{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,27]],"date-time":"2026-05-27T11:06:42Z","timestamp":1779880002586,"version":"3.53.1"},"reference-count":44,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2025,3,1]],"date-time":"2025-03-01T00:00:00Z","timestamp":1740787200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2025,3,1]],"date-time":"2025-03-01T00:00:00Z","timestamp":1740787200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,3,1]],"date-time":"2026-03-01T00:00:00Z","timestamp":1772323200000},"content-version":"vor","delay-in-days":365,"URL":"http:\/\/www.elsevier.com\/open-access\/userlicense\/1.0\/"},{"start":{"date-parts":[[2025,3,1]],"date-time":"2025-03-01T00:00:00Z","timestamp":1740787200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-017"},{"start":{"date-parts":[[2025,3,1]],"date-time":"2025-03-01T00:00:00Z","timestamp":1740787200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2025,3,1]],"date-time":"2025-03-01T00:00:00Z","timestamp":1740787200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-012"},{"start":{"date-parts":[[2025,3,1]],"date-time":"2025-03-01T00:00:00Z","timestamp":1740787200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2025,3,1]],"date-time":"2025-03-01T00:00:00Z","timestamp":1740787200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-004"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01-AR067203"],"award-info":[{"award-number":["R01-AR067203"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01-AR081081"],"award-info":[{"award-number":["R01-AR081081"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01-AR083352"],"award-info":[{"award-number":["R01-AR083352"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Developmental Biology"],"published-print":{"date-parts":[[2025,3]]},"DOI":"10.1016\/j.ydbio.2024.12.012","type":"journal-article","created":{"date-parts":[[2024,12,19]],"date-time":"2024-12-19T02:56:27Z","timestamp":1734576987000},"page":"96-105","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":2,"special_numbering":"C","title":["Mesenchymal cell contractility regulates villus morphogenesis and intestinal architecture"],"prefix":"10.1016","volume":"519","author":[{"given":"Taylor D.","family":"Hinnant","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Caroline","family":"Joo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Terry","family":"Lechler","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.ydbio.2024.12.012_bib1","doi-asserted-by":"crossref","first-page":"297","DOI":"10.1159\/000242668","article-title":"Cell proliferation in developing human jejunum","volume":"51","author":"Arsenault","year":"2009","journal-title":"Biol. Neonate"},{"key":"10.1016\/j.ydbio.2024.12.012_bib2","doi-asserted-by":"crossref","first-page":"15020","DOI":"10.1021\/bi061566o","article-title":"Importance of force linkage in mechanochemistry of adhesion receptors","volume":"45","author":"Astrof","year":"2006","journal-title":"Biochemistry"},{"key":"10.1016\/j.ydbio.2024.12.012_bib3","doi-asserted-by":"crossref","DOI":"10.1242\/dev.201251","article-title":"Maf family transcription factors are required for nutrient uptake in the mouse neonatal gut","volume":"149","author":"Bara","year":"2022","journal-title":"Development"},{"key":"10.1016\/j.ydbio.2024.12.012_bib4","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1016\/j.devcel.2017.08.019","article-title":"YAP\/TAZ and hedgehog coordinate growth and patterning in gastrointestinal mesenchyme","volume":"43","author":"Cotton","year":"2017","journal-title":"Dev. Cell"},{"key":"10.1016\/j.ydbio.2024.12.012_bib5","doi-asserted-by":"crossref","first-page":"1315","DOI":"10.1016\/j.stem.2024.06.012","article-title":"Construction of multilayered small intestine-like tissue by reproducing interstitial flow","volume":"31","author":"Deguchi","year":"2024","journal-title":"Cell Stem Cell"},{"key":"10.1016\/j.ydbio.2024.12.012_bib6","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1038\/nature10137","article-title":"Role of YAP\/TAZ in mechanotransduction","volume":"474","author":"Dupont","year":"2011","journal-title":"Nature"},{"key":"10.1016\/j.ydbio.2024.12.012_bib7","doi-asserted-by":"crossref","first-page":"918","DOI":"10.1039\/C6IB00046K","article-title":"Coordination of signaling and tissue mechanics during morphogenesis of murine intestinal villi: a role for mitotic cell rounding","volume":"8","author":"Freddo","year":"2016","journal-title":"Integr. Biol."},{"key":"10.1016\/j.ydbio.2024.12.012_bib8","doi-asserted-by":"crossref","first-page":"1543","DOI":"10.1053\/j.gastro.2013.02.037","article-title":"Hippo signaling regulates differentiation and maintenance in the exocrine pancreas","volume":"144","author":"Gao","year":"2013","journal-title":"Gastroenterology"},{"key":"10.1016\/j.ydbio.2024.12.012_bib9","doi-asserted-by":"crossref","first-page":"1684","DOI":"10.1126\/science.1093587","article-title":"De novo crypt formation and juvenile polyposis on BMP inhibition in mouse intestine","volume":"303","author":"Haramis","year":"2004","journal-title":"Science"},{"key":"10.1016\/j.ydbio.2024.12.012_bib10","doi-asserted-by":"crossref","first-page":"1117","DOI":"10.1038\/ng1430","article-title":"BMP signaling inhibits intestinal stem cell self-renewal through suppression of Wnt\u2013\u03b2-catenin signaling","volume":"36","author":"He","year":"2004","journal-title":"Nat. Genet."},{"key":"10.1016\/j.ydbio.2024.12.012_bib11","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pgen.1010899","article-title":"Compartment specific responses to contractility in the small intestinal epithelium","volume":"20","author":"Hinnant","year":"2024","journal-title":"PLoS Genet."},{"key":"10.1016\/j.ydbio.2024.12.012_bib12","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1016\/j.devcel.2017.12.004","article-title":"Engineered tissue folding by mechanical compaction of the mesenchyme","volume":"44","author":"Hughes","year":"2018","journal-title":"Dev. Cell"},{"key":"10.1016\/j.ydbio.2024.12.012_bib13","doi-asserted-by":"crossref","DOI":"10.1016\/j.taap.2021.115669","article-title":"Selectivity of the collagen-binding integrin inhibitors, TC-I-15 and obtustatin","volume":"428","author":"Hunter","year":"2021","journal-title":"Toxicol. Appl. Pharmacol."},{"key":"10.1016\/j.ydbio.2024.12.012_bib14","doi-asserted-by":"crossref","first-page":"3072","DOI":"10.1016\/j.cell.2024.04.039","article-title":"Patterning and folding of intestinal villi by active mesenchymal dewetting","volume":"187","author":"Huycke","year":"2024","journal-title":"Cell"},{"key":"10.1016\/j.ydbio.2024.12.012_bib15","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1016\/j.mayocp.2017.10.025","article-title":"Not all that flattens villi is celiac disease: a review of enteropathies","volume":"93","author":"Jansson-Knodell","year":"2018","journal-title":"Mayo Clin. Proc."},{"key":"10.1016\/j.ydbio.2024.12.012_bib16","doi-asserted-by":"crossref","first-page":"3457","DOI":"10.1242\/dev.127.16.3457","article-title":"Abnormal gastrointestinal development in PDGF-A and PDGFR-(alpha) deficient mice implicates a novel mesenchymal structure with putative instructive properties in villus morphogenesis","volume":"127","author":"Karlsson","year":"2000","journal-title":"Development"},{"key":"10.1016\/j.ydbio.2024.12.012_bib17","doi-asserted-by":"crossref","first-page":"893","DOI":"10.1053\/j.gastro.2010.06.014","article-title":"Indian hedgehog regulates intestinal stem cell fate through Epithelial\u2212Mesenchymal interactions during development","volume":"139","author":"Kosinski","year":"2010","journal-title":"Gastroenterology"},{"key":"10.1016\/j.ydbio.2024.12.012_bib18","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1016\/j.stem.2023.03.004","article-title":"Graded BMP signaling within intestinal crypt architecture directs self-organization of the Wnt-secreting stem cell niche","volume":"30","author":"Kraiczy","year":"2023","journal-title":"Cell Stem Cell"},{"key":"10.1016\/j.ydbio.2024.12.012_bib19","doi-asserted-by":"crossref","first-page":"705","DOI":"10.1126\/science.aau3429","article-title":"Active cell migration is critical for steady-state epithelial turnover in the gut","volume":"365","author":"Krndija","year":"2019","journal-title":"Science"},{"key":"10.1016\/j.ydbio.2024.12.012_bib20","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.36865","article-title":"Pdgfra marks a cellular lineage with distinct contributions to myofibroblasts in lung maturation and injury response","volume":"7","author":"Li","year":"2018","journal-title":"Elife"},{"key":"10.1016\/j.ydbio.2024.12.012_bib21","doi-asserted-by":"crossref","first-page":"279","DOI":"10.1242\/dev.01576","article-title":"Epithelial hedgehog signals pattern the intestinal crypt-villus axis","volume":"132","author":"Madison","year":"2005","journal-title":"Development"},{"key":"10.1016\/j.ydbio.2024.12.012_bib22","doi-asserted-by":"crossref","DOI":"10.1038\/s41467-022-28369-7","article-title":"Mesenchymal-epithelial crosstalk shapes intestinal regionalisation via Wnt and Shh signalling","volume":"13","author":"Maimets","year":"2022","journal-title":"Nat. Commun."},{"key":"10.1016\/j.ydbio.2024.12.012_bib23","doi-asserted-by":"crossref","first-page":"391","DOI":"10.1016\/j.stem.2020.01.008","article-title":"Distinct mesenchymal cell populations generate the essential intestinal BMP signaling gradient","volume":"26","author":"McCarthy","year":"2020","journal-title":"Cell Stem Cell"},{"key":"10.1016\/j.ydbio.2024.12.012_bib24","doi-asserted-by":"crossref","first-page":"550","DOI":"10.1016\/j.devcel.2023.02.012","article-title":"Smooth muscle contributes to the development and function of a layered intestinal stem cell niche","volume":"58","author":"McCarthy","year":"2023","journal-title":"Dev. Cell"},{"key":"10.1016\/j.ydbio.2024.12.012_bib25","first-page":"719","volume":"vol. 106","author":"Miller","year":"2009"},{"key":"10.1016\/j.ydbio.2024.12.012_bib26","doi-asserted-by":"crossref","first-page":"5117","DOI":"10.1038\/onc.2012.5","article-title":"YAP induces malignant mesothelioma cell proliferation by upregulating transcription of cell cycle-promoting genes","volume":"31","author":"Mizuno","year":"2012","journal-title":"Oncogene"},{"key":"10.1016\/j.ydbio.2024.12.012_bib27","doi-asserted-by":"crossref","first-page":"581","DOI":"10.1016\/j.bpg.2012.11.013","article-title":"Diarrhoea due to small bowel diseases","volume":"26","author":"Murray","year":"2012","journal-title":"Best Pract. Res. Clin. Gastroenterol."},{"key":"10.1016\/j.ydbio.2024.12.012_bib28","doi-asserted-by":"crossref","first-page":"436","DOI":"10.1016\/j.stem.2020.11.002","article-title":"Differentiated daughter cells regulate stem cell proliferation and fate through intra-tissue tension","volume":"28","author":"Ning","year":"2021","journal-title":"Cell Stem Cell"},{"key":"10.1016\/j.ydbio.2024.12.012_bib29","doi-asserted-by":"crossref","DOI":"10.1038\/ncomms13119","article-title":"Coordinated integrin activation by actin-dependent force during T-cell migration","volume":"7","author":"Nordenfelt","year":"2016","journal-title":"Nat. Commun."},{"key":"10.1016\/j.ydbio.2024.12.012_bib30","doi-asserted-by":"crossref","first-page":"166","DOI":"10.1016\/j.devcel.2021.12.012","article-title":"Adaptive differentiation promotes intestinal villus recovery","volume":"57","author":"Ohara","year":"2022","journal-title":"Dev. Cell"},{"key":"10.1016\/j.ydbio.2024.12.012_bib31","doi-asserted-by":"crossref","first-page":"1960","DOI":"10.1016\/j.cell.2022.04.023","article-title":"Reciprocal cell-ECM dynamics generate supracellular fluidity underlying spontaneous follicle patterning","volume":"185","author":"Palmquist","year":"2022","journal-title":"Cell"},{"key":"10.1016\/j.ydbio.2024.12.012_bib32","doi-asserted-by":"crossref","first-page":"647","DOI":"10.1016\/j.devcel.2020.02.003","article-title":"Hedgehog-activated Fat4 and PCP pathways mediate mesenchymal cell clustering and villus formation in gut development","volume":"52","author":"Rao-Bhatia","year":"2020","journal-title":"Dev. Cell"},{"key":"10.1016\/j.ydbio.2024.12.012_bib33","doi-asserted-by":"crossref","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":"10.1016\/j.ydbio.2024.12.012_bib34","doi-asserted-by":"crossref","first-page":"910","DOI":"10.1038\/s41596-021-00674-3","article-title":"3D in vitro morphogenesis of human intestinal epithelium in a gut-on-a-chip or a hybrid chip with a cell culture insert","volume":"17","author":"Shin","year":"2022","journal-title":"Nat. Protoc."},{"key":"10.1016\/j.ydbio.2024.12.012_bib35","doi-asserted-by":"crossref","first-page":"242","DOI":"10.1038\/s41586-018-0084-4","article-title":"Subepithelial telocytes are an important source of Wnts that supports intestinal crypts","volume":"557","author":"Shoshkes-Carmel","year":"2018","journal-title":"Nature"},{"key":"10.1016\/j.ydbio.2024.12.012_bib36","doi-asserted-by":"crossref","first-page":"569","DOI":"10.1016\/j.cell.2015.03.041","article-title":"Bending gradients: how the intestinal stem cell gets its home","volume":"161","author":"Shyer","year":"2015","journal-title":"Cell"},{"key":"10.1016\/j.ydbio.2024.12.012_bib37","doi-asserted-by":"crossref","first-page":"1197","DOI":"10.1007\/s12551-022-00997-9","article-title":"Molecular determinants of intrinsic cellular stiffness in health and disease","volume":"14","author":"Su","year":"2022","journal-title":"Biophy. Rev."},{"key":"10.1016\/j.ydbio.2024.12.012_bib38","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1016\/j.devcel.2018.03.024","article-title":"Morphogenesis and compartmentalization of the intestinal crypt","volume":"45","author":"Sumigray","year":"2018","journal-title":"Dev. Cell"},{"key":"10.1016\/j.ydbio.2024.12.012_bib39","doi-asserted-by":"crossref","first-page":"359","DOI":"10.1126\/science.1092436","article-title":"Defining the epithelial stem cell niche in skin","volume":"303","author":"Tumbar","year":"2004","journal-title":"Science"},{"key":"10.1016\/j.ydbio.2024.12.012_bib40","doi-asserted-by":"crossref","first-page":"990","DOI":"10.1038\/40187","article-title":"Calcium sensitization of smooth muscle mediated by a Rho-associated protein kinase in hypertension","volume":"389","author":"Uehata","year":"1997","journal-title":"Nature"},{"key":"10.1016\/j.ydbio.2024.12.012_bib41","doi-asserted-by":"crossref","DOI":"10.1038\/ncomms14396","article-title":"Optogenetic control of cellular forces and mechanotransduction","volume":"8","author":"Valon","year":"2017","journal-title":"Nat. Commun."},{"key":"10.1016\/j.ydbio.2024.12.012_bib42","doi-asserted-by":"crossref","first-page":"15817","DOI":"10.1073\/pnas.1205669109","article-title":"Hedgehog-responsive mesenchymal clusters direct patterning and emergence of intestinal villi","volume":"109","author":"Walton","year":"2012","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.ydbio.2024.12.012_bib43","first-page":"427","article-title":"Villification in the mouse: Bmp signals control intestinal villus patterning","volume":"143","author":"Walton","year":"2015","journal-title":"Development"},{"key":"10.1016\/j.ydbio.2024.12.012_bib44","doi-asserted-by":"crossref","first-page":"665","DOI":"10.1038\/onc.2012.611","article-title":"Stem cell CD44v isoforms promote intestinal cancer formation in Apc(min) mice downstream of Wnt signaling","volume":"33","author":"Zeilstra","year":"2014","journal-title":"Oncogene"}],"container-title":["Developmental Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0012160624002847?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0012160624002847?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,3,1]],"date-time":"2026-03-01T05:20:06Z","timestamp":1772342406000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0012160624002847"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,3]]},"references-count":44,"alternative-id":["S0012160624002847"],"URL":"https:\/\/doi.org\/10.1016\/j.ydbio.2024.12.012","relation":{},"ISSN":["0012-1606"],"issn-type":[{"value":"0012-1606","type":"print"}],"subject":[],"published":{"date-parts":[[2025,3]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Mesenchymal cell contractility regulates villus morphogenesis and intestinal architecture","name":"articletitle","label":"Article Title"},{"value":"Developmental Biology","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.ydbio.2024.12.012","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2024 Elsevier Inc.","name":"copyright","label":"Copyright"}]}}