Cell structure of barb ridges in down feathers and juvenile wing feathers of the developing chick embryo: barb ridge modification in relation to feather evolution.

The present study deals with the cell structure and three-dimensional organization of barb and barbule cells within barb ridges of down feathers and juvenile feathers in the chick embryo. Juvenile feathers represent the second generation of feathers in the wing, and replace down feathers some weeks after hatching. Within the follicle of juvenile feathers, at 16-18 days of embryonic development, barb ridges are more numerous than in down feathers. Barb ridges of juvenile feathers contain more cells in their barbule and axial plates with respect to barb ridges of down feathers. This condition determines the formation of longer barbules inserted in the rami of juvenile feathers than barbules of down feathers. Barb ridges of juvenile feathers merge with the rachidial ridge so that pennaceous feathers are formed. Barbule cells are surrounded by cytoplasmic elongation from barb vane ridge cells located in the axial plate, which constitute most of the axial plate. The degeneration of supportive cells among barbule cells branching from barbs determine the formation of spaces between barbules. The study emphasizes that, in addition to the size of the dermal papilla, it is the length of barb ridges and the infiltration of barb ridge vane cells among barbule cells that determine the size and length of feathers. The knowledge of the cell structure of barb ridges allows understanding not only of how feathers develop but also gives insights into their evolution. Based on changes of the process of barb ridge morphogenesis some hypotheses on the evolution of plumulaceous and pennaceous feathers are presented. Feathers derived from the process of carving-out supportive cells within barb ridges and from the specific pattern of fusion of barb/barbule cells. This process initially produced variably branched down feathers and later, after barb ridge fusion, a rachis. From the modulation in the pattern of barb ridge formation various pennaceous feathers later evolved.