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These structural parameters were measured in mid diaphyseal undemineralized cross\u2010sections and analyzed using uni\u2010, bi\u2010, and multivariate (principal components analysis) data analysis techniques. We found that the four variables are significantly and positively correlated, and that covariation between variables accounts for as much as 58% of the total variation. These results suggest that torsion is a main determinant of the macro\u2010 and microstructural design of long bones in birds. Humerus, ulna, and femur generally possess torsion\u2010resisting features, while other bones (radius, carpometacarpus, tibiotarsus, tarsometatarsus, and foot phalanx) rather show bending\/axial load\u2010resisting structural properties. These results are congruent with in vivo strain data from the literature, which reported high torsional loading in humerus and ulna during flapping flight, but also in the subhorizontal avian femur during terrestrial locomotion. The precise function of the laminar tissue spatial arrangement, the role of pneumatization, and the influence of flight mode are discussed. \u00a9 2004 Wiley\u2010Liss, Inc.<\/jats:p>","DOI":"10.1002\/ar.a.20141","type":"journal-article","created":{"date-parts":[[2004,12,6]],"date-time":"2004-12-06T16:46:33Z","timestamp":1102351593000},"page":"49-66","source":"Crossref","is-referenced-by-count":115,"title":["Torsional resistance as a principal component of the structural design of long bones: Comparative multivariate evidence in birds"],"prefix":"10.1002","volume":"282A","author":[{"given":"Emmanuel","family":"De Margerie","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sophie","family":"Sanchez","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jorge","family":"Cubo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jacques","family":"Castanet","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2004,12,6]]},"reference":[{"key":"e_1_2_7_2_1","volume-title":"Animal mechanics","author":"Alexander RMN","year":"1968"},{"key":"e_1_2_7_3_1","first-page":"315","article-title":"La structure du tissu osseux envisag\u00e9e comme expression de diff\u00e9rences dans la vitesse de l'accroissement","volume":"58","author":"Amprino R","year":"1947","journal-title":"Arch Biol"},{"key":"e_1_2_7_4_1","first-page":"329","article-title":"La struttura delle ossa nei vertebrati","volume":"9","author":"Amprino R","year":"1947","journal-title":"Pontifica Acad Sci"},{"key":"e_1_2_7_5_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0022-5193(88)80122-X"},{"key":"e_1_2_7_6_1","doi-asserted-by":"publisher","DOI":"10.1016\/0021-9290(91)90374-V"},{"key":"e_1_2_7_7_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.1052250106"},{"key":"e_1_2_7_8_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.10015"},{"key":"e_1_2_7_9_1","first-page":"197","article-title":"Strength of the limb bones of birds and small mammals in bending and twisting: annales des Sciences naturelles","volume":"12","author":"Bou J","year":"1991","journal-title":"Zoologie Paris"},{"key":"e_1_2_7_10_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.b.10031"},{"key":"e_1_2_7_11_1","doi-asserted-by":"publisher","DOI":"10.1017\/S0094837300020108"},{"key":"e_1_2_7_12_1","doi-asserted-by":"publisher","DOI":"10.1002\/(SICI)1097-4687(199906)240:3<237::AID-JMOR3>3.0.CO;2-N"},{"key":"e_1_2_7_13_1","doi-asserted-by":"publisher","DOI":"10.1016\/0021-9290(80)90005-6"},{"key":"e_1_2_7_14_1","doi-asserted-by":"publisher","DOI":"10.1016\/S1095-6433(01)00463-9"},{"key":"e_1_2_7_15_1","first-page":"301","article-title":"Expression de la dynamique de croissance osseuse dans la structure de l'os p\u00e9riostique chez Anas platyrhynchos","volume":"319","author":"Castanet J","year":"1996","journal-title":"C R Acad Sci III Vie"},{"key":"e_1_2_7_16_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0764-4469(00)00181-5"},{"key":"e_1_2_7_17_1","doi-asserted-by":"publisher","DOI":"10.1046\/j.1439-0361.2002.02021.x"},{"key":"e_1_2_7_18_1","doi-asserted-by":"publisher","DOI":"10.1076\/ejom.36.1.19.9029"},{"key":"e_1_2_7_19_1","doi-asserted-by":"publisher","DOI":"10.1076\/0924-3860(200004)38:2;1-F;FT112"},{"key":"e_1_2_7_20_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1096-3642.2000.tb02198.x"},{"key":"e_1_2_7_21_1","first-page":"351","article-title":"Differences in the blood supply of bone of different histological types","volume":"101","author":"Currey JD","year":"1960","journal-title":"Q J Micr Sci"},{"key":"e_1_2_7_22_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7998.1985.tb03551.x"},{"key":"e_1_2_7_23_1","doi-asserted-by":"publisher","DOI":"10.1515\/9781400849505"},{"key":"e_1_2_7_24_1","first-page":"3084","article-title":"Etude de la r\u00e9partition des types de tissus osseux chez les Vert\u00e9br\u00e9s t\u00e9trapodes au moyen de l'analyse factorielle des correspondances","volume":"274","author":"de Bonis L","year":"1972","journal-title":"C R Acad Sci D"},{"key":"e_1_2_7_25_1","doi-asserted-by":"publisher","DOI":"10.1046\/j.1469-7580.2002.00118.x"},{"key":"e_1_2_7_26_1","doi-asserted-by":"publisher","DOI":"10.1016\/S1631-0691(02)01429-4"},{"key":"e_1_2_7_27_1","doi-asserted-by":"publisher","DOI":"10.1242\/jeb.00841"},{"key":"e_1_2_7_28_1","first-page":"51","article-title":"Recherches pal\u00e9ohistologiques sur les os longs des T\u00e9trapodes: VII, sur la classification, la signification fonctionnelle et l'histoire des tissus osseux des T\u00e9trapodes","volume":"61","author":"de Ricql\u00e8s A","year":"1975","journal-title":"Premi\u00e8re partie\u2014structures. 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