{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T16:59:23Z","timestamp":1791392363630,"version":"4.3.3"},"reference-count":84,"publisher":"Wiley","issue":"11","license":[{"start":{"date-parts":[[2025,12,3]],"date-time":"2025-12-03T00:00:00Z","timestamp":1764720000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"},{"start":{"date-parts":[[2025,12,3]],"date-time":"2025-12-03T00:00:00Z","timestamp":1764720000000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/501100003977","name":"Israel Science Foundation","doi-asserted-by":"publisher","award":["2150\/25"],"award-info":[{"award-number":["2150\/25"]}],"id":[{"id":"10.13039\/501100003977","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["anatomypubs.onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["The Anatomical Record"],"published-print":{"date-parts":[[2026,11]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>The lips play a central role in both facial expression and speech, yet their microarchitecture remains largely unexamined across primates. This study presents the first systematic, histological, and morphometric investigation of labial structure in 15 primate species, including humans, non\u2010human great apes, Cercopithecidae, Cebidae, and Lemuridae. We analyzed sagittal lip sections stained with Masson's trichrome, applying deep learning\u2010based segmentation and manual annotations to quantify the relative proportions of muscle, adipose, and connective tissue. We also assessed the morphology of the orbicularis oris muscle (OOm) and the anatomical configuration of direct labial tractors (DLTs). Our results reveal that humans possess a uniquely curved, hook\u2010shaped OOm, a dermis\u2010inserting DLT system segregated from the OOm, and a distinctive tissue profile marked by high adipose and low connective tissue content. These features differ markedly from other primates, which typically exhibit a planar OOm, overlapping DLT architecture, and connective tissue\u2010rich labial composition. Interestingly, Lemuridae share aspects of the human soft tissue profile but lack the muscular specialization. We propose that these derived human traits\u2014especially the curved marginal OOm and anatomically decoupled DLTs\u2014enhance the lips' capacity for fine motor control. These features may have supported the evolution of speech\u2010related articulatory precision. While not providing direct functional evidence, our findings offer a new anatomical framework for understanding the evolution of facial expressivity and speech. This study underscores the value of soft tissue microanatomy in reconstructing the evolutionary pathways of hominin communication.<\/jats:p>","DOI":"10.1002\/ar.70103","type":"journal-article","created":{"date-parts":[[2025,12,3]],"date-time":"2025-12-03T11:21:46Z","timestamp":1764760906000},"page":"3276-3296","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Patterns of interspecific variation in labial microarchitecture among anthropoid primates and the evolution of the hominin lips"],"prefix":"10.1002","volume":"309","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7544-013X","authenticated-orcid":false,"given":"Liat","family":"Rotenstreich","sequence":"first","affiliation":[{"name":"Laboratory for Anatomy and Human Evolution, The Farkas Family Center for Anatomical Research and Education (CARE), Department of Neuroscience, Rappaport Faculty of Medicine Technion\u2014Israel Institute of Technology  Haifa Israel"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kfir","family":"Eliyahu","sequence":"additional","affiliation":[{"name":"Laboratory for Anatomy and Human Evolution, The Farkas Family Center for Anatomical Research and Education (CARE), Department of Neuroscience, Rappaport Faculty of Medicine Technion\u2014Israel Institute of Technology  Haifa Israel"},{"name":"Faculty of Data and Decision Sciences Technion\u2014Israel Institute of Technology  Haifa Israel"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nir","family":"Edery","sequence":"additional","affiliation":[{"name":"Department of Pathology Kimron Veterinary Institute  Bet Dagan Israel"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lior","family":"Moss","sequence":"additional","affiliation":[{"name":"Department of Pathology Kimron Veterinary Institute  Bet Dagan Israel"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Saleem","family":"Eben Bari","sequence":"additional","affiliation":[{"name":"Department of Pathology Kimron Veterinary Institute  Bet Dagan Israel"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0537-4661","authenticated-orcid":false,"given":"Assaf","family":"Marom","sequence":"additional","affiliation":[{"name":"Laboratory for Anatomy and Human Evolution, The Farkas Family Center for Anatomical Research and Education (CARE), Department of Neuroscience, Rappaport Faculty of Medicine Technion\u2014Israel Institute of Technology  Haifa Israel"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2025,12,3]]},"reference":[{"key":"e_1_2_7_2_1","doi-asserted-by":"publisher","DOI":"10.1163\/156853963X00220"},{"key":"e_1_2_7_3_1","doi-asserted-by":"publisher","DOI":"10.1002\/bies.20719"},{"key":"e_1_2_7_4_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.21355"},{"key":"e_1_2_7_5_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.22913"},{"key":"e_1_2_7_6_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0110523"},{"key":"e_1_2_7_7_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.23451"},{"key":"e_1_2_7_8_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.a.10093"},{"key":"e_1_2_7_9_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.23393"},{"key":"e_1_2_7_10_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7580.2009.01113.x"},{"key":"e_1_2_7_11_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7580.2006.00523.x"},{"key":"e_1_2_7_12_1","volume-title":"DeJong's the neurologic examination","author":"Campbell W. W.","year":"2020"},{"key":"e_1_2_7_13_1","doi-asserted-by":"crossref","unstructured":"Chen T. &Guestrin C.(2016).Xgboost: A scalable tree boosting system. arXiv preprint.https:\/\/arxiv.org\/abs\/1603.02754","DOI":"10.1145\/2939672.2939785"},{"key":"e_1_2_7_14_1","volume-title":"The Cat Facial Action Coding System manual (CatFACS)","author":"Correia Caeiro C.","year":"2013"},{"key":"e_1_2_7_15_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0266442"},{"key":"e_1_2_7_16_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0308790"},{"key":"e_1_2_7_17_1","doi-asserted-by":"publisher","DOI":"10.1037\/10001-000"},{"key":"e_1_2_7_18_1","doi-asserted-by":"publisher","DOI":"10.3758\/s13423-016-1130-6"},{"key":"e_1_2_7_19_1","doi-asserted-by":"publisher","DOI":"10.1038\/s42003-023-04945-5"},{"key":"e_1_2_7_20_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7580.2008.00953.x"},{"key":"e_1_2_7_21_1","doi-asserted-by":"publisher","DOI":"10.1201\/b14592"},{"key":"e_1_2_7_22_1","doi-asserted-by":"publisher","DOI":"10.1201\/b10299"},{"key":"e_1_2_7_23_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7580.2011.01403.x"},{"key":"e_1_2_7_24_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7580.2009.01111.x"},{"key":"e_1_2_7_25_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajpa.21007"},{"key":"e_1_2_7_26_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajpa.20902"},{"key":"e_1_2_7_27_1","first-page":"349","article-title":"A note on sections of the lips of the primates","volume":"44","author":"Duckworth W.","year":"1910","journal-title":"Journal of Anatomy and Physiology"},{"key":"e_1_2_7_28_1","doi-asserted-by":"publisher","DOI":"10.1037\/0003\u2010066X.48.4.384"},{"key":"e_1_2_7_29_1","volume-title":"Facial action coding system","author":"Ekman P.","year":"1978"},{"key":"e_1_2_7_30_1","doi-asserted-by":"publisher","DOI":"10.1016\/S1364-6613(00)01494-7"},{"key":"e_1_2_7_31_1","volume-title":"Human facial expression: An evolutionary view","author":"Fridlund A. J.","year":"2014"},{"key":"e_1_2_7_32_1","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2009.0142"},{"key":"e_1_2_7_33_1","doi-asserted-by":"publisher","DOI":"10.1002\/aja.1001200207"},{"key":"e_1_2_7_34_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00266-003-3065-3"},{"key":"e_1_2_7_35_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00676.2019"},{"key":"e_1_2_7_36_1","doi-asserted-by":"publisher","DOI":"10.1097\/00006534-199607000-00009"},{"key":"e_1_2_7_37_1","doi-asserted-by":"publisher","DOI":"10.1007\/s002669900113"},{"key":"e_1_2_7_38_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2111876118"},{"key":"e_1_2_7_39_1","doi-asserted-by":"publisher","DOI":"10.1086\/394355"},{"key":"e_1_2_7_40_1","doi-asserted-by":"publisher","DOI":"10.1086\/394364"},{"key":"e_1_2_7_41_1","volume-title":"Evolution of facial musculature and facial expression","author":"Huber E.","year":"1931"},{"key":"e_1_2_7_42_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.25650"},{"key":"e_1_2_7_43_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.wocn.2020.100978"},{"key":"e_1_2_7_44_1","doi-asserted-by":"publisher","DOI":"10.1086\/509092"},{"key":"e_1_2_7_45_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-642-39979-4_79"},{"key":"e_1_2_7_46_1","first-page":"1","article-title":"Facial muscles: The modiolus and muscles surrounding the rima oris with some remarks about the panniculus adiposus","volume":"60","author":"Lightoller G.","year":"1925","journal-title":"Journal of Anatomy"},{"key":"e_1_2_7_47_1","first-page":"19","article-title":"The facial muscles of three orang utans and two cercopithecidae","volume":"63","author":"Lightoller G.","year":"1928","journal-title":"Journal of Anatomy"},{"key":"e_1_2_7_48_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7998.1934.tb07754.x"},{"key":"e_1_2_7_49_1","volume-title":"Junqueira's basic histology: Text and atlas","author":"Mescher A. L.","year":"2018"},{"key":"e_1_2_7_50_1","doi-asserted-by":"publisher","DOI":"10.1097\/00006534-197607000-00013"},{"key":"e_1_2_7_51_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1467-7687.2012.01149.x"},{"key":"e_1_2_7_52_1","first-page":"236","volume-title":"Primate ethology","author":"Moynihan M.","year":"1967"},{"key":"e_1_2_7_53_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajpa.22206"},{"key":"e_1_2_7_54_1","doi-asserted-by":"publisher","DOI":"10.1123\/mcj.15.1.141"},{"key":"e_1_2_7_55_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.25172"},{"key":"e_1_2_7_56_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajpa.21401"},{"key":"e_1_2_7_57_1","first-page":"30","article-title":"The facial muscles of Papio ursinus","volume":"75","author":"Pellat A.","year":"1979","journal-title":"South African Journal of Science"},{"key":"e_1_2_7_58_1","first-page":"436","article-title":"The facial muscles of three African primates contrasted with those of Papio ursinus","volume":"75","author":"Pellat A.","year":"1979","journal-title":"South African Journal of Science"},{"key":"e_1_2_7_59_1","doi-asserted-by":"publisher","DOI":"10.1093\/brain\/60.4.389"},{"key":"e_1_2_7_60_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1001342"},{"key":"e_1_2_7_61_1","volume-title":"The anatomy of the gorilla","author":"Raven H. C.","year":"1950"},{"key":"e_1_2_7_62_1","unstructured":"Ravi N. Gabeur V. Hu Y.\u2010T. Hu R. Ryali C. Ma T. Khedr H. R\u00e4dle R. Rolland C. &Gustafson L.(2024).Sam 2: Segment anything in images and videos. arXiv Preprint.https:\/\/arxiv.org\/abs\/2408.00714"},{"key":"e_1_2_7_63_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7580.2008.01004.x"},{"key":"e_1_2_7_64_1","doi-asserted-by":"publisher","DOI":"10.1096\/fj.202300927R"},{"key":"e_1_2_7_65_1","volume-title":"Morphologisches Jahrbuch: Zeitschrift F\u00fcr Anatomie Und Entwicklungsgeschichte","author":"Ruge G.","year":"1876"},{"key":"e_1_2_7_66_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.aanat.2018.11.004"},{"key":"e_1_2_7_67_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.aanat.2023.152161"},{"key":"e_1_2_7_68_1","doi-asserted-by":"publisher","DOI":"10.1002\/ca.23355"},{"key":"e_1_2_7_69_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.aanat.2018.02.007"},{"key":"e_1_2_7_70_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.aanat.2019.151414"},{"key":"e_1_2_7_71_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajpa.20001"},{"key":"e_1_2_7_72_1","doi-asserted-by":"publisher","DOI":"10.1136\/jnnp.56.4.329"},{"key":"e_1_2_7_73_1","volume-title":"Oral anatomy","author":"Sicher H.","year":"1975"},{"key":"e_1_2_7_74_1","volume-title":"Gray's anatomy e\u2010book","author":"Standring S.","year":"2021"},{"key":"e_1_2_7_75_1","doi-asserted-by":"publisher","DOI":"10.1044\/1092-4388(2012\/12-0200)"},{"key":"e_1_2_7_76_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.1090290303"},{"key":"e_1_2_7_77_1","doi-asserted-by":"publisher","DOI":"10.1186\/s13227-018-0103-4"},{"key":"e_1_2_7_78_1","first-page":"9","volume-title":"Social evolution in primates: The role of ecological factors and male behaviour","author":"Schaik C. P.","year":"1996"},{"key":"e_1_2_7_79_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10919-006-0017-z"},{"key":"e_1_2_7_80_1","volume-title":"DogFACS: The dog facial action coding system","author":"Waller B. M.","year":"2013"},{"key":"e_1_2_7_81_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neubiorev.2016.09.005"},{"key":"e_1_2_7_82_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0131738"},{"key":"e_1_2_7_83_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.25519"},{"key":"e_1_2_7_84_1","doi-asserted-by":"publisher","DOI":"10.1006\/jpho.1997.0050"},{"key":"e_1_2_7_85_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-1-4757-9930-9_9"}],"container-title":["The Anatomical Record"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/anatomypubs.onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/ar.70103","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/anatomypubs.onlinelibrary.wiley.com\/doi\/full-xml\/10.1002\/ar.70103","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/anatomypubs.onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/ar.70103","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T16:45:02Z","timestamp":1791391502000},"score":1,"resource":{"primary":{"URL":"https:\/\/anatomypubs.onlinelibrary.wiley.com\/doi\/10.1002\/ar.70103"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,12,3]]},"references-count":84,"journal-issue":{"issue":"11","published-print":{"date-parts":[[2026,11]]}},"alternative-id":["10.1002\/ar.70103"],"URL":"https:\/\/doi.org\/10.1002\/ar.70103","archive":["Portico"],"relation":{},"ISSN":["1932-8486","1932-8494"],"issn-type":[{"value":"1932-8486","type":"print"},{"value":"1932-8494","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,12,3]]},"assertion":[{"value":"2025-08-11","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2025-11-13","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2025-12-03","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}]}}