Development, structure, and protein composition of the corneous beak in turtles

The beak or rhamphotheca in turtles is a horny lamina that replaces the teeth. Its origin, development, structure, and protein composition are here presented. At mid-development stages, the epidermis of the maxilla and mandible gives rise to placodes that enlarge and merge into laminae through an intense cell proliferation. In these expanding laminae, the epidermis gives rise to 5-8 layers of embryonic epidermis where coarse filaments accumulate for the initial keratinization of cells destined to be sloughed before hatching. Underneath the embryonic epidermis of the beak numerous layers of spindle-shaped beta-cells are produced while they are absent in other skin regions. Beta-cells contain hard corneous material and give rise to the corneous layer of the beak whose external layers desquamate due to wearing and mechanical abrasion. Beta-catenin is present in nuclei of proliferating keratinocytes of the germinal layer likely responding to a wnt signal, but also is part of the adhesive junctions located among beak keratinocytes. The thick corneous layer is made of mature corneocytes connected one to another along their irregular perimeter by an unknown cementing material and junctional remnants. Immunolabeling shows that the main components of the horny beak are Corneous Beta Proteins (CBPs) of 10-15 kDa which genes are located in the Epidermal Differentiation Complex (EDC) of the turtle genome. Specific CBPs, in addition to a lower amount of Intermediate Filament Keratins, accumulate in the horny beak. Compaction of the main proteins with other unknown, minor proteins give rise to the hard corneous material of the beak.