Skin structure and cornification proteins in the soft-shelled turtle Trionyx spiniferus

In contrast to most chelonians, the fully aquatic soft-shelled turtles have a smooth, unscaled, and pliable shell. The skin of the shell, tail, limbs, and neck of juveniles of Trionyx spiniferus has been studied by ultrastructural, immunocytochemical, and immunoblotting methods. The epidermis of the carapace and plastron has a thick corneous layer composed of alpha-corneocytes surrounded by a cornified cell envelope. The softer epidermis is similar to that of the shell but the epidermis and corneous layer are much thinner. Pre-corneous cells in both soft and shell epidermis are rich in vesicles produced in the Golgi apparatus and smooth endoplasmic vesicles, and contain numerous dense-core mucus-like and vesicular (lamellar) bodies. Secreted material is present among corneocytes where it probably forms an extensive intercellular lipid-mucus waterproof barrier. The dermis is very thick and composed of several layers of collagen bundles that form a plywood-patterned dermis. This dermis constitutes a strong mechanical barrier that compensates for the low content in beta-keratin, and lack of cornified scutes and dermal bones. The growth of the shell mainly occurs along the lateral margins. Immunocytochemistry reveals the presence of some beta-keratin in soft and shell epidermis, and this is confirmed by immunoblotting where bands at 18 and 32-35 kDa are present. Other proteins of the cornified cell envelope (loricrin and sciellin) or associated to lipid trafficking (caveolin-1) are also detected by immunoblotting. Loricrin positive bands at 24 and 57 kDa are present while bands cross-reactive for sciellin are seen at 45 and 53 kDa. Caveolin-1 positive bands are seen at 21-22 kDa. The presence of these proteins indicates that the epidermis is both coriaceous and waterproof. These results suggest that the shell of Trionyx is tough enough to be as mechanically efficient as the hard shell of the other turtles. At the same time, a soft shell is lighter, its shape is more easily controlled by muscles, and it allows a more controlled closure and retraction of limbs and neck inside the shell. Thus, the shell makes the animal more streamlined for swimming.