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American Zoologist 2000 40(4):631-639; doi:10.1093/icb/40.4.631
© 2000 by The Society for Integrative and Comparative Biology
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Evolving a Protofeather and Feather Diversity1

Alan H. Brush2,1
1 92 High St., Mystic, Connecticut 06355

It is likely that feathers evolved from a conical shaped tubercle rather than a plate-like structure. Although the morphology of the presumably most primitive feather is unknown, minimal conditions for its production include the cellular capacity to synthesize feather proteins (={phi}-keratin) which provides the molecular phenotype, and a follicular mechanism for production and assembly of molecular and gross structure. Once the minimal structural element, presumably recognizable as a barb, existed, a variety of phenotypes followed rapidly. A tubercular growth center of appropriate size could produce a simple barb-like element, with cortex and medulla. This might be recognized externally as a bristle, but need never existed as a separate morphological unit. Rather, if individual placodes gave rise to multiple barb ridges that fused proximally, a structure resembling natal down would have resulted. Subsequent differentiation is controlled by the follicular symmetry, and the feather shape is regulated by barb length. Barb length is directly related to growth period. As feathers appear to grow at roughly similar, size independent rates, shape is determined by individual barb growth periods. The simple fusion of individual proto-barbs would produce a morphology identifiable as natal down. Although this might be the simplest feather structure, others could emerge quickly, perhaps simultaneously, a consequence of the same redundant processing. Once the machinery existed, broad phenotypic plasticity was possible. I constructed a feather phylogram based on these conditions, the fossil record, and ontogeny. I organized the subsequent changes in morphology by perceived complexity. The changes are simply individual responses to similar processes that might be time (when in ontogeny) and space (where on body) dependent.


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