
Reproductive System
The structure of the genitalia in Murdochia pallidum may be outlined briefly. In the male, the vas deferens is a narrow tube closed throughout its whole length, as far as the tip of the prominent cephalic penis. It leaves the thin-walled testis sac as a lightly coiled, non-glandular tube, which emerges ventrally to the renal organ, to run the whole length of the pallial cavity, just below the rectum, along the attachment of the mantle to the trunk musculature. The closure of the pallial portion of the vas deferens—primitively an open ciliated groove—is to be regarded as correlated with the terrestrial habit. The wall of the pallial vas deferens is thickened and glandular, as far forward as the base of the penis, its lumen narrow, rounded or slightly flattened. This portion of the male duct evidently performs the role of a “prostate” gland: its lining cells are stoutly columnar, filled with minute, non-staining secretion granules, and having rounded, basal nuclei; Regularly alternating with the glandular cells are narrow, darkly compressed, ciliated cells, about half the length of the secreting cells, and with dark-staining elongate nuclei. Their distal surfaces are narrow, and cilia are borne also by the glandular cells, the whole of the lumen being uniformly lined. The penis is attached to the head immediately behind the right tentacle, forming a stout muscular column, flattened or cylindrical, with strong annular rugae, and yellowish-white in colour. It is generally reflected back within the pallial cavity. lying transversely across the trunk, or along the right side. It is perforated throughout by a narrow ejaculatory duct, continuous with the vas deferens, proceeding to the distal tip of the penis, which becomes narrow and cylindrical, and forms the intromittent part of the male organ. This structure is freely extensible, and can be withdrawn into the haemocoelic blood space enclosed by the stouter basal portion of the penis. (Fig. 3.) It can be strongly protruded by the pressure of the muscular wall of the basal part of the penis on the blood contained in the haemocoelic space. The terminal intromittent portion of the penis is supported by a flexible rod of elastic connective tissue, which runs alongside, and finally extends completely around the ejaculatory duct. It is made up of long, vacuolated cells, radially inserted at the base of the

epithelium of the ejaculatory duct. The basal part of the penis has a stout layer of muscle fibres, circular and longitudinal elements intermingled, and its external surface is traversed along one side by a broad tract of glandular epithelium, mucus being produced by rounded, subepithelial secretory cells, sending up narrow ducts between the cells of the columnar epithelium.
Text-fig. 3—Transverse section of the penis, with the terminal intromittent portion invaginated.
CART, cartilage-like supporting rod; EP, external epithelium of the penis; EJ.D, ejaculatory duct; G.EP, tract of glandular epithelium, GLD, subepithehal gland cells; HMC, haemocoele occupied by sparse connective tissue, INV, Invagination
The female genital duct is divisible into two portions—an extremely narrow, much convoluted posterior part, leading forward from the ovary to open into the stouter, glandular-walled pallial oviduct. The functions of the pallial oviduct have not yet been properly ascertained. It appears to combine the secretory role of both the albumen gland and the capsule gland as recognised by Fretter (1946) for marine prosobranchs. But the two regions are not structurally separable in Murdochia.

On leaving the ovary, the posterior portion of the female duct is at first very narrow, thin-walled and translucent, the cells lightly staining, and with long cilia; the duct is here strongly distensible with the passage of eggs. A little further forward the proximal oviduct becomes thrown into a number of coils and closely convoluted, opaque white in appearance. It is here noticeably muscular, its coat of circular fibres being well developed and the epithelial lining being thrown into 4–5 longitudinal folds by cell height differences. The epithelium contains plump, ovoid gland cells, with basal nuclei and mucous contents staining lightly in haematoxylin. Ciliated cells are also present, especially at the crests of the folds, centrally constricted and with rod-like central nuclei. The posterior-most part of the coiled oviduct is of different structure from the rest, and functions as a receptaculum seminis. The epithelium here is not folded, and the cells are of one type only, regularly columnar, clear or lightly staining, with a single row of basal nuclei. A coating of close-pressed sperm heads is found attached to the distal surfaces of the cells, the tails occupying the middle of the lumen. It is presumably in this part of the duct that fertilisation takes place before the addition of glandular secretions to the egg.
The pallial oviduct runs forward immediately below the rectum. It is pale yellow in colour, thick-walled and laterally compressed, thrown into a succession of deep transverse folds which project into the lumen as ridges of the glandular wall. The epithelium is extremely regular, the gland cells tall and narrowly columnar, with basal nuclei, and with small triangular or wedge-shaped ciliated cells inserted between their tips. The character of the secretion changes about halfway along the duct. Posteriorly the cells are filled with rows of large, finely granular vesicles or spherules, each occupying the whole width of a cell, and staining light pinkish-brown with Van Gieson's. Further forward, the secretion is in little droplets, much smaller, and staining deeper purplish, with a stronger affinity for haematoxylm. The aperture of the pallial oviduct lies far forward, just below the anus, and near the opening the histology changes. The wall becomes somewhat folded by differences in the length of the cells. The ciliated cells are longer with their cilia more prominent, and the contents of the gland cells—in the basal two-thirds—are tiny refractile spherules, staining deeply black with haematoxylin.
