{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T21:44:01Z","timestamp":1792791841899,"version":"4.5.0"},"reference-count":69,"publisher":"National Academy of Sciences","issue":"49","license":[{"start":{"date-parts":[[2016,5,23]],"date-time":"2016-05-23T00:00:00Z","timestamp":1463961600000},"content-version":"vor","delay-in-days":182,"URL":"http:\/\/www.pnas.org\/site\/misc\/userlicense.xhtml"}],"funder":[{"name":"HHS | NIH | National Institute of Arthritis and Musculoskeletal and Skin Diseases","award":["AR 47364"],"award-info":[{"award-number":["AR 47364"]}],"id":[{"id":"https:\/\/ror.org\/006zn3t30","id-type":"ROR","asserted-by":"crossref"}]},{"DOI":"10.13039\/100000069","name":"HHS | NIH | National Institute of Arthritis and Musculoskeletal and Skin Diseases","doi-asserted-by":"publisher","award":["AR 60306"],"award-info":[{"award-number":["AR 60306"]}],"id":[{"id":"10.13039\/100000069","id-type":"DOI","asserted-by":"publisher"},{"id":"https:\/\/ror.org\/006zn3t30","id-type":"ROR","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100004663","name":"Ministry of Science and Technology, Taiwan","doi-asserted-by":"crossref","award":["MOST 104-2621-B-001-003-MY3"],"award-info":[{"award-number":["MOST 104-2621-B-001-003-MY3"]}],"id":[{"id":"10.13039\/501100004663","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["www.pnas.org"],"crossmark-restriction":true},"short-container-title":["Proc. Natl. Acad. Sci. U.S.A."],"published-print":{"date-parts":[[2015,12,8]]},"abstract":"<jats:title>Significance<\/jats:title>\n                  <jats:p>Avian skin appendages include feathers, scales, claws, and beaks. They are mainly composed of \u03b1-keratins, found in all vertebrates, and \u03b2-keratins, found only in birds and reptiles. Scientists have wondered how keratins are interwoven to form different skin appendages. By studying keratin gene expression patterns in different chicken skin appendages, we found \u03b1- and \u03b2-keratin interactions crucial for appendage morphogenesis. Mutations in either \u03b1- or \u03b2-keratins can disrupt keratin expression and cause structural defects. Thus, different combinations of \u03b1- and \u03b2-keratins contribute to the structural diversity of feathers. The expansion of \u03b2-keratin genes during bird evolution might have greatly increased skin appendage diversity because it increased the possible interactions between \u03b1- and \u03b2-keratins.<\/jats:p>","DOI":"10.1073\/pnas.1520566112","type":"journal-article","created":{"date-parts":[[2015,11,23]],"date-time":"2015-11-23T22:19:00Z","timestamp":1448317140000},"update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":82,"title":["Topographical mapping of \u03b1- and \u03b2-keratins on developing chicken skin integuments: Functional interaction and evolutionary perspectives"],"prefix":"10.1073","volume":"112","author":[{"given":"Ping","family":"Wu","sequence":"first","affiliation":[{"name":"Department of Pathology, Keck School of Medicine, University of Southern California, Los Angeles, CA 90033;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chen Siang","family":"Ng","sequence":"additional","affiliation":[{"name":"Biodiversity Research Center, Academia Sinica, Taipei 11529, Taiwan;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jie","family":"Yan","sequence":"additional","affiliation":[{"name":"Department of Pathology, Keck School of Medicine, University of Southern California, Los Angeles, CA 90033;"},{"name":"Jiangsu Key Laboratory for Biodiversity and Biotechnology, College of Life Sciences, Nanjing Normal University, Nanjing 210023, China;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yung-Chih","family":"Lai","sequence":"additional","affiliation":[{"name":"Department of Pathology, Keck School of Medicine, University of Southern California, Los Angeles, CA 90033;"},{"name":"Research Center for Developmental Biology and Regenerative Medicine, National Taiwan University, Taipei 10041, Taiwan;"},{"name":"Integrative Stem Cell Center, China Medical University, Taichung 40447, Taiwan;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chih-Kuan","family":"Chen","sequence":"additional","affiliation":[{"name":"Biodiversity Research Center, Academia Sinica, Taipei 11529, Taiwan;"},{"name":"Institute of Ecology and Evolutionary Biology, National Taiwan University, Taipei 10617, Taiwan;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yu-Ting","family":"Lai","sequence":"additional","affiliation":[{"name":"Biodiversity Research Center, Academia Sinica, Taipei 11529, Taiwan;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Siao-Man","family":"Wu","sequence":"additional","affiliation":[{"name":"Biodiversity Research Center, Academia Sinica, Taipei 11529, Taiwan;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiun-Jie","family":"Chen","sequence":"additional","affiliation":[{"name":"Biodiversity Research Center, Academia Sinica, Taipei 11529, Taiwan;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Weiqi","family":"Luo","sequence":"additional","affiliation":[{"name":"Department of Pathology, Keck School of Medicine, University of Southern California, Los Angeles, CA 90033;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Randall B.","family":"Widelitz","sequence":"additional","affiliation":[{"name":"Department of Pathology, Keck School of Medicine, University of Southern California, Los Angeles, CA 90033;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wen-Hsiung","family":"Li","sequence":"additional","affiliation":[{"name":"Biodiversity Research Center, Academia Sinica, Taipei 11529, Taiwan;"},{"name":"Department of Ecology and Evolution, University of Chicago, Chicago, IL 60637;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cheng-Ming","family":"Chuong","sequence":"additional","affiliation":[{"name":"Department of Pathology, Keck School of Medicine, University of Southern California, Los Angeles, CA 90033;"},{"name":"Research Center for Developmental Biology and Regenerative Medicine, National Taiwan University, Taipei 10041, Taiwan;"},{"name":"Integrative Stem Cell Center, China Medical University, Taichung 40447, Taiwan;"},{"name":"Center for the Integrative and Evolutionary Galliform Genomics, National Chung Hsing University, Taichung 40227, Taiwan"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"341","published-online":{"date-parts":[[2015,11,23]]},"reference":[{"key":"e_1_3_4_1_2","doi-asserted-by":"crossref","first-page":"194","DOI":"10.1007\/978-3-662-00989-5_11","volume-title":"Biology of the Integument 2 Vertebrates","author":"Sawyer RH","year":"1986","unstructured":"RH Sawyer, LW Knapp, WM O\u2019Guin, Epidermis, dermis and appendages. Biology of the Integument 2 Vertebrates, eds J Bereiter-Hahn, AG Matoltsy, KS Richards (Springer, Berlin) Vol 2, 194\u2013238 (1986)."},{"key":"e_1_3_4_2_2","first-page":"159","article-title":"What is the \u2018true\u2019 function of skin?","volume":"11","author":"Chuong CM","year":"2002","unstructured":"CM Chuong, , What is the \u2018true\u2019 function of skin? Exp Dermatol 11, 159\u2013187 (2002).","journal-title":"Exp Dermatol"},{"key":"e_1_3_4_3_2","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1002\/jez.b.23","article-title":"Development and evolution of the amniote integument: Current landscape and future horizon","volume":"298","author":"Chuong CM","year":"2003","unstructured":"CM Chuong, DG Homberger, Development and evolution of the amniote integument: Current landscape and future horizon. J Exp Zoolog B Mol Dev Evol 298, 1\u201311 (2003).","journal-title":"J Exp Zoolog B Mol Dev Evol"},{"key":"e_1_3_4_4_2","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1387\/ijdb.15272390","article-title":"Evo-Devo of amniote integuments and appendages","volume":"48","author":"Wu P","year":"2004","unstructured":"P Wu, , Evo-Devo of amniote integuments and appendages. Int J Dev Biol 48, 249\u2013270 (2004).","journal-title":"Int J Dev Biol"},{"key":"e_1_3_4_5_2","doi-asserted-by":"crossref","first-page":"1252","DOI":"10.1073\/pnas.1423685112","article-title":"Seeing is believing: Dynamic evolution of gene families","volume":"112","author":"Harris RM","year":"2015","unstructured":"RM Harris, HA Hofmann, Seeing is believing: Dynamic evolution of gene families. Proc Natl Acad Sci USA 112, 1252\u20131253 (2015).","journal-title":"Proc Natl Acad Sci USA"},{"key":"e_1_3_4_6_2","doi-asserted-by":"crossref","DOI":"10.1007\/978-3-642-86659-3","volume-title":"Evolution by Gene Duplication","author":"Ohno S","year":"1970","unstructured":"S Ohno Evolution by Gene Duplication (Springer, New York, 1970)."},{"key":"e_1_3_4_7_2","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1146\/annurev-animal-022513-114127","article-title":"Development, regeneration, and evolution of feathers","volume":"3","author":"Chen CF","year":"2015","unstructured":"CF Chen, , Development, regeneration, and evolution of feathers. Annu Rev Anim Biosci 3, 169\u2013195 (2015).","journal-title":"Annu Rev Anim Biosci"},{"key":"e_1_3_4_8_2","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1387\/ijdb.15272383","article-title":"The biology of feather follicles","volume":"48","author":"Yu M","year":"2004","unstructured":"M Yu, , The biology of feather follicles. Int J Dev Biol 48, 181\u2013191 (2004).","journal-title":"Int J Dev Biol"},{"key":"e_1_3_4_9_2","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1111\/j.1432-0436.1988.tb00205.x","article-title":"Identification, expression, and localization of beta keratin gene products during development of avian scutate scales","volume":"38","author":"Shames RB","year":"1988","unstructured":"RB Shames, LW Knapp, WE Carver, RH Sawyer, Identification, expression, and localization of beta keratin gene products during development of avian scutate scales. Differentiation 38, 115\u2013123 (1988).","journal-title":"Differentiation"},{"key":"e_1_3_4_10_2","doi-asserted-by":"crossref","first-page":"66","DOI":"10.1002\/aja.1001840108","article-title":"Immunocytochemical localization and biochemical analysis of alpha and beta keratins in the avian lingual epithelium","volume":"184","author":"Carver WE","year":"1989","unstructured":"WE Carver, RH Sawyer, Immunocytochemical localization and biochemical analysis of alpha and beta keratins in the avian lingual epithelium. Am J Anat 184, 66\u201375 (1989).","journal-title":"Am J Anat"},{"key":"e_1_3_4_11_2","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.proghi.2008.01.001","article-title":"Cytochemical and molecular characteristics of the process of cornification during feather morphogenesis","volume":"43","author":"Alibardi L","year":"2008","unstructured":"L Alibardi, M Toni, Cytochemical and molecular characteristics of the process of cornification during feather morphogenesis. Prog Histochem Cytochem 43, 1\u201369 (2008).","journal-title":"Prog Histochem Cytochem"},{"key":"e_1_3_4_12_2","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1093\/molbev\/msu309","article-title":"Feather development genes and associated regulatory innovation predate the origin of Dinosauria","volume":"32","author":"Lowe CB","year":"2015","unstructured":"CB Lowe, JA Clarke, AJ Baker, D Haussler, SV Edwards, Feather development genes and associated regulatory innovation predate the origin of Dinosauria. Mol Biol Evol 32, 23\u201328 (2015).","journal-title":"Mol Biol Evol"},{"key":"e_1_3_4_13_2","doi-asserted-by":"crossref","first-page":"1311","DOI":"10.1126\/science.1251385","article-title":"Comparative genomics reveals insights into avian genome evolution and adaptation","volume":"346","author":"Zhang G","year":"2014","unstructured":"G Zhang, , Comparative genomics reveals insights into avian genome evolution and adaptation. Science; Avian Genome Consortium 346, 1311\u20131320 (2014).","journal-title":"Science"},{"key":"e_1_3_4_14_2","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1186\/s12862-014-0249-1","article-title":"Dynamic evolution of the alpha (\u03b1) and beta (\u03b2) keratins has accompanied integument diversification and the adaptation of birds into novel lifestyles","volume":"14","author":"Greenwold MJ","year":"2014","unstructured":"MJ Greenwold, , Dynamic evolution of the alpha (\u03b1) and beta (\u03b2) keratins has accompanied integument diversification and the adaptation of birds into novel lifestyles. BMC Evol Biol 14, 249 (2014).","journal-title":"BMC Evol Biol"},{"key":"e_1_3_4_15_2","doi-asserted-by":"crossref","first-page":"995","DOI":"10.1093\/molbev\/msr269","article-title":"Radiation and functional diversification of alpha keratins during early vertebrate evolution","volume":"29","author":"Vandebergh W","year":"2012","unstructured":"W Vandebergh, F Bossuyt, Radiation and functional diversification of alpha keratins during early vertebrate evolution. Mol Biol Evol 29, 995\u20131004 (2012).","journal-title":"Mol Biol Evol"},{"key":"e_1_3_4_16_2","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1002\/(SICI)1097-010X(19991215)285:4<291::AID-JEZ1>3.0.CO;2-9","article-title":"Development and evolutionary origin of feathers","volume":"285","author":"Prum RO","year":"1999","unstructured":"RO Prum, Development and evolutionary origin of feathers. J Exp Zool 285, 291\u2013306 (1999).","journal-title":"J Exp Zool"},{"key":"e_1_3_4_17_2","doi-asserted-by":"crossref","first-page":"570","DOI":"10.1002\/jez.b.21073","article-title":"Evolution of the morphological innovations of feathers","volume":"304","author":"Prum RO","year":"2005","unstructured":"RO Prum, Evolution of the morphological innovations of feathers. J Exp Zoolog B Mol Dev Evol 304, 570\u2013579 (2005).","journal-title":"J Exp Zoolog B Mol Dev Evol"},{"key":"e_1_3_4_18_2","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1086\/341993","article-title":"The evolutionary origin and diversification of feathers","volume":"77","author":"Prum RO","year":"2002","unstructured":"RO Prum, AH Brush, The evolutionary origin and diversification of feathers. Q Rev Biol 77, 261\u2013295 (2002).","journal-title":"Q Rev Biol"},{"key":"e_1_3_4_19_2","doi-asserted-by":"crossref","first-page":"1253293","DOI":"10.1126\/science.1253293","article-title":"An integrative approach to understanding bird origins","volume":"346","author":"Xu X","year":"2014","unstructured":"X Xu, , An integrative approach to understanding bird origins. Science 346, 1253293 (2014).","journal-title":"Science"},{"key":"e_1_3_4_20_2","doi-asserted-by":"crossref","first-page":"951","DOI":"10.1073\/pnas.0506894103","article-title":"Wnt3a gradient converts radial to bilateral feather symmetry via topological arrangement of epithelia","volume":"103","author":"Yue Z","year":"2006","unstructured":"Z Yue, TX Jiang, RB Widelitz, CM Chuong, Wnt3a gradient converts radial to bilateral feather symmetry via topological arrangement of epithelia. Proc Natl Acad Sci USA 103, 951\u2013955 (2006).","journal-title":"Proc Natl Acad Sci USA"},{"key":"e_1_3_4_21_2","doi-asserted-by":"crossref","first-page":"307","DOI":"10.1016\/j.diff.2011.03.004","article-title":"From buds to follicles: Matrix metalloproteinases in developmental tissue remodeling during feather morphogenesis","volume":"81","author":"Jiang TX","year":"2011","unstructured":"TX Jiang, TL Tuan, P Wu, RB Widelitz, CM Chuong, From buds to follicles: Matrix metalloproteinases in developmental tissue remodeling during feather morphogenesis. Differentiation 81, 307\u2013314 (2011).","journal-title":"Differentiation"},{"key":"e_1_3_4_22_2","doi-asserted-by":"crossref","first-page":"1026","DOI":"10.1038\/nature04222","article-title":"Mapping stem cell activities in the feather follicle","volume":"438","author":"Yue Z","year":"2005","unstructured":"Z Yue, TX Jiang, RB Widelitz, CM Chuong, Mapping stem cell activities in the feather follicle. Nature 438, 1026\u20131029 (2005).","journal-title":"Nature"},{"key":"e_1_3_4_23_2","doi-asserted-by":"crossref","first-page":"1212","DOI":"10.1016\/j.cell.2014.08.019","article-title":"SnapShot: Branching morphogenesis","volume":"158","author":"Chuong CM","year":"2014","unstructured":"CM Chuong, R Bhat, RB Widelitz, MJ Bissell, SnapShot: Branching morphogenesis. Cell 158, 1212\u20131212.e1 (2014).","journal-title":"Cell"},{"key":"e_1_3_4_24_2","doi-asserted-by":"crossref","first-page":"148","DOI":"10.1186\/1471-2148-10-148","article-title":"Genomic organization and molecular phylogenies of the beta (beta) keratin multigene family in the chicken (Gallus gallus) and zebra finch (Taeniopygia guttata): Implications for feather evolution","volume":"10","author":"Greenwold MJ","year":"2010","unstructured":"MJ Greenwold, RH Sawyer, Genomic organization and molecular phylogenies of the beta (beta) keratin multigene family in the chicken (Gallus gallus) and zebra finch (Taeniopygia guttata): Implications for feather evolution. BMC Evol Biol 10, 148 (2010).","journal-title":"BMC Evol Biol"},{"key":"e_1_3_4_25_2","doi-asserted-by":"crossref","first-page":"2258","DOI":"10.1093\/gbe\/evu181","article-title":"Genomic organization, transcriptomic analysis, and functional characterization of avian \u03b1- and \u03b2-keratins in diverse feather forms","volume":"6","author":"Ng CS","year":"2014","unstructured":"CS Ng, , Genomic organization, transcriptomic analysis, and functional characterization of avian \u03b1- and \u03b2-keratins in diverse feather forms. Genome Biol Evol 6, 2258\u20132273 (2014).","journal-title":"Genome Biol Evol"},{"key":"e_1_3_4_26_2","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1002\/jez.b.22514","article-title":"Molecular evolution and expression of archosaurian \u03b2-keratins: Diversification and expansion of archosaurian \u03b2-keratins and the origin of feather \u03b2-keratins","volume":"320","author":"Greenwold MJ","year":"2013","unstructured":"MJ Greenwold, RH Sawyer, Molecular evolution and expression of archosaurian \u03b2-keratins: Diversification and expansion of archosaurian \u03b2-keratins and the origin of feather \u03b2-keratins. J Exp Zoolog B Mol Dev Evol 320, 393\u2013405 (2013).","journal-title":"J Exp Zoolog B Mol Dev Evol"},{"key":"e_1_3_4_27_2","doi-asserted-by":"crossref","first-page":"609","DOI":"10.1002\/jez.b.21436","article-title":"Linking the molecular evolution of avian beta (\u03b2) keratins to the evolution of feathers","volume":"316","author":"Greenwold MJ","year":"2011","unstructured":"MJ Greenwold, RH Sawyer, Linking the molecular evolution of avian beta (\u03b2) keratins to the evolution of feathers. J Exp Zoolog B Mol Dev Evol 316, 609\u2013616 (2011).","journal-title":"J Exp Zoolog B Mol Dev Evol"},{"key":"e_1_3_4_28_2","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1002\/jez.b.25","article-title":"Adaptation to the sky: Defining the feather with integument fossils from mesozoic China and experimental evidence from molecular laboratories","volume":"298","author":"Chuong CM","year":"2003","unstructured":"CM Chuong, , Adaptation to the sky: Defining the feather with integument fossils from mesozoic China and experimental evidence from molecular laboratories. J Exp Zoolog B Mol Dev Evol 298, 42\u201356 (2003).","journal-title":"J Exp Zoolog B Mol Dev Evol"},{"key":"e_1_3_4_29_2","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1016\/j.gene.2014.03.027","article-title":"Identification of a feather \u03b2-keratin gene exclusively expressed in pennaceous barbule cells of contour feathers in chicken","volume":"542","author":"Kowata K","year":"2014","unstructured":"K Kowata, , Identification of a feather \u03b2-keratin gene exclusively expressed in pennaceous barbule cells of contour feathers in chicken. Gene 542, 23\u201328 (2014).","journal-title":"Gene"},{"key":"e_1_3_4_30_2","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1002\/jmor.1050880104","article-title":"A series of normal stages in the development of the chick embryo","volume":"88","author":"Hamburger V","year":"1951","unstructured":"V Hamburger, HL Hamilton, A series of normal stages in the development of the chick embryo. J Morphol 88, 49\u201392 (1951).","journal-title":"J Morphol"},{"key":"e_1_3_4_31_2","unstructured":"AM Lucas PR Stettenheim Avian anatomy. Integument. Agriculture Handbook 362 (US Dept Ag Washington DC). (1972)."},{"key":"e_1_3_4_32_2","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1002\/jez.1401900106","article-title":"Avian scale development. IV. Ultrastructure of the anterior shank skin of the scaleless mutant","volume":"190","author":"Sawyer RH","year":"1974","unstructured":"RH Sawyer, UK Abbott, GN Fry, Avian scale development. IV. Ultrastructure of the anterior shank skin of the scaleless mutant. J Exp Zool 190, 71\u201378 (1974).","journal-title":"J Exp Zool"},{"key":"e_1_3_4_33_2","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1002\/jez.1401900105","article-title":"Avian scale development. III. Ultrastructure of the keratinizing cells of the outer and inner epidermal surfaces of the scale ridge","volume":"190","author":"Sawyer RH","year":"1974","unstructured":"RH Sawyer, UK Abbott, GN Fry, Avian scale development. III. Ultrastructure of the keratinizing cells of the outer and inner epidermal surfaces of the scale ridge. J Exp Zool 190, 57\u201370 (1974).","journal-title":"J Exp Zool"},{"key":"e_1_3_4_34_2","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1002\/jez.b.26","article-title":"Avian skin development and the evolutionary origin of feathers","volume":"298","author":"Sawyer RH","year":"2003","unstructured":"RH Sawyer, LW Knapp, Avian skin development and the evolutionary origin of feathers. J Exp Zoolog B Mol Dev Evol 298, 57\u201372 (2003).","journal-title":"J Exp Zoolog B Mol Dev Evol"},{"key":"e_1_3_4_35_2","doi-asserted-by":"crossref","first-page":"256","DOI":"10.1002\/dvdy.20291","article-title":"Evolutionary origin of the feather epidermis","volume":"232","author":"Sawyer RH","year":"2005","unstructured":"RH Sawyer, L Rogers, L Washington, TC Glenn, LW Knapp, Evolutionary origin of the feather epidermis. Dev Dyn 232, 256\u2013267 (2005).","journal-title":"Dev Dyn"},{"key":"e_1_3_4_36_2","first-page":"175","article-title":"Beta-keratin localization in developing alligator scales and feathers in relation to the development and evolution of feathers","volume":"38","author":"Alibardi L","year":"2006","unstructured":"L Alibardi, LW Knapp, RH Sawyer, Beta-keratin localization in developing alligator scales and feathers in relation to the development and evolution of feathers. J Submicrosc Cytol Pathol 38, 175\u2013192 (2006).","journal-title":"J Submicrosc Cytol Pathol"},{"key":"e_1_3_4_37_2","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1186\/s12862-015-0360-y","article-title":"Convergent evolution of cysteine-rich proteins in feathers and hair","volume":"15","author":"Strasser B","year":"2015","unstructured":"B Strasser, V Mlitz, M Hermann, E Tschachler, L Eckhart, Convergent evolution of cysteine-rich proteins in feathers and hair. BMC Evol Biol 15, 82 (2015).","journal-title":"BMC Evol Biol"},{"key":"e_1_3_4_38_2","doi-asserted-by":"crossref","first-page":"771","DOI":"10.1021\/pr301036k","article-title":"Chicken corneocyte cross-linked proteome","volume":"12","author":"Rice RH","year":"2013","unstructured":"RH Rice, BR Winters, BP Durbin-Johnson, DM Rocke, Chicken corneocyte cross-linked proteome. J Proteome Res 12, 771\u2013776 (2013).","journal-title":"J Proteome Res"},{"key":"e_1_3_4_39_2","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1016\/j.jdermsci.2009.12.002","article-title":"Novel and recurrent mutations in Keratin 5 and 14 in Korean patients with Epidermolysis bullosa simplex","volume":"57","author":"Kang TW","year":"2010","unstructured":"TW Kang, JS Lee, SE Kim, SW Oh, SC Kim, Novel and recurrent mutations in Keratin 5 and 14 in Korean patients with Epidermolysis bullosa simplex. J Dermatol Sci 57, 90\u201394 (2010).","journal-title":"J Dermatol Sci"},{"key":"e_1_3_4_40_2","doi-asserted-by":"crossref","first-page":"1083","DOI":"10.1111\/j.0022-202X.2005.23739.x","article-title":"A novel deletion mutation in keratin 5 causing the removal of 5 amino acids and elevated mutant mRNA levels in Dowling-Meara epidermolysis bullosa simplex","volume":"124","author":"Kemp MW","year":"2005","unstructured":"MW Kemp, , A novel deletion mutation in keratin 5 causing the removal of 5 amino acids and elevated mutant mRNA levels in Dowling-Meara epidermolysis bullosa simplex. J Invest Dermatol 124, 1083\u20131085 (2005).","journal-title":"J Invest Dermatol"},{"key":"e_1_3_4_41_2","doi-asserted-by":"crossref","first-page":"293","DOI":"10.1038\/sj.ejhg.5200292","article-title":"Donor splice site mutation in keratin 5 causes in-frame removal of 22 amino acids of H1 and 1A rod domains in Dowling-Meara epidermolysis bullosa simplex","volume":"7","author":"Rugg EL","year":"1999","unstructured":"EL Rugg, , Donor splice site mutation in keratin 5 causes in-frame removal of 22 amino acids of H1 and 1A rod domains in Dowling-Meara epidermolysis bullosa simplex. Eur J Hum Genet 7, 293\u2013300 (1999).","journal-title":"Eur J Hum Genet"},{"key":"e_1_3_4_42_2","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1111\/j.1749-6632.1985.tb50425.x","article-title":"Genes for hair and avian keratins","volume":"455","author":"Rogers GE","year":"1985","unstructured":"GE Rogers, Genes for hair and avian keratins. Ann N Y Acad Sci 455, 403\u2013425 (1985).","journal-title":"Ann N Y Acad Sci"},{"key":"e_1_3_4_43_2","doi-asserted-by":"crossref","DOI":"10.1007\/978-3-662-00989-5","volume-title":"Biology of the Integument 2 Vertebrates","author":"Bereiter-Hahn J","year":"1986","unstructured":"J Bereiter-Hahn, AG Matoltsy, KS Richards Biology of the Integument 2 Vertebrates (Springer, Berlin, 1986)."},{"key":"e_1_3_4_44_2","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1002\/aja.1001960411","article-title":"Region-specific patterns of beta keratin expression during avian skin development","volume":"196","author":"Knapp LW","year":"1993","unstructured":"LW Knapp, RB Shames, GL Barnes, RH Sawyer, Region-specific patterns of beta keratin expression during avian skin development. Dev Dyn 196, 283\u2013290 (1993).","journal-title":"Dev Dyn"},{"key":"e_1_3_4_45_2","doi-asserted-by":"crossref","first-page":"549","DOI":"10.1016\/0022-2836(89)90593-7","article-title":"Avian keratin genes. I. A molecular analysis of the structure and expression of a group of feather keratin genes","volume":"209","author":"Presland RB","year":"1989","unstructured":"RB Presland, , Avian keratin genes. I. A molecular analysis of the structure and expression of a group of feather keratin genes. J Mol Biol 209, 549\u2013559 (1989).","journal-title":"J Mol Biol"},{"key":"e_1_3_4_46_2","doi-asserted-by":"crossref","first-page":"561","DOI":"10.1016\/0022-2836(89)90594-9","article-title":"Avian keratin genes. II. Chromosomal arrangement and close linkage of three gene families","volume":"209","author":"Presland RB","year":"1989","unstructured":"RB Presland, LA Whitbread, GE Rogers, Avian keratin genes. II. Chromosomal arrangement and close linkage of three gene families. J Mol Biol 209, 561\u2013576 (1989).","journal-title":"J Mol Biol"},{"key":"e_1_3_4_47_2","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1016\/0378-1119(91)90415-8","article-title":"The structure and expression of a gene encoding chick claw keratin","volume":"101","author":"Whitbread LA","year":"1991","unstructured":"LA Whitbread, K Gregg, GE Rogers, The structure and expression of a gene encoding chick claw keratin. Gene 101, 223\u2013229 (1991).","journal-title":"Gene"},{"key":"e_1_3_4_48_2","doi-asserted-by":"crossref","first-page":"560","DOI":"10.1111\/j.1469-7580.2009.01045.x","article-title":"Evolution of hard proteins in the sauropsid integument in relation to the cornification of skin derivatives in amniotes","volume":"214","author":"Alibardi L","year":"2009","unstructured":"L Alibardi, L Dalla Valle, A Nardi, M Toni, Evolution of hard proteins in the sauropsid integument in relation to the cornification of skin derivatives in amniotes. J Anat 214, 560\u2013586 (2009).","journal-title":"J Anat"},{"key":"e_1_3_4_49_2","doi-asserted-by":"crossref","first-page":"1320","DOI":"10.1126\/science.1253451","article-title":"Whole-genome analyses resolve early branches in the tree of life of modern birds","volume":"346","author":"Jarvis ED","year":"2014","unstructured":"ED Jarvis, , Whole-genome analyses resolve early branches in the tree of life of modern birds. Science 346, 1320\u20131331 (2014).","journal-title":"Science"},{"key":"e_1_3_4_50_2","doi-asserted-by":"crossref","first-page":"e1002748","DOI":"10.1371\/journal.pgen.1002748","article-title":"The chicken frizzle feather is due to an \u03b1-keratin (KRT75) mutation that causes a defective rachis","volume":"8","author":"Ng CS","year":"2012","unstructured":"CS Ng, , The chicken frizzle feather is due to an \u03b1-keratin (KRT75) mutation that causes a defective rachis. PLoS Genet 8, e1002748 (2012).","journal-title":"PLoS Genet"},{"key":"e_1_3_4_51_2","doi-asserted-by":"crossref","first-page":"1981","DOI":"10.1016\/j.yexcr.2007.04.007","article-title":"Intermediate filaments: A historical perspective","volume":"313","author":"Oshima RG","year":"2007","unstructured":"RG Oshima, Intermediate filaments: A historical perspective. Exp Cell Res 313, 1981\u20131994 (2007).","journal-title":"Exp Cell Res"},{"key":"e_1_3_4_52_2","doi-asserted-by":"crossref","first-page":"629","DOI":"10.1007\/s12038-007-0062-8","article-title":"Keratins: Markers of cell differentiation or regulators of cell differentiation?","volume":"32","author":"Vaidya MM","year":"2007","unstructured":"MM Vaidya, D Kanojia, Keratins: Markers of cell differentiation or regulators of cell differentiation? J Biosci 32, 629\u2013634 (2007).","journal-title":"J Biosci"},{"key":"e_1_3_4_53_2","doi-asserted-by":"crossref","first-page":"2255","DOI":"10.1016\/j.yexcr.2007.02.030","article-title":"Intermediate filaments: A role in epithelial polarity","volume":"313","author":"Oriolo AS","year":"2007","unstructured":"AS Oriolo, FA Wald, VP Ramsauer, PJ Salas, Intermediate filaments: A role in epithelial polarity. Exp Cell Res 313, 2255\u20132264 (2007).","journal-title":"Exp Cell Res"},{"key":"e_1_3_4_54_2","doi-asserted-by":"crossref","first-page":"pe53","DOI":"10.1126\/stke.3662006pe53","article-title":"Intermediate filaments as signaling platforms","volume":"2006","author":"Pallari HM","year":"2006","unstructured":"HM Pallari, JE Eriksson, Intermediate filaments as signaling platforms. Sci STKE 2006, pe53 (2006).","journal-title":"Sci STKE"},{"key":"e_1_3_4_55_2","doi-asserted-by":"crossref","first-page":"1581","DOI":"10.1101\/gad.1552107","article-title":"Intermediate filament scaffolds fulfill mechanical, organizational, and signaling functions in the cytoplasm","volume":"21","author":"Kim S","year":"2007","unstructured":"S Kim, PA Coulombe, Intermediate filament scaffolds fulfill mechanical, organizational, and signaling functions in the cytoplasm. Genes Dev 21, 1581\u20131597 (2007).","journal-title":"Genes Dev"},{"key":"e_1_3_4_56_2","doi-asserted-by":"crossref","first-page":"565","DOI":"10.1038\/nrm3175","article-title":"Deconstructing the skin: Cytoarchitectural determinants of epidermal morphogenesis","volume":"12","author":"Simpson CL","year":"2011","unstructured":"CL Simpson, DM Patel, KJ Green, Deconstructing the skin: Cytoarchitectural determinants of epidermal morphogenesis. Nat Rev Mol Cell Biol 12, 565\u2013580 (2011).","journal-title":"Nat Rev Mol Cell Biol"},{"key":"e_1_3_4_57_2","doi-asserted-by":"crossref","first-page":"1174","DOI":"10.1053\/jhep.2001.29374","article-title":"Disturbances in hepatic cell-cycle regulation in mice with assembly-deficient keratins 8\/18","volume":"34","author":"Toivola DM","year":"2001","unstructured":"DM Toivola, , Disturbances in hepatic cell-cycle regulation in mice with assembly-deficient keratins 8\/18. Hepatology 34, 1174\u20131183 (2001).","journal-title":"Hepatology"},{"key":"e_1_3_4_58_2","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/j.ceb.2012.10.018","article-title":"The expanding significance of keratin intermediate filaments in normal and diseased epithelia","volume":"25","author":"Pan X","year":"2013","unstructured":"X Pan, RP Hobbs, PA Coulombe, The expanding significance of keratin intermediate filaments in normal and diseased epithelia. Curr Opin Cell Biol 25, 47\u201356 (2013).","journal-title":"Curr Opin Cell Biol"},{"key":"e_1_3_4_59_2","doi-asserted-by":"crossref","first-page":"1763","DOI":"10.1172\/JCI38339","article-title":"Introducing intermediate filaments: From discovery to disease","volume":"119","author":"Eriksson JE","year":"2009","unstructured":"JE Eriksson, , Introducing intermediate filaments: From discovery to disease. J Clin Invest 119, 1763\u20131771 (2009).","journal-title":"J Clin Invest"},{"key":"e_1_3_4_60_2","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1016\/j.tree.2009.12.004","article-title":"Evolutionary stasis: The stable chromosomes of birds","volume":"25","author":"Ellegren H","year":"2010","unstructured":"H Ellegren, Evolutionary stasis: The stable chromosomes of birds. Trends Ecol Evol 25, 283\u2013291 (2010).","journal-title":"Trends Ecol Evol"},{"key":"e_1_3_4_61_2","doi-asserted-by":"crossref","first-page":"923","DOI":"10.1093\/gbe\/evt060","article-title":"Rapid evolution of beta-keratin genes contribute to phenotypic differences that distinguish turtles and birds from other reptiles","volume":"5","author":"Li YI","year":"2013","unstructured":"YI Li, L Kong, CP Ponting, W Haerty, Rapid evolution of beta-keratin genes contribute to phenotypic differences that distinguish turtles and birds from other reptiles. Genome Biol Evol 5, 923\u2013933 (2013).","journal-title":"Genome Biol Evol"},{"key":"e_1_3_4_62_2","doi-asserted-by":"crossref","first-page":"938","DOI":"10.1038\/nrg2482","article-title":"Turning a hobby into a job: How duplicated genes find new functions","volume":"9","author":"Conant GC","year":"2008","unstructured":"GC Conant, KH Wolfe, Turning a hobby into a job: How duplicated genes find new functions. Nat Rev Genet 9, 938\u2013950 (2008).","journal-title":"Nat Rev Genet"},{"key":"e_1_3_4_63_2","doi-asserted-by":"crossref","first-page":"449","DOI":"10.1016\/S0959-437X(00)00111-8","article-title":"Evo-devo of feathers and scales: Building complex epithelial appendages","volume":"10","author":"Chuong CM","year":"2000","unstructured":"CM Chuong, R Chodankar, RB Widelitz, TX Jiang, Evo-devo of feathers and scales: Building complex epithelial appendages. Curr Opin Genet Dev 10, 449\u2013456 (2000).","journal-title":"Curr Opin Genet Dev"},{"key":"e_1_3_4_64_2","doi-asserted-by":"crossref","first-page":"690","DOI":"10.1002\/jez.641","article-title":"The peacock's train (Pavo cristatus and Pavo cristatus mut. alba) I. structure, mechanics, and chemistry of the tail feather coverts","volume":"313","author":"Weiss IM","year":"2010","unstructured":"IM Weiss, HO Kirchner, The peacock's train (Pavo cristatus and Pavo cristatus mut. alba) I. structure, mechanics, and chemistry of the tail feather coverts. J Exp Zool A Ecol Genet Physiol 313, 690\u2013703 (2010).","journal-title":"J Exp Zool A Ecol Genet Physiol"},{"key":"e_1_3_4_65_2","first-page":"395","volume-title":"Molecular Basis of Epithelial Appendage Morphogenesis","author":"Jiang T-X","year":"1998","unstructured":"T-X Jiang, S Stott, RB Widelitz, C-M Chuong, Current methods in the study of avian skin appendages. Molecular Basis of Epithelial Appendage Morphogenesis, ed C-M Chuong (Landes Bioscience, Austin, TX), pp. 395\u2013408 (1998)."},{"key":"e_1_3_4_66_2","doi-asserted-by":"crossref","first-page":"166","DOI":"10.1093\/bioinformatics\/btu638","article-title":"HTSeq--a Python framework to work with high-throughput sequencing data","volume":"31","author":"Anders S","year":"2015","unstructured":"S Anders, PT Pyl, W Huber, HTSeq--a Python framework to work with high-throughput sequencing data. Bioinformatics 31, 166\u2013169 (2015).","journal-title":"Bioinformatics"},{"key":"e_1_3_4_67_2","doi-asserted-by":"crossref","first-page":"R25","DOI":"10.1186\/gb-2010-11-3-r25","article-title":"A scaling normalization method for differential expression analysis of RNA-seq data","volume":"11","author":"Robinson MD","year":"2010","unstructured":"MD Robinson, A Oshlack, A scaling normalization method for differential expression analysis of RNA-seq data. Genome Biol 11, R25 (2010).","journal-title":"Genome Biol"},{"key":"e_1_3_4_68_2","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1093\/bioinformatics\/btp616","article-title":"edgeR: A Bioconductor package for differential expression analysis of digital gene expression data","volume":"26","author":"Robinson MD","year":"2010","unstructured":"MD Robinson, DJ McCarthy, GK Smyth, edgeR: A Bioconductor package for differential expression analysis of digital gene expression data. Bioinformatics 26, 139\u2013140 (2010).","journal-title":"Bioinformatics"},{"key":"e_1_3_4_69_2","doi-asserted-by":"crossref","first-page":"639","DOI":"10.1111\/1523-1747.ep12584254","article-title":"Early events during avian skin appendage regeneration: Dependence on epithelial-mesenchymal interaction and order of molecular reappearance","volume":"107","author":"Chuong CM","year":"1996","unstructured":"CM Chuong, RB Widelitz, S Ting-Berreth, TX Jiang, Early events during avian skin appendage regeneration: Dependence on epithelial-mesenchymal interaction and order of molecular reappearance. J Invest Dermatol 107, 639\u2013646 (1996).","journal-title":"J Invest Dermatol"}],"container-title":["Proceedings of the National Academy of Sciences"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/www.pnas.org\/syndication\/doi\/10.1073\/pnas.1520566112","content-type":"unspecified","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/pnas.org\/doi\/pdf\/10.1073\/pnas.1520566112","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,6,12]],"date-time":"2024-06-12T12:56:16Z","timestamp":1718196976000},"score":1,"resource":{"primary":{"URL":"https:\/\/pnas.org\/doi\/full\/10.1073\/pnas.1520566112"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2015,11,23]]},"references-count":69,"journal-issue":{"issue":"49","published-print":{"date-parts":[[2015,12,8]]}},"alternative-id":["10.1073\/pnas.1520566112"],"URL":"https:\/\/doi.org\/10.1073\/pnas.1520566112","relation":{},"ISSN":["0027-8424","1091-6490"],"issn-type":[{"value":"0027-8424","type":"print"},{"value":"1091-6490","type":"electronic"}],"subject":[],"published":{"date-parts":[[2015,11,23]]},"assertion":[{"value":"2015-11-23","order":2,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}]}}