Molecular shifts in limb identity underlie development of feathered feet in two domestic avian species
Animals ranging from fish to birds display dramatic diversity within and among species; yet remarkably little is known about the genetic and developmental mechanisms that underlie this variation. In birds and their extinct dinosaur relatives, the distribution of scales and feathers on the feet is a highly variable trait. Different breeds of domestic pigeon all belong to the same species but have feet that can be feathery or scaly to different extents. Classical genetics experiments suggested that only a few genes are involved in the transition from scaled to feathered skin on the feet of pigeons. However, the molecular basis for this transition was unknown. Domyan et al. set out to identify the genes involved in the transition from scaled to feathered feet by mating different breeds of pigeon in the laboratory and then sequencing the birds’ DNA. They also surveyed the entire DNA sequences of many additional pigeon breeds with and without feathered feet. This combined approach showed that two regions of the pigeon genome have a profound effect on the number and size of feathers on the feet of domestic pigeons. These regions contain genes that are known to play key roles in controlling the growth of a limb and whether it develops into a leg or a wing. In developing pigeon embryos, Domyan et al. found that a gene called Pitx1, which is typically considered a hindlimb gene, is expressed at lower levels in the developing legs of breeds with feathered feet than in a breed with scaled feet. The experiments also showed that Tbx5, a gene that is expressed in the forelimbs of many animals, is expressed abnormally in the embryonic hindlimbs of breeds of pigeon and chicken with feathery feet. Together, these findings suggest that the hindlimbs of domestic birds with feathery feet are more like wings at the molecular level, which results in them being covered in feathers rather than scales. Future work will now aim to discover the specific DNA sequences that alter the expression of Pitx1 and Tbx5 in feather-footed breeds, and whether the same genes control the foot feathers of other species of birds.