Reproductive System

The sexes are separate, and when mature are sometimes distinguished by small differences of colour in the genital region. Both male and female gonads consist of more or less lobulated hollow sacs connected with the epidermis by short ducts. In their disposition they are either uniserial, biserial or multiserial. They occur in the branchial region, and also extend to a variable distance behind it. In exceptional cases they are either confined to the branchial region or excluded from it. When they are arranged in uniserial or biserial rows the genital ducts open into or near the branchial grooves in the region of the pharynx and in a corresponding position in the post-branchial region. An important feature is the occurrence in some species (Ptychoderidae) of paired longitudinal pleural or lateral folds of the body which are mobile, and can be approximated at their free edges so as to close in the dorsal surface, embracing both the median dorsal nerve-tract and the branchial grooves with the gill-pores, so as to form a temporary peri-branchial and medullary tube, open behind where the folds cease.

On the other hand, they can be spread out horizontally so as to expose their own upper side as well as the dorsal surface of the body (fig. 1). These folds are called the genital pleurae because they contain the bulk of the gonads. Correlated with the presence of the genital pleurae there is a pair of vascular folds of the basement membrane proceeding from the dorsal wall of the gut in the post-branchial portion of the branchio-genital region, and from the dorsal angles made by the pleural folds with the body-wall in the pharyngeal region; they pass, in their most fully developed condition, to the free border of the genital pleurae. These vascular membranes are called the lateral septa. Since there are many species which do not possess these genital pleurae, the question arises as to whether their presence or their absence is the more primitive condition. Without attempting to answer this question categorically, it may be pointed out that within the limits of the family (Ptychoderidae) which is especially characterized by their presence there are some species in which the genital pleurae are quite obsolete, and yet lateral septa occur (e.g. Ptychodera ruficollis), seeming to indicate that the pleural folds have in such cases been secondarily suppressed.


The development of Balanoglossus takes place according to two different schemes, known as direct and indirect, correlated with the occurrence in the group of two kinds of ova, large and small. Direct development, in which the adult form is achieved without striking metamorphosis by a gradual succession of stages, seems to be confined to the family Balanoglossidae. The remaining two families of Enteropneusta, Ptychoderidae and Spengelidae, contain species of which probably all pursue an indirect course of development, culminating in a metamorphosis by which the adult form is attained. In these cases the larva, called Tornaria, is pelagic and transparent, and possesses a complicated ciliated seam, the longitudinal ciliated band, often drawn out into convoluted bays and lappets. In addition to this ciliated band the form of the Tornaria is quite characteristic and unlike the adult. The Tornaria larva offers a certain similarity to larvae of Echinoderms (sea-urchins, star-fishes, and sea-cucumbers), and when first discovered was so described.

It is within the bounds of possibility that Tornaria actually does indicate a remote affinity on the part of the Enteropneusta to the Echinoderms, not only on account of its external form, but also by reason of the possession of a dorsal water-pore communicating with the anterior body-cavity. In the direct development Bateson showed that the three divisions of the coelom arise as pouches constricted off from the archenteron or primitive gut, thus resembling the development of the mesoblastic somites of Amphioxus. It would appear that while the direct development throws light upon the special plan of organization of the Enteropneusta, the indirect development affords a clue to their possible derivation. However this may be, it is sufficiently remarkable that a small and circumscribed group like the Enteropneusta, which presents such a comparatively uniform plan of composition and of external form, should follow two such diverse methods of development.