Ultrastructure of the feather follicle in relation to the formation of the rachis in pennaceous feathers

The present ultrastructural study on follicle of regenerating feathers of four different avian species focuses on the formation and cytology of the rachis. Epithelial cells within the bottom part of the follicle (the collar) are contacted from mesenchymal cells of the dermal papilla. The most basal part of the collar is formed by a circular epithelium containing germinal cells, while in the upper ramogenic part of the collar barb ridges are generated. Epithelial cells rest upon a basement membrane that is stretched in actively forming barb ridges among which anchored mesenchymal cells send thin elongation. This observation suggests that an intense exchange of molecules with the epithelium occurs. The process of formation of the rachis occurs by fusion of barb ridges with the nonsegmented, dorsal or anterior part of the collar. The latter becomes the rachidial ridge, the upper part of the collar where barbs form the branches of the pennaceous feather. The rachis grows and matures into an external cortical part, containing compact corneous material (feather keratin, as confirmed by immunocytochemistry), and a vacuolated medulla with a process similar to that occurring in rami of single barbs. The extension of the medulla and cortex varies along the rachis in different species. In general a thin cortex is formed in those sections of the rachis where barbs are absent, and the feather keratin positive layer increases in the basal part of the feather, the calamus.