{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,29]],"date-time":"2026-06-29T13:46:38Z","timestamp":1782740798111,"version":"3.54.5"},"reference-count":88,"publisher":"The Company of Biologists","issue":"12","license":[{"start":{"date-parts":[[2026,6,29]],"date-time":"2026-06-29T00:00:00Z","timestamp":1782691200000},"content-version":"vor","delay-in-days":14,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/100000069","name":"National Institute of Arthritis and Musculoskeletal and Skin Diseases","doi-asserted-by":"crossref","award":["R01AR067739"],"award-info":[{"award-number":["R01AR067739"]}],"id":[{"id":"10.13039\/100000069","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100000072","name":"National Institute of Dental and Craniofacial Research","doi-asserted-by":"crossref","award":["F31DE033222"],"award-info":[{"award-number":["F31DE033222"]}],"id":[{"id":"10.13039\/100000072","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100000057","name":"National Institute of General Medical Sciences","doi-asserted-by":"crossref","award":["T32GM145441"],"award-info":[{"award-number":["T32GM145441"]}],"id":[{"id":"10.13039\/100000057","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100008893","name":"University of Iowa","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100008893","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100016025","name":"American Association for Anatomy","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100016025","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,6,15]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>Orofacial clefts, including cleft palate (CP), are among the most common types of birth defects. CP results from a failure of palatal shelf fusion during development. Previous human studies have shown that variants of the RhoA GTPase activating protein 29 (ARHGAP29) gene are linked to CP, yet the role and tissue-specific requirements for ARHGAP29 during palatogenesis remain unknown. Here, we use tissue-specific deletion of Arhgap29 in mice to provide the first direct evidence that ARHGAP29 is essential for palatal elevation and fusion. We demonstrate that ectodermal conditional loss of Arhgap29 induces a significant delay in palatogenesis at embryonic day (E) 14.5 and a significant, complete cleft of the secondary palate at E18.5, which is not observed when Arhgap29 is lost later in development using K14-Cre and K6-Cre. Phenotypic analyses of palatal shelves at E14.5 reveal a disorganized and thicker epithelium. Loss of Arhgap29 increases palatal epithelial cell area and upregulates markers of contractility including alpha-smooth muscle actin and phospho-myosin regulatory light chain, implicating cell morphology and contractility as potential drivers of CP.<\/jats:p>","DOI":"10.1242\/dev.205636","type":"journal-article","created":{"date-parts":[[2026,6,11]],"date-time":"2026-06-11T08:30:23Z","timestamp":1781166623000},"source":"Crossref","is-referenced-by-count":0,"title":["Ectodermal loss of ARHGAP29 alters epithelial morphology and disrupts murine palatogenesis"],"prefix":"10.1242","volume":"153","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4724-3179","authenticated-orcid":false,"given":"Emily","family":"Adelizzi","sequence":"first","affiliation":[{"name":"The University of Iowa 1 Department of Anatomy and Cell Biology , , 51 Newton Road, Iowa City, IA 52245 ,","place":["USA"]},{"name":"The University of Iowa 2 Interdisciplinary Graduate Program in Genetics , , 51 Newton Road, Iowa City, IA 52245 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-7604-9988","authenticated-orcid":false,"given":"Lindsey","family":"Rhea","sequence":"additional","affiliation":[{"name":"The University of Iowa 1 Department of Anatomy and Cell Biology , , 51 Newton Road, Iowa City, IA 52245 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0009-6676-9158","authenticated-orcid":false,"given":"Campbell","family":"Mitvalsky","sequence":"additional","affiliation":[{"name":"The University of Iowa 1 Department of Anatomy and Cell Biology , , 51 Newton Road, Iowa City, IA 52245 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Samuel","family":"Pek","sequence":"additional","affiliation":[{"name":"The University of Iowa 1 Department of Anatomy and Cell Biology , , 51 Newton Road, Iowa City, IA 52245 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bethany","family":"Doolittle","sequence":"additional","affiliation":[{"name":"The University of Iowa 1 Department of Anatomy and Cell Biology , , 51 Newton Road, Iowa City, IA 52245 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5182-8874","authenticated-orcid":false,"given":"Martine","family":"Dunnwald","sequence":"additional","affiliation":[{"name":"The University of Iowa 1 Department of Anatomy and Cell Biology , , 51 Newton Road, Iowa City, IA 52245 ,","place":["USA"]},{"name":"The University of Iowa 2 Interdisciplinary Graduate Program in Genetics , , 51 Newton Road, Iowa City, IA 52245 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2026,6,29]]},"reference":[{"key":"2026062909015445600_DEV205636C1","doi-asserted-by":"publisher","first-page":"2868","DOI":"10.1038\/s41467-023-38526-1","article-title":"Neural crest E-cadherin loss drives cleft lip\/palate by epigenetic modulation via pro-inflammatory gene-environment interaction","volume":"14","author":"Alvizi","year":"2023","journal-title":"Nat. Commun."},{"key":"2026062909015445600_DEV205636C2","doi-asserted-by":"publisher","first-page":"697","DOI":"10.1177\/10556656221135926","article-title":"Damaging mutations in AFDN contribute to risk of nonsyndromic cleft lip with or without cleft palate","volume":"61","author":"Awotoye","year":"2024","journal-title":"Cleft Palate Craniofac. J."},{"key":"2026062909015445600_DEV205636C3","doi-asserted-by":"publisher","first-page":"810","DOI":"10.1161\/CIRCRESAHA.116.309094","article-title":"Rasip1-mediated Rho GTPase signaling regulates blood vessel tubulogenesis via nonmuscle myosin II","volume":"119","author":"Barry","year":"2016","journal-title":"Circ. Res."},{"key":"2026062909015445600_DEV205636C4","doi-asserted-by":"publisher","first-page":"21","DOI":"10.3390\/biology9010021","article-title":"Apoptotic bodies: particular extracellular vesicles involved in intercellular communication","volume":"9","author":"Battistelli","year":"2020","journal-title":"Biology"},{"key":"2026062909015445600_DEV205636C5","doi-asserted-by":"publisher","first-page":"525","DOI":"10.1038\/ng.580","article-title":"A genome-wide association study of cleft lip with and without cleft palate identifies risk variants near MAFB and ABCA4","volume":"42","author":"Beaty","year":"2010","journal-title":"Nat. Genet."},{"key":"2026062909015445600_DEV205636C6","doi-asserted-by":"publisher","first-page":"2840","DOI":"10.1242\/jcs.139246","article-title":"Interferon regulatory factor 6 regulates keratinocyte migration","volume":"127","author":"Biggs","year":"2014","journal-title":"J. Cell Sci."},{"key":"2026062909015445600_DEV205636C7","doi-asserted-by":"publisher","first-page":"3275","DOI":"10.1016\/j.cub.2021.05.021","article-title":"Desmosomes polarize and integrate chemical and mechanical signaling to govern epidermal tissue form and function","volume":"31","author":"Broussard","year":"2021","journal-title":"Curr. Biol."},{"key":"2026062909015445600_DEV205636C8","doi-asserted-by":"publisher","first-page":"231","DOI":"10.1242\/dev.067082","article-title":"Palatogenesis: morphogenetic and molecular mechanisms of secondary palate development","volume":"139","author":"Bush","year":"2012","journal-title":"Development"},{"key":"2026062909015445600_DEV205636C9","doi-asserted-by":"publisher","first-page":"101404","DOI":"10.1016\/j.gendis.2024.101404","article-title":"Arhgap29 deficiency causes EEC like syndrome in mice","volume":"12","author":"Chi","year":"2025","journal-title":"Genes Dis."},{"key":"2026062909015445600_DEV205636C10","doi-asserted-by":"publisher","first-page":"392","DOI":"10.3389\/fphys.2016.00392","article-title":"Mesenchymal remodeling during palatal shelf elevation revealed by extracellular matrix and F-actin expression patterns","volume":"7","author":"Chiquet","year":"2016","journal-title":"Front. Physiol."},{"key":"2026062909015445600_DEV205636C11","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1002\/ar.1091510202","article-title":"Development of the rat palate","volume":"151","author":"Coleman","year":"1965","journal-title":"Anat. Rec."},{"key":"2026062909015445600_DEV205636C12","doi-asserted-by":"publisher","first-page":"106125","DOI":"10.1016\/j.isci.2023.106125","article-title":"GRHL2 and AP2a coordinate early surface ectoderm lineage commitment during development","volume":"26","author":"Collier","year":"2023","journal-title":"iScience"},{"key":"2026062909015445600_DEV205636C13","doi-asserted-by":"publisher","first-page":"22","DOI":"10.1186\/s10020-021-00283-6","article-title":"TGF-\u03b2-induced \u03b1-SMA expression is mediated by C\/EBP\u03b2 acetylation in human alveolar epithelial cells","volume":"27","author":"Ding","year":"2021","journal-title":"Mol. Med."},{"key":"2026062909015445600_DEV205636C14","doi-asserted-by":"publisher","first-page":"167","DOI":"10.1038\/nrg2933","article-title":"Cleft lip and palate: understanding genetic and environmental influences","volume":"12","author":"Dixon","year":"2011","journal-title":"Nat. Rev. Genet."},{"key":"2026062909015445600_DEV205636C15","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1038\/s41467-023-35881-x","article-title":"Annexin A1 is a polarity cue that directs mitotic spindle orientation during mammalian epithelial morphogenesis","volume":"14","author":"Fankhaenel","year":"2023","journal-title":"Nat. Commun."},{"key":"2026062909015445600_DEV205636C16","doi-asserted-by":"publisher","first-page":"732","DOI":"10.1111\/1523-1747.ep12371684","article-title":"Annexin-1 localization in human skin: possible association with cytoskeletal elements in keratinocytes of the stratum spinosum","volume":"101","author":"Fava","year":"1993","journal-title":"J. Invest. Dermatol."},{"key":"2026062909015445600_DEV205636C17","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1242\/dev.103.Supplement.41","article-title":"Palate development","volume":"103","author":"Ferguson","year":"1988","journal-title":"Development"},{"key":"2026062909015445600_DEV205636C18","doi-asserted-by":"publisher","first-page":"522","DOI":"10.1597\/1545-1569_1995_032_0522_dotfap_2.3.co_2","article-title":"Development of the face and palate","volume":"32","author":"Ferguson","year":"1995","journal-title":"Cleft. Palate-Cran. J."},{"key":"2026062909015445600_DEV205636C19","doi-asserted-by":"publisher","first-page":"455","DOI":"10.1016\/S0012-1606(89)80017-X","article-title":"Medial edge epithelium transforms to mesenchyme after embryonic palatal shelves fuse","volume":"131","author":"Fitchett","year":"1989","journal-title":"Dev. Biol."},{"key":"2026062909015445600_DEV205636C20","doi-asserted-by":"publisher","first-page":"6915","DOI":"10.1523\/JNEUROSCI.6087-10.2011","article-title":"Neural crest and ectodermal cells intermix in the nasal placode to give rise to GnRH-1 neurons, sensory neurons, and olfactory ensheathing cells","volume":"31","author":"Forni","year":"2011","journal-title":"J. Neurosci."},{"key":"2026062909015445600_DEV205636C21","doi-asserted-by":"publisher","first-page":"2524","DOI":"10.1093\/hmg\/dds071","article-title":"Tbx1 regulates oral epithelial adhesion and palatal development","volume":"21","author":"Funato","year":"2012","journal-title":"Hum. Mol. Genet."},{"key":"2026062909015445600_DEV205636C22","doi-asserted-by":"publisher","first-page":"237","DOI":"10.1387\/ijdb.10410903","article-title":"Midline fusion in the formation of the secondary palate anticipated by upregulation of keratin K5\/6 and localized expression of vimentin mRNA in medial edge epithelium","volume":"43","author":"Gibbins","year":"1999","journal-title":"Int. J. Dev. Biol."},{"key":"2026062909015445600_DEV205636C23","doi-asserted-by":"publisher","first-page":"1532448","DOI":"10.3389\/fcell.2025.1532448","article-title":"Novel insights into palatal shelf elevation dynamics in normal mouse embryos","volume":"13","author":"Goering","year":"2025","journal-title":"Front. Cell Dev. Biol."},{"key":"2026062909015445600_DEV205636C24","doi-asserted-by":"publisher","first-page":"1509","DOI":"10.1002\/dvdy.21534","article-title":"Mice with an anterior cleft of the palate survive neonatal lethality","volume":"237","author":"Gu","year":"2008","journal-title":"Dev. Dyn."},{"key":"2026062909015445600_DEV205636C25","doi-asserted-by":"publisher","first-page":"272","DOI":"10.3390\/genes14020272","article-title":"Effect of the Rho-Kinase\/ROCK signaling pathway on cytoskeleton components","volume":"14","author":"Guan","year":"2023","journal-title":"Genes"},{"key":"2026062909015445600_DEV205636C26","doi-asserted-by":"publisher","first-page":"845","DOI":"10.1093\/hmg\/ddaa002","article-title":"SPECC1L regulates palate development downstream of IRF6","volume":"29","author":"Hall","year":"2020","journal-title":"Hum. Mol. Genet."},{"key":"2026062909015445600_DEV205636C27","doi-asserted-by":"publisher","first-page":"75","DOI":"10.1016\/j.semcdb.2017.08.021","article-title":"Periderm: life-cycle and function during orofacial and epidermal development","volume":"91","author":"Hammond","year":"2019","journal-title":"Semin. Cell Dev. Biol."},{"key":"2026062909015445600_DEV205636C28","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1002\/dvdy.70105","article-title":"The role of Rho GTPases in facial morphogenesis","author":"Ibrahim","year":"2026","journal-title":"Dev. Dyn."},{"key":"2026062909015445600_DEV205636C29","doi-asserted-by":"publisher","first-page":"1335","DOI":"10.1038\/ng1903","article-title":"Abnormal skin, limb and craniofacial morphogenesis in mice deficient for interferon regulatory factor 6 (Irf6)","volume":"38","author":"Ingraham","year":"2006","journal-title":"Nat. Genet."},{"key":"2026062909015445600_DEV205636C30","doi-asserted-by":"publisher","first-page":"585","DOI":"10.1016\/j.celrep.2018.09.059","article-title":"Single-cell transcriptomics of traced epidermal and hair follicle stem cells reveals rapid adaptations during wound healing","volume":"25","author":"Joost","year":"2018","journal-title":"Cell Rep."},{"key":"2026062909015445600_DEV205636C31","doi-asserted-by":"publisher","first-page":"e1002122","DOI":"10.1371\/journal.pbio.1002122","article-title":"Convergence and extrusion are required for normal fusion of the mammalian secondary palate","volume":"13","author":"Kim","year":"2015","journal-title":"PLoS Biol."},{"key":"2026062909015445600_DEV205636C32","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1126\/science.273.5272.245","article-title":"Regulation of myosin phosphatase by Rho and Rho-associated kinase (Rho-kinase)","volume":"273","author":"Kimura","year":"1996","journal-title":"Science"},{"key":"2026062909015445600_DEV205636C33","doi-asserted-by":"publisher","first-page":"1169","DOI":"10.1002\/dvdy.23800","article-title":"Multi-layered hypertrophied MEE formation by microtubule disruption via GEF-H1\/RhoA\/ROCK signaling pathway","volume":"241","author":"Kitase","year":"2012","journal-title":"Dev. Dyn."},{"key":"2026062909015445600_DEV205636C34","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1387\/ijdb.120117yk","article-title":"Microtubule disassembly prevents palatal fusion and alters regulation of the E-cadherin\/catenin complex","volume":"57","author":"Kitase","year":"2013","journal-title":"Int. J. Dev. Biol."},{"key":"2026062909015445600_DEV205636C35","doi-asserted-by":"publisher","first-page":"2616","DOI":"10.3390\/cells9122616","article-title":"Influence of ARHGAP29 on the invasion of mesenchymal-transformed breast cancer cells","volume":"9","author":"Kolb","year":"2020","journal-title":"Cells"},{"key":"2026062909015445600_DEV205636C36","doi-asserted-by":"publisher","first-page":"285","DOI":"10.1038\/ng985","article-title":"Mutations in IRF6 cause Van der Woude and popliteal pterygium syndromes","volume":"32","author":"Kondo","year":"2002","journal-title":"Nat. Genet."},{"key":"2026062909015445600_DEV205636C37","doi-asserted-by":"publisher","first-page":"713","DOI":"10.2119\/molmed.2016.00041","article-title":"RhoA\/ROCK signaling regulates TGF\u03b2-induced epithelial-mesenchymal transition of lens epithelial cells through MRTF-A","volume":"22","author":"Korol","year":"2016","journal-title":"Mol. Med."},{"key":"2026062909015445600_DEV205636C38","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1016\/bs.ctdb.2015.07.002","article-title":"Cellular and molecular mechanisms of palatogenesis","volume":"115","author":"Lan","year":"2015","journal-title":"Curr. Top. Dev. Biol."},{"key":"2026062909015445600_DEV205636C39","doi-asserted-by":"publisher","first-page":"e28","DOI":"10.1136\/jmg.2005.036442","article-title":"A mutation of the p63 gene in non-syndromic cleft lip","volume":"43","author":"Leoyklang","year":"2006","journal-title":"J. Med. Genet."},{"key":"2026062909015445600_DEV205636C40","doi-asserted-by":"publisher","first-page":"934","DOI":"10.1002\/bdra.23076","article-title":"Expression and mutation analyses implicate ARHGAP29 as the etiologic gene for the cleft lip with or without cleft palate locus identified by genome-wide association on chromosome 1p22","volume":"94","author":"Leslie","year":"2012","journal-title":"Birth Defects Res. A Clin. Mol. Teratol"},{"key":"2026062909015445600_DEV205636C41","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1002\/tera.1420090114","article-title":"Presence of contractile proteins in mouse fetal palate prior to shelf elevation","volume":"9","author":"Lessard","year":"1974","journal-title":"Teratology"},{"key":"2026062909015445600_DEV205636C42","doi-asserted-by":"publisher","first-page":"229","DOI":"10.1016\/j.ydbio.2013.04.026","article-title":"The widely used Wnt1-Cre transgene causes developmental phenotypes by ectopic activation of Wnt signaling","volume":"379","author":"Lewis","year":"2013","journal-title":"Dev. Biol."},{"key":"2026062909015445600_DEV205636C43","doi-asserted-by":"publisher","first-page":"14759","DOI":"10.1038\/ncomms14759","article-title":"Identification of common non-coding variants at 1p22 that are functional for non-syndromic orofacial clefting","volume":"8","author":"Liu","year":"2017","journal-title":"Nat. Commun."},{"key":"2026062909015445600_DEV205636C44","doi-asserted-by":"publisher","first-page":"dev189241","DOI":"10.1242\/dev.189241","article-title":"Disruption of the nectin-afadin complex recapitulates features of the human cleft lip\/palate syndrome CLPED1","volume":"147","author":"Lough","year":"2020","journal-title":"Development"},{"key":"2026062909015445600_DEV205636C45","doi-asserted-by":"publisher","first-page":"341","DOI":"10.1074\/jbc.M005505200","article-title":"Smooth muscle differentiation marker gene expression is regulated by RhoA-mediated actin polymerization","volume":"276","author":"Mack","year":"2001","journal-title":"J. Biol. Chem."},{"key":"2026062909015445600_DEV205636C46","doi-asserted-by":"publisher","first-page":"343","DOI":"10.1006\/dbio.2000.9644","article-title":"Medial edge epithelial cell fate during palatal fusion","volume":"220","author":"Martinez-Alvarez","year":"2000","journal-title":"Dev. Biol."},{"key":"2026062909015445600_DEV205636C47","doi-asserted-by":"publisher","first-page":"F911","DOI":"10.1152\/ajprenal.00183.2002","article-title":"Central role for Rho in TGF-\u03b21-induced \u03b1-smooth muscle actin expression during epithelial-mesenchymal transition","volume":"284","author":"Masszi","year":"2003","journal-title":"Am. J. Physiol. Renal. Physiol."},{"key":"2026062909015445600_DEV205636C48","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1080\/21541248.2016.1276999","article-title":"Molecular control of Rab activity by GEFs, GAPs and GDI","volume":"9","author":"M\u00fcller","year":"2018","journal-title":"Small GTPases"},{"key":"2026062909015445600_DEV205636C49","doi-asserted-by":"publisher","first-page":"26226","DOI":"10.1038\/srep26226","article-title":"Cell-cell contact and matrix adhesion promote \u03b1SMA expression during TGF\u03b21-induced epithelial-myofibroblast transition via Notch and MRTF-A","volume":"6","author":"O'Connor","year":"2016","journal-title":"Sci. Rep."},{"key":"2026062909015445600_DEV205636C50","doi-asserted-by":"publisher","first-page":"713","DOI":"10.1002\/dvdy.573","article-title":"Single cell sequencing of the mouse anterior palate reveals mesenchymal heterogeneity","volume":"252","author":"Ozekin","year":"2023","journal-title":"Dev. Dyn."},{"key":"2026062909015445600_DEV205636C51","doi-asserted-by":"publisher","first-page":"1298","DOI":"10.1177\/0022034517726079","article-title":"ARHGAP29 mutation is associated with abnormal oral epithelial adhesions","volume":"96","author":"Paul","year":"2017","journal-title":"J. Dent. Res."},{"key":"2026062909015445600_DEV205636C52","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1016\/j.ajhg.2013.11.009","article-title":"Dominant mutations in GRHL3 cause Van der Woude Syndrome and disrupt oral periderm development","volume":"94","author":"Peyrard-Janvid","year":"2014","journal-title":"Am. J. Hum. Genet."},{"key":"2026062909015445600_DEV205636C53","doi-asserted-by":"publisher","first-page":"11427","DOI":"10.1073\/pnas.1306595110","article-title":"Rasip1 mediates Rap1 regulation of Rho in endothelial barrier function through ArhGAP29","volume":"110","author":"Post","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2026062909015445600_DEV205636C54","doi-asserted-by":"publisher","first-page":"RP100961","DOI":"10.7554\/eLife.100961","article-title":"Molecular and mechanical signatures contributing to mouse epidermal differentiation and barrier formation","volume":"13","author":"Prado-Mantilla","year":"2025","journal-title":"eLife"},{"key":"2026062909015445600_DEV205636C55","doi-asserted-by":"publisher","first-page":"1495","DOI":"10.1016\/j.celrep.2017.04.075","article-title":"YAP regulates actin dynamics through ARHGAP29 and promotes metastasis","volume":"19","author":"Qiao","year":"2017","journal-title":"Cell Rep."},{"key":"2026062909015445600_DEV205636C56","doi-asserted-by":"publisher","first-page":"73","DOI":"10.1597\/10-178","article-title":"Genetics of nonsyndromic orofacial clefts","volume":"49","author":"Rahimov","year":"2012","journal-title":"Cleft Palate Craniofac. J."},{"key":"2026062909015445600_DEV205636C57","doi-asserted-by":"publisher","first-page":"310","DOI":"10.1002\/dvdy.759","article-title":"ARHGAP29 promotes keratinocyte proliferation and migration in vitro and is dispensable for in vivo wound healing","volume":"254","author":"Rhea","year":"2025","journal-title":"Dev. Dyn."},{"key":"2026062909015445600_DEV205636C58","doi-asserted-by":"publisher","first-page":"949","DOI":"10.1042\/BST20220461","article-title":"SPECC1L: a cytoskeletal protein that regulates embryonic tissue dynamics","volume":"51","author":"Saadi","year":"2023","journal-title":"Biochem. Soc. Trans."},{"key":"2026062909015445600_DEV205636C59","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1007\/978-3-030-67171-6_4","article-title":"Apoptotic bodies: mechanism of formation, isolation and functional relevance","volume":"97","author":"Santavanond","year":"2021","journal-title":"Subcell. Biochem."},{"key":"2026062909015445600_DEV205636C60","doi-asserted-by":"publisher","first-page":"24333","DOI":"10.1074\/jbc.272.39.24333","article-title":"A novel GTPase-activating protein for Rho interacts with a PDZ domain of the protein-tyrosine phosphatase PTPL1","volume":"272","author":"Saras","year":"1997","journal-title":"J. Biol. Chem."},{"key":"2026062909015445600_DEV205636C61","doi-asserted-by":"publisher","first-page":"459","DOI":"10.1177\/00220345221146454","article-title":"Periderm fate during palatogenesis: TGF-\u03b2 and periderm dedifferentiation","volume":"102","author":"Saroya","year":"2023","journal-title":"J. Dent. Res."},{"key":"2026062909015445600_DEV205636C62","doi-asserted-by":"publisher","first-page":"11208","DOI":"10.1038\/s41598-018-29681-3","article-title":"Runx1-Stat3-Tgfb3 signaling network regulating the anterior palatal development","volume":"8","author":"Sarper","year":"2018","journal-title":"Sci. Rep."},{"key":"2026062909015445600_DEV205636C63","doi-asserted-by":"publisher","first-page":"683","DOI":"10.1111\/cge.12823","article-title":"Impact of rare variants in ARHGAP29 to the etiology of oral clefts: role of loss-of-function vs missense variants","volume":"91","author":"Savastano","year":"2017","journal-title":"Clin. Genet."},{"key":"2026062909015445600_DEV205636C64","doi-asserted-by":"publisher","first-page":"676","DOI":"10.1038\/nmeth.2019","article-title":"Fiji: an open-source platform for biological-image analysis","volume":"9","author":"Schindelin","year":"2012","journal-title":"Nat. Methods"},{"key":"2026062909015445600_DEV205636C65","doi-asserted-by":"publisher","first-page":"e0174206","DOI":"10.1371\/journal.pone.0174206","article-title":"A tissue-specific role for intraflagellar transport genes during craniofacial development","volume":"12","author":"Schock","year":"2017","journal-title":"PLoS ONE"},{"key":"2026062909015445600_DEV205636C66","doi-asserted-by":"publisher","first-page":"1045","DOI":"10.3390\/ijms19041045","article-title":"Annexin A1: uncovering the many talents of an old protein","volume":"19","author":"Sheikh","year":"2018","journal-title":"Int. J. Mol. Sci."},{"key":"2026062909015445600_DEV205636C67","doi-asserted-by":"publisher","first-page":"2735","DOI":"10.3892\/or.2020.7811","article-title":"ARHGAP29 expression may be a novel prognostic factor of cell proliferation and invasion in prostate cancer","volume":"44","author":"Shimizu","year":"2020","journal-title":"Oncol. Rep."},{"key":"2026062909015445600_DEV205636C68","doi-asserted-by":"publisher","first-page":"51","DOI":"10.7150\/ijbs.1.51","article-title":"The GAPs, GEFs, and GDIs of heterotrimeric G-protein alpha subunits","volume":"1","author":"Siderovski","year":"2005","journal-title":"Int. J. Biol. Sci."},{"key":"2026062909015445600_DEV205636C69","doi-asserted-by":"publisher","first-page":"8239258","DOI":"10.1155\/2016\/8239258","article-title":"Annexin A1 and the resolution of inflammation: modulation of neutrophil recruitment, apoptosis, and clearance","volume":"2016","author":"Sugimoto","year":"2016","journal-title":"J. Immunol. Res."},{"key":"2026062909015445600_DEV205636C70","doi-asserted-by":"publisher","first-page":"1159","DOI":"10.1161\/ATVBAHA.118.310991","article-title":"Afadin facilitates vascular endothelial growth factor-induced network formation and migration of vascular endothelial cells by inactivating rho-associated kinase through ArhGAP29","volume":"38","author":"Tagashira","year":"2018","journal-title":"Arterioscler. Thromb. Vasc. Biol."},{"key":"2026062909015445600_DEV205636C71","doi-asserted-by":"publisher","first-page":"2791","DOI":"10.1016\/j.cub.2020.05.032","article-title":"The Mammalian crumbs complex defines a distinct polarity domain apical of epithelial tight junctions","volume":"30","author":"Tan","year":"2020","journal-title":"Curr. Biol."},{"key":"2026062909015445600_DEV205636C72","doi-asserted-by":"publisher","first-page":"2602","DOI":"10.1016\/j.jid.2021.04.008","article-title":"Laminin 332 is indispensable for homeostatic epidermal differentiation programs","volume":"141","author":"Tayem","year":"2021","journal-title":"J. Invest. Dermatol."},{"key":"2026062909015445600_DEV205636C73","doi-asserted-by":"publisher","first-page":"dev200181","DOI":"10.1242\/dev.200181","article-title":"A unique form of collective epithelial migration is crucial for tissue fusion in the secondary palate and can overcome loss of epithelial apoptosis","volume":"149","author":"Teng","year":"2022","journal-title":"Development"},{"key":"2026062909015445600_DEV205636C74","doi-asserted-by":"publisher","first-page":"e202503070","DOI":"10.1083\/jcb.20250307","article-title":"Actomyosin contractility and a threshold of cadherin cell adhesion are required during tissue fusion","volume":"225","author":"Teng","year":"2025","journal-title":"J. Cell Biol."},{"key":"2026062909015445600_DEV205636C75","doi-asserted-by":"publisher","first-page":"273","DOI":"10.1016\/j.ydbio.2008.06.030","article-title":"Facial clefting in Tp63 deficient mice results from altered Bmp4, Fgf8 and Shh signaling","volume":"321","author":"Thomason","year":"2008","journal-title":"Dev. Biol."},{"key":"2026062909015445600_DEV205636C76","doi-asserted-by":"publisher","first-page":"e38972","DOI":"10.1371\/journal.pone.0038972","article-title":"FGFR1-induced epithelial to mesenchymal transition through MAPK\/PLCgamma\/COX-2-mediated mechanisms","volume":"7","author":"Tomlinson","year":"2012","journal-title":"PLoS ONE"},{"key":"2026062909015445600_DEV205636C77","doi-asserted-by":"publisher","first-page":"1032","DOI":"10.1101\/gad.978802","article-title":"Role of Rho family GTPases in epithelial morphogenesis","volume":"16","author":"Van Aelst","year":"2002","journal-title":"Genes Dev."},{"key":"2026062909015445600_DEV205636C78","doi-asserted-by":"publisher","first-page":"76","DOI":"10.1016\/S0955-0674(00)00177-0","article-title":"Actin dynamics and cell-cell adhesion in epithelia","volume":"13","author":"Vasioukhin","year":"2001","journal-title":"Curr. Opin. Cell Biol."},{"key":"2026062909015445600_DEV205636C79","doi-asserted-by":"publisher","first-page":"8551","DOI":"10.1073\/pnas.96.15.8551","article-title":"The magical touch: genome targeting in epidermal stem cells induced by tamoxifen application to mouse skin","volume":"96","author":"Vasioukhin","year":"1999","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2026062909015445600_DEV205636C80","first-page":"176","article-title":"Closure of the secondary palate in three strains of mice","volume":"4","author":"Walker","year":"1956","journal-title":"J. Embryol. Exp. Morph."},{"key":"2026062909015445600_DEV205636C81","doi-asserted-by":"publisher","first-page":"205","DOI":"10.1016\/j.yexcr.2005.11.004","article-title":"Multiple roles of \u03b1-smooth muscle actin in mechanotransduction","volume":"312","author":"Wang","year":"2006","journal-title":"Exp. Cell Res."},{"key":"2026062909015445600_DEV205636C82","doi-asserted-by":"publisher","first-page":"3493","DOI":"10.1111\/odi.14408","article-title":"Mesenchymal \u03b2-catenin signaling affects palatogenesis by regulating \u03b1-actinin-4 and F-actin","volume":"29","author":"Wang","year":"2023","journal-title":"Oral Dis."},{"key":"2026062909015445600_DEV205636C83","first-page":"453","article-title":"The surface layer of human foetal skin and oral mucosa: a study by scanning and transmission electron microscopy","volume":"108","author":"Whittaker","year":"1971","journal-title":"J. Anat."},{"key":"2026062909015445600_DEV205636C84","doi-asserted-by":"publisher","first-page":"526","DOI":"10.1016\/j.devcel.2011.02.010","article-title":"Blood vessel tubulogenesis requires Rasip1 regulation of GTPase signaling","volume":"20","author":"Xu","year":"2011","journal-title":"Dev. Cell"},{"key":"2026062909015445600_DEV205636C85","doi-asserted-by":"publisher","first-page":"463","DOI":"10.1038\/s41419-020-2625-7","article-title":"Annexin-A1 promotes RIG-I-dependent signaling and apoptosis via regulation of the IRF3-IFNAR-STAT1-IFIT1 pathway in A549 lung epithelial cells","volume":"11","author":"Yap","year":"2020","journal-title":"Cell Death Dis."},{"key":"2026062909015445600_DEV205636C86","doi-asserted-by":"publisher","first-page":"4397","DOI":"10.1242\/dev.02013","article-title":"Shox2-deficient mice exhibit a rare type of incomplete clefting of the secondary palate","volume":"132","author":"Yu","year":"2005","journal-title":"Development"},{"key":"2026062909015445600_DEV205636C87","doi-asserted-by":"publisher","first-page":"4647","DOI":"10.3390\/ijms26104647","article-title":"Arhgap29 deficiency directly leads to systemic and craniofacial skeletal abnormalities","volume":"26","author":"Zhang","year":"2025","journal-title":"Int. J. Mol. Sci."},{"key":"2026062909015445600_DEV205636C88","doi-asserted-by":"publisher","first-page":"769","DOI":"10.1056\/NEJMoa032909","article-title":"Interferon regulatory factor 6 (IRF6) gene variants and the risk of isolated cleft lip or palate","volume":"351","author":"Zucchero","year":"2004","journal-title":"N. Engl. J. Med."}],"container-title":["Development"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.biologists.com\/dev\/article-pdf\/doi\/10.1242\/dev.205636\/3703604\/dev205636.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/dev\/article-pdf\/doi\/10.1242\/dev.205636\/3706162\/dev205636.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/dev\/article-pdf\/doi\/10.1242\/dev.205636\/3706162\/dev205636.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,29]],"date-time":"2026-06-29T13:02:04Z","timestamp":1782738124000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/dev\/article\/153\/12\/dev205636\/372102\/Ectodermal-loss-of-ARHGAP29-alters-epithelial"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6,15]]},"references-count":88,"journal-issue":{"issue":"12","published-print":{"date-parts":[[2026,6,15]]}},"URL":"https:\/\/doi.org\/10.1242\/dev.205636","relation":{},"ISSN":["0950-1991","1477-9129"],"issn-type":[{"value":"0950-1991","type":"print"},{"value":"1477-9129","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,6,15]]},"article-number":"dev205636"}}