Developmental integration of feather growth and pigmentation and its implications for the evolution of diet-derived coloration

Variation in avian coloration is produced by coordinated pigmentation of thousands of growing feathers that vary in shape and size. Although the functional consequences of avian coloration are frequently studied, little is known about its developmental basis, and, specifically, the rules that link feather growth to pigment uptake and synthesis. Here, we combine biochemical, modeling, and morphometric techniques to examine the developmental basis of feather pigmentation in house finches (Carpodacus mexicanus)-a species with extensive variation in both growth dynamics of ornamental feathers and their carotenoid pigmentation. We found that the rate of carotenoid uptake was constant across a wide range of feather sizes and shapes, and the relative pigmented area of feathers was independent of the total amount of deposited carotenoids. Analysis of the developmental linkage of feather growth and pigment uptake showed that the mechanisms behind partitioning the feather into pigmented and nonpigmented parts and the mechanisms regulating carotenoid uptake into growing feathers are partially independent. Carotenoid uptake strongly covaried with early elements of feather differentiation (the barb addition rate and diameter), whereas the pigmented area was most closely associated with the rate of feather growth. We suggest that strong effects of carotenoid uptake on genetically integrated mechanisms of feather growth and differentiation provide a likely route for genetic assimilation of diet-dependent coloration.