Ontogeny of the Molluscan Shell Field: a Review

In the gastropod, scaphopod, lamellibranch, and cephalopod gastrulae a thickened portion of the posttrochal region is referred to as the embryonic shell field. It invaginates and gives rise to the shell gland. In species with an at least temporarily external shell, the shell gland evaginates and again forms a shell field. In lamellibranchs, the shell field grows into two halves connected by the ligament‐secreting isthmus. In polyplacophorans plate fields are produced without invagination. Slugs and endocochleate cephalopods overgrow the embryonic shell field to form an internal shell sac. The calcified part of the shell is secreted by the flattened central region. The periostracum has its origin in the permanently thickened peripheral region of the shell field. In many forms, this region is depressed in a periostracal groove. If the shell is external, the central region of flattened cells, the mantle roof, along with the two or three marginal folds of the free mantle edge and, in species with internal shell, the shell sac are parts of the mantle. The shell field descends from the first somatoblasts. Either of 2 d or 2 c alone is able to form the shell field. There are arguments that the formation of the embryonic shell field is not autonomic, but induced by the entoderm during a period of contact. The shell gland and the shell field grow by mitotic cell divisions. Cells secreting organic material are highly prismatic, have a well developed ergastoplasm and large dictyosornes, and contain much peroxidase. The secretion of calcium manifests itself in very flat cells, rich in alkaline phosphatase and glycogen. The shell gland and the rosette of ectocochleate conchifera together are homologous to the proximal part of the shell sac in slugs and endocochleate cephalopods.