structures, which forces its hinder part into a discoid expansion. These cenogenetic modifications seem to be so great that until twenty years ago these important processes were totally misunderstood. It was generally believed that the body-cavity in man and the higher vertebrates was due to the division of a simple middle layer, and that the latter arose by cleavage from one or both of the primary germinal layers. The truth was brought to light at last by the comparative embryological research of the Hertwigs. They showed in their Cœlom Theory (1881) that all vertebrates are true enterocœla, and that in every case a pair of cœlom-pouches are developed from the primitive gut by folding. The cenogenetic chordula-forms of the craniotes must therefore be derived from the palingenetic embryology of the amphioxus in the same way as I had previously proved for their gastrula-forms.
The chief difference between the cœlomation of the acrania (amphioxus) and the other vertebrates (with skulls—craniotes) is that the two cœlom-folds of the primitive gut in the former are from the first hollow vesicles, filled with fluid, but in the latter are empty pouches, the layers of which (inner and outer) close with each other. In common parlance we still call a pouch or pocket by that name, whether it is full or empty. It is different in ontogeny; in some of our embryological literature ordinary logic does not count for very much. In many of the manuals and large treatises on this science it is proved that vesicles, pouches, or sacs deserve that name only when they are inflated and filled with a clear fluid. When they are not so filled (for instance, when the primitive gut of the gastrula is filled with yelk, or when the walls of the empty cœlom-pouches are pressed together), these vesicles must not be cavities any longer, but “solid structures.”