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In order to understand these models, knowledge of the development of the avian integument must be put into an evolutionary context. Thus, we present a review of avian scale and feather development, which summarizes the morphogenetic events involved, as well as the expression of the beta (\u03b2) keratin multigene family that characterizes the epidermal appendages of reptiles and birds. First we review information on the evolution of the ectodermal epidermis and its beta (\u03b2) keratins. Then we examine the morphogenesis of scutate scales and feathers including studies in which the extraembryonic ectoderm of the chorion is used to examine dermal induction. We also present studies on the scaleless (<jats:italic>sc<\/jats:italic>) mutant, and, because of the recent discovery of \u201cfour\u2010winged\u201d dinosaurs, we review earlier studies of a chicken strain, Silkie, that expresses ptilopody (<jats:italic>pti<\/jats:italic>), \u201cfeathered feet.\u201d We conclude that the ability of the ectodermal epidermis to generate discrete cell populations capable of forming functional structural elements consisting of specific members of the \u03b2 keratin multigene family was a plesiomorphic feature of the archosaurian ancestor of crocodilians and birds. Evidence suggests that the discrete epidermal lineages that make up the <jats:italic>embryonic feather filament<\/jats:italic> of extant birds are homologous with similar embryonic lineages of the developing scutate scales of birds and the scales of alligators. We believe that the early expression of conserved signaling modules in the embryonic skin of the avian ancestor led to the early morphogenesis of the <jats:italic>embryonic feather filament<\/jats:italic>, with its periderm, sheath, and barb ridge lineages forming the first protofeather. Invagination of the epidermis of the protofeather led to formation of the follicle providing for feather renewal and diversification. The observations that scale formation in birds involves an inhibition of feather formation coupled with observations on the feathered feet of the scaleless (High\u2010line) and Silkie strains support the view that the ancestor of modern birds may have had feathered hind limbs similar to those recently discovered in nonavian dromaeosaurids. And finally, our recent observation on the bristles of the wild turkey beard raises the possibility that similar integumentary appendages may have adorned nonavian dinosaurs, and thus all <jats:italic>filamentous integumentary appendages<\/jats:italic> may not be homologous to modern feathers. <jats:italic>J. Exp. Zool. (Mol. Dev. Evol.) 298B:57\u201372, 2003.<\/jats:italic> \u00a9 2003 Wiley\u2010Liss, Inc.<\/jats:p>","DOI":"10.1002\/jez.b.26","type":"journal-article","created":{"date-parts":[[2003,8,27]],"date-time":"2003-08-27T13:42:26Z","timestamp":1061991746000},"page":"57-72","source":"Crossref","is-referenced-by-count":121,"title":["Avian skin development and the evolutionary origin of feathers"],"prefix":"10.1002","volume":"298B","author":[{"given":"Roger H.","family":"Sawyer","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Loren W.","family":"Knapp","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2003,8,26]]},"reference":[{"key":"e_1_2_1_2_1","first-page":"1347","article-title":"Selection for feather number in scaleless chickens","volume":"44","author":"Abbott UK","year":"1965","journal-title":"Poult Sci"},{"key":"e_1_2_1_3_1","volume-title":"Methods in developmental biology","author":"Abbott UK","year":"1967"},{"key":"e_1_2_1_4_1","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1093\/oxfordjournals.jhered.a106685","article-title":"Scaleless, an inherited ectodermal defect in the domestic fowl","volume":"18","author":"Abbott UK","year":"1957","journal-title":"J Hered"},{"key":"e_1_2_1_5_1","doi-asserted-by":"publisher","DOI":"10.1046\/j.1523-1747.1999.00592.x"},{"key":"e_1_2_1_6_1","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1007\/BF00335521","article-title":"Comparison of \u03b1\u2010 and \u03b2\u2010keratin in reptiles","volume":"110","author":"Alexander NJ","year":"1970","journal-title":"Z Zellforsch"},{"key":"e_1_2_1_7_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF00335586"},{"key":"e_1_2_1_8_1","doi-asserted-by":"publisher","DOI":"10.1054\/tice.2001.0198"},{"key":"e_1_2_1_9_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.10079"},{"key":"e_1_2_1_10_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.10076"},{"key":"e_1_2_1_11_1","doi-asserted-by":"publisher","DOI":"10.1002\/jez.10145"},{"key":"e_1_2_1_12_1","doi-asserted-by":"publisher","DOI":"10.1046\/j.1469-7580.2001.19830265.x"},{"key":"e_1_2_1_13_1","doi-asserted-by":"crossref","unstructured":"AltmanPL KatzDD.1979.Biological Handbooks III. 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