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R. Soc. Interface."],"published-print":{"date-parts":[[2018,6]]},"abstract":"<jats:p>Animal horns play an important role during intraspecific combat. This work investigates the microstructure and mechanical properties of horns from four representative ruminant species: the bighorn sheep (<jats:italic>Ovis canadensis<\/jats:italic>), domestic sheep (<jats:italic>Ovis aries<\/jats:italic>), mountain goat (<jats:italic>Oreamnos americanus<\/jats:italic>) and pronghorn (<jats:italic>Antilocapra americana<\/jats:italic>), aiming to understand the relation between evolved microstructures and mechanical properties. Microstructural similarity is found where disc-shaped keratin cells attach edge-to-edge along the growth direction of the horn core (longitudinal direction) forming a lamella; multiple lamellae are layered face to face along the impact direction (radial direction, perpendicular to horn core growth direction), forming a wavy pattern surrounding a common feature, the tubules. Differences among species include the number and shape of the tubules, the orientation of aligned lamellae and the shape of keratin cells. Water absorption tests reveal that the pronghorn horn has the largest water-absorbing ability due to the presence of nanopores in the keratin cells. The loading direction (compressive and tensile) and level of hydration vary among the horns from different species. The differences in mechanical properties among species may relate to their different fighting behaviours: high stiffness and strength in mountain goat to support the forces during stabbing; high tensile strength in pronghorn for interlocked pulling; impact energy absorption properties in domestic and bighorn sheep to protect the skull during butting. These design rules based on evolutionary modifications among species can be applied in synthetic materials to meet different mechanical requirements.<\/jats:p>","DOI":"10.1098\/rsif.2018.0093","type":"journal-article","created":{"date-parts":[[2018,6,6]],"date-time":"2018-06-06T07:31:07Z","timestamp":1528270267000},"page":"20180093","update-policy":"https:\/\/doi.org\/10.1098\/crossmark-policy","source":"Crossref","is-referenced-by-count":53,"title":["Microstructure and mechanical properties of different keratinous horns"],"prefix":"10.1098","volume":"15","author":[{"given":"Yuchen","family":"Zhang","sequence":"first","affiliation":[{"name":"Materials Science and Engineering Program, University of California, San Diego, 9500 Gilman Drive, La Jolla, CA 92093, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3023-7610","authenticated-orcid":false,"given":"Wei","family":"Huang","sequence":"additional","affiliation":[{"name":"Materials Science and Engineering Program, University of California, San Diego, 9500 Gilman Drive, La Jolla, CA 92093, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cheryl","family":"Hayashi","sequence":"additional","affiliation":[{"name":"Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"John","family":"Gatesy","sequence":"additional","affiliation":[{"name":"Division of Vertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1442-3472","authenticated-orcid":false,"given":"Joanna","family":"McKittrick","sequence":"additional","affiliation":[{"name":"Materials Science and Engineering Program, University of California, San Diego, 9500 Gilman Drive, La Jolla, CA 92093, USA"},{"name":"Mechanical and Aerospace Engineering, University of California, San Diego, 9500 Gilman Drive, La Jolla, CA 92093, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"175","published-online":{"date-parts":[[2018,6,6]]},"reference":[{"key":"e_1_3_6_2_2","doi-asserted-by":"publisher","DOI":"10.2307\/1382822"},{"key":"e_1_3_6_3_2","first-page":"292","volume-title":"Biomechanics in animal behaviour","author":"Kitchener A","year":"2000"},{"key":"e_1_3_6_4_2","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.ecolsys.39.110707.173502"},{"key":"e_1_3_6_5_2","doi-asserted-by":"publisher","DOI":"10.1007\/s11837-012-0302-8"},{"key":"e_1_3_6_6_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.pmatsci.2015.06.001"},{"key":"e_1_3_6_7_2","doi-asserted-by":"publisher","DOI":"10.1038\/199531a0"},{"key":"e_1_3_6_8_2","doi-asserted-by":"publisher","DOI":"10.1098\/rspa.1995.0075"},{"key":"e_1_3_6_9_2","doi-asserted-by":"publisher","DOI":"10.1080\/14786430410001680935"},{"key":"e_1_3_6_10_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.pmatsci.2007.05.002"},{"key":"e_1_3_6_11_2","doi-asserted-by":"publisher","DOI":"10.1016\/0892-0354(91)90016-6"},{"key":"e_1_3_6_12_2","doi-asserted-by":"publisher","DOI":"10.1016\/S0955-0674(01)00301-5"},{"key":"e_1_3_6_13_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.pmatsci.2012.03.001"},{"key":"e_1_3_6_14_2","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1242\/jeb.130.1.121","article-title":"Functional design of horse hoof keratin: the modulation of mechanical properties through hydration effects","volume":"130","author":"Bertram J","year":"1987","journal-title":"J. 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