{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T20:33:56Z","timestamp":1792787636053,"version":"4.5.0"},"reference-count":36,"publisher":"SAGE Publications","license":[{"start":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T00:00:00Z","timestamp":1790726400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"},{"start":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T00:00:00Z","timestamp":1790726400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/journals.sagepub.com\/page\/policies\/text-and-data-mining-license"}],"funder":[{"DOI":"10.13039\/100005648","name":"American Association of Orthodontists Foundation","doi-asserted-by":"publisher","award":["BRA"],"award-info":[{"award-number":["BRA"]}],"id":[{"id":"10.13039\/100005648","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Bethel Musculoskeletal Research Center","award":["Fellowship"],"award-info":[{"award-number":["Fellowship"]}],"id":[{"id":"https:\/\/ror.org\/02729ja77","id-type":"ROR","asserted-by":"crossref"}]},{"name":"King's College London Dental Institute","award":["Studentship"],"award-info":[{"award-number":["Studentship"]}],"id":[{"id":"https:\/\/ror.org\/0220mzb33","id-type":"ROR","asserted-by":"crossref"}]},{"DOI":"10.13039\/100000072","name":"National Institute of Dental and Craniofacial Research","doi-asserted-by":"publisher","award":["K08DE028011"],"award-info":[{"award-number":["K08DE028011"]}],"id":[{"id":"10.13039\/100000072","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["journals.sagepub.com"],"crossmark-restriction":true},"short-container-title":["J Dent Res"],"abstract":"<jats:p>\n                    Cleft palate can be caused by failure of elevation of the palatal shelves over the tongue during development. The physical mechanism of elevation is poorly understood. It is thought that elevation is obstructed by the tongue in a condition known as Pierre Robin sequence (PRS), in which the mandible is smaller (micrognathia) and\/or further back than normal (retrognathia). Some PRS cases are associated with mutations in the regulatory and coding regions of\n                    <jats:italic toggle=\"yes\">SOX9<\/jats:italic>\n                    . Here, we first examined the timing of palatal shelf elevation in mouse embryonic explants. While elevation in vivo occurs between embryonic days E13.5 and E14.5, we found that palate elevation occurred in both E12.5 and E13.5 explants immediately upon dissection, indicating release of elastic prestress by removal of the tongue. These findings suggest that palatal shelves are programmed ab initio to grow horizontally (as they do in birds and most reptiles) but are deflected in mammals by the tongue. Next, to examine this process in utero, we generated a new mt\n                    <jats:italic toggle=\"yes\">Hand2<\/jats:italic>\n                    <jats:sup>Cre<\/jats:sup>\n                    mouse line to delete\n                    <jats:italic toggle=\"yes\">Sox9<\/jats:italic>\n                    or to express diphtheria toxin specifically in the mandibular mesenchyme.\n                    <jats:italic toggle=\"yes\">Sox9<\/jats:italic>\n                    deletion resulted in a hypoplastic, dysmorphic, retrognathic mandible with fully penetrant, complete cleft of the secondary palate. In contrast, most\n                    <jats:italic toggle=\"yes\">R26R<\/jats:italic>\n                    <jats:sup>DTA\/+<\/jats:sup>\n                    ;mt\n                    <jats:italic toggle=\"yes\">Hand2<\/jats:italic>\n                    <jats:sup>Cre<\/jats:sup>\n                    embryos showed a reduced mandible (micrognathia) but also displayed a dramatically smaller tongue (microglossia). With a nonobstructing tongue, these mutants exhibited horizontal palatal shelf growth and formed a normal palate. These results are consistent with a mechanism in which physical obstruction by the tongue is the primary cause of downward development of the palatal shelves, far earlier than previously appreciated, and that normal elevation occurs by the release of elastic prestress. We experimentally demonstrate that in PRS, micrognathia causes cleft palate through tongue malpositioning that prevents release of this palatal shelf prestress.\n                  <\/jats:p>","DOI":"10.1177\/00220345261448467","type":"journal-article","created":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T07:51:11Z","timestamp":1790754671000},"update-policy":"https:\/\/doi.org\/10.1177\/sage-journals-update-policy","source":"Crossref","is-referenced-by-count":0,"title":["Intrinsic Tension Drives Palatal Shelf Reorientation post Tongue Retraction"],"prefix":"10.1177","author":[{"ORCID":"https:\/\/orcid.org\/0009-0003-7824-3571","authenticated-orcid":false,"given":"C.C.","family":"Chen","sequence":"first","affiliation":[{"name":"Department of Orofacial Sciences, University of California San Francisco, School of Dentistry, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4706-5509","authenticated-orcid":false,"given":"S.","family":"Zhang","sequence":"additional","affiliation":[{"name":"Department of Orthodontics and Dentofacial Orthopedics, University of Pittsburgh, School of Dental Medicine, Pittsburgh, PA, USA"},{"name":"Center for Craniofacial Regeneration, University of Pittsburgh, School of Dental Medicine, Pittsburgh, PA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9122-9403","authenticated-orcid":false,"given":"J.D.","family":"Morgan","sequence":"additional","affiliation":[{"name":"Centre for Craniofacial Regeneration and Biology, King\u2019s College London, London, England, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4148-5492","authenticated-orcid":false,"given":"C.","family":"Liu","sequence":"additional","affiliation":[{"name":"Department of Cell and Tissue Biology, University of California San Francisco, School of Dentistry, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-2431-5969","authenticated-orcid":false,"given":"M.A.","family":"Nye","sequence":"additional","affiliation":[{"name":"Department of Orthodontics and Dentofacial Orthopedics, University of Pittsburgh, School of Dental Medicine, Pittsburgh, PA, USA"},{"name":"Center for Craniofacial Regeneration, University of Pittsburgh, School of Dental Medicine, Pittsburgh, PA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"N.","family":"Nguyen","sequence":"additional","affiliation":[{"name":"Department of Orofacial Sciences, University of California San Francisco, School of Dentistry, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-0634-2116","authenticated-orcid":false,"given":"I.","family":"Mangoba","sequence":"additional","affiliation":[{"name":"Department of Orofacial Sciences, University of California San Francisco, School of Dentistry, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"M.M.","family":"Blanco","sequence":"additional","affiliation":[{"name":"Department of Orthodontics and Dentofacial Orthopedics, University of Pittsburgh, School of Dental Medicine, Pittsburgh, PA, USA"},{"name":"Center for Craniofacial Regeneration, University of Pittsburgh, School of Dental Medicine, Pittsburgh, PA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"D.J.","family":"Volzer","sequence":"additional","affiliation":[{"name":"Department of Orthodontics and Dentofacial Orthopedics, University of Pittsburgh, School of Dental Medicine, Pittsburgh, PA, USA"},{"name":"Center for Craniofacial Regeneration, University of Pittsburgh, School of Dental Medicine, Pittsburgh, PA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0800-444X","authenticated-orcid":false,"given":"C.P.","family":"Chen","sequence":"additional","affiliation":[{"name":"Department of Orofacial Sciences, University of California San Francisco, School of Dentistry, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8120-4501","authenticated-orcid":false,"given":"D.L.","family":"Cuylear","sequence":"additional","affiliation":[{"name":"Department of Orofacial Sciences, University of California San Francisco, School of Dentistry, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2014-2757","authenticated-orcid":false,"given":"A.H.","family":"Jheon","sequence":"additional","affiliation":[{"name":"Department of Orthodontics and Dentofacial Orthopedics, University of Pittsburgh, School of Dental Medicine, Pittsburgh, PA, USA"},{"name":"Center for Craniofacial Regeneration, University of Pittsburgh, School of Dental Medicine, Pittsburgh, PA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2857-0315","authenticated-orcid":false,"given":"M.T.","family":"Cobourne","sequence":"additional","affiliation":[{"name":"Centre for Craniofacial Regeneration and Biology, King\u2019s College London, London, England, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6053-8756","authenticated-orcid":false,"given":"J.O.","family":"Bush","sequence":"additional","affiliation":[{"name":"Department of Cell and Tissue Biology, University of California San Francisco, School of Dentistry, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6102-2620","authenticated-orcid":false,"given":"J.B.A.","family":"Green","sequence":"additional","affiliation":[{"name":"Centre for Craniofacial Regeneration and Biology, King\u2019s College London, London, England, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-5920-1549","authenticated-orcid":false,"given":"A.F.","family":"Goodwin","sequence":"additional","affiliation":[{"name":"Department of Orthodontics and Dentofacial Orthopedics, University of Pittsburgh, School of Dental Medicine, Pittsburgh, PA, 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