{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T09:09:18Z","timestamp":1790759358547,"version":"4.1.0"},"reference-count":46,"publisher":"Elsevier BV","issue":"1","license":[{"start":{"date-parts":[[2020,4,1]],"date-time":"2020-04-01T00:00:00Z","timestamp":1585699200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2020,4,1]],"date-time":"2020-04-01T00:00:00Z","timestamp":1585699200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2021,4,6]],"date-time":"2021-04-06T00:00:00Z","timestamp":1617667200000},"content-version":"vor","delay-in-days":370,"URL":"http:\/\/www.elsevier.com\/open-access\/userlicense\/1.0\/"}],"funder":[{"DOI":"10.13039\/100000050","name":"National Heart, Lung, and Blood Institute","doi-asserted-by":"publisher","award":["HL143256"],"award-info":[{"award-number":["HL143256"]}],"id":[{"id":"10.13039\/100000050","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000050","name":"National Heart, Lung, and Blood Institute","doi-asserted-by":"publisher","award":["R01 HL142215"],"award-info":[{"award-number":["R01 HL142215"]}],"id":[{"id":"10.13039\/100000050","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000050","name":"National Heart, Lung, and Blood Institute","doi-asserted-by":"publisher","award":["HL119946"],"award-info":[{"award-number":["HL119946"]}],"id":[{"id":"10.13039\/100000050","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000050","name":"National Heart, Lung, and Blood Institute","doi-asserted-by":"publisher","award":["HL122406"],"award-info":[{"award-number":["HL122406"]}],"id":[{"id":"10.13039\/100000050","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000065","name":"National Institute of Neurological Disorders and Stroke","doi-asserted-by":"publisher","award":["NS047101"],"award-info":[{"award-number":["NS047101"]}],"id":[{"id":"10.13039\/100000065","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["DGE-1747503"],"award-info":[{"award-number":["DGE-1747503"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["-1747503"],"award-info":[{"award-number":["-1747503"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["cell.com","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Developmental Cell"],"published-print":{"date-parts":[[2020,4]]},"DOI":"10.1016\/j.devcel.2020.02.001","type":"journal-article","created":{"date-parts":[[2020,3,9]],"date-time":"2020-03-09T03:41:58Z","timestamp":1583725318000},"page":"73-85.e5","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":62,"title":["Smooth Muscle Differentiation Is Essential for Airway Size, Tracheal Cartilage Segmentation, but Dispensable for Epithelial Branching"],"prefix":"10.1016","volume":"53","author":[{"given":"Randee E.","family":"Young","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mary-Kayt","family":"Jones","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Elizabeth A.","family":"Hines","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rongbo","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yongfeng","family":"Luo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wei","family":"Shi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jamie M.","family":"Verheyden","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xin","family":"Sun","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"key":"10.1016\/j.devcel.2020.02.001_bib46","article-title":"A high throughput in situ hybridization method to characterize mRNA expression patterns in the fetal mouse lower urogenital tract.","volume":"54","author":"Abler","year":"2011","journal-title":"J Vis Exp"},{"key":"10.1016\/j.devcel.2020.02.001_bib43","doi-asserted-by":"crossref","first-page":"W147","DOI":"10.1093\/nar\/gkw419","article-title":"Heatmapper: web-enabled heat mapping for all","volume":"44","author":"Babicki","year":"2016","journal-title":"Nucleic Acid Res"},{"key":"10.1016\/j.devcel.2020.02.001_bib1","doi-asserted-by":"crossref","first-page":"484","DOI":"10.1016\/j.ymthe.2005.02.020","article-title":"Factors influencing adenovirus-mediated airway transduction in fetal mice","volume":"12","author":"Buckley","year":"2005","journal-title":"Mol. Ther."},{"key":"10.1016\/j.devcel.2020.02.001_bib2","doi-asserted-by":"crossref","first-page":"801","DOI":"10.1172\/JCI70291","article-title":"Prenatal retinoid deficiency leads to airway hyperresponsiveness in adult mice","volume":"124","author":"Chen","year":"2014","journal-title":"J.\u00a0Clin. Invest."},{"key":"10.1016\/j.devcel.2020.02.001_bib3","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1242\/dev.163014","article-title":"Niche-mediated BMP\/SMAD signaling regulates lung alveolar stem cell proliferation and differentiation","volume":"145","author":"Chung","year":"2018","journal-title":"Development"},{"key":"10.1016\/j.devcel.2020.02.001_bib40","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1093\/bioinformatics\/bts635","article-title":"STAR: ultrafast universal RNA-seq aligner","volume":"29","author":"Dobin","year":"2013","journal-title":"Bioinformatics"},{"key":"10.1016\/j.devcel.2020.02.001_bib4","doi-asserted-by":"crossref","first-page":"481","DOI":"10.1136\/thoraxjnl-2016-209598","article-title":"Lung Gene expression analysis (LGEA): an integrative web portal for comprehensive gene expression data analysis in lung development","volume":"72","author":"Du","year":"2017","journal-title":"Thorax"},{"key":"10.1016\/j.devcel.2020.02.001_bib5","doi-asserted-by":"crossref","first-page":"4271","DOI":"10.1242\/dev.151266","article-title":"Mechanical regulation of musculoskeletal system development","volume":"144","author":"Felsenthal","year":"2017","journal-title":"Development"},{"key":"10.1016\/j.devcel.2020.02.001_bib6","doi-asserted-by":"crossref","first-page":"dev181172","DOI":"10.1242\/dev.181172","article-title":"Smooth muscle differentiation shapes domain branches during mouse lung development","volume":"146","author":"Goodwin","year":"2019","journal-title":"Development"},{"key":"10.1016\/j.devcel.2020.02.001_bib7","doi-asserted-by":"crossref","first-page":"16287","DOI":"10.1073\/pnas.0902274106","article-title":"\u03b2-catenin promotes respiratory progenitor identity in mouse foregut","volume":"106","author":"Harris-Johnson","year":"2009","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.devcel.2020.02.001_bib45","doi-asserted-by":"crossref","first-page":"610","DOI":"10.1053\/j.gastro.2008.05.032","article-title":"Myosin light chain kinase is central to smooth muscle contraction and required for gastrointestinal motility in mice","volume":"135","author":"He","year":"2008","journal-title":"Gastroenterology"},{"key":"10.1016\/j.devcel.2020.02.001_bib8","doi-asserted-by":"crossref","first-page":"19444","DOI":"10.1073\/pnas.1313223110","article-title":"Establishment of smooth muscle and cartilage juxtaposition in the developing mouse upper airways","volume":"110","author":"Hines","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.devcel.2020.02.001_bib9","first-page":"515","article-title":"Myocardin regulates expression of contractile genes in smooth muscle cells and is required for closure of the ductus arteriosus in mice","volume":"118","author":"Huang","year":"2008","journal-title":"J.\u00a0Clin. Invest."},{"key":"10.1016\/j.devcel.2020.02.001_bib44","doi-asserted-by":"crossref","first-page":"18734","DOI":"10.1073\/pnas.0910749106","article-title":"Myocardin is required for cardiomyocyte survival and maintenance of heart function","volume":"106","author":"Huang","year":"2009","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.devcel.2020.02.001_bib10","doi-asserted-by":"crossref","first-page":"L559","DOI":"10.1152\/ajplung.00498.2005","article-title":"Peristalsis of airway smooth muscle is developmentally regulated and uncoupled from hypoplastic lung growth","volume":"291","author":"Jesudason","year":"2006","journal-title":"Am. J. Physiol. Lung Cell. Mol. Physiol."},{"key":"10.1016\/j.devcel.2020.02.001_bib11","doi-asserted-by":"crossref","first-page":"719","DOI":"10.1016\/j.devcel.2015.08.012","article-title":"Localized smooth muscle differentiation is essential for epithelial bifurcation during branching morphogenesis of the mammalian lung","volume":"34","author":"Kim","year":"2015","journal-title":"Dev. Cell"},{"key":"10.1016\/j.devcel.2020.02.001_bib12","doi-asserted-by":"crossref","first-page":"2816","DOI":"10.1038\/s41467-018-05189-2","article-title":"Synchronized mesenchymal cell polarization and differentiation shape the formation of the murine trachea and esophagus","volume":"9","author":"Kishimoto","year":"2018","journal-title":"Nat. Commun."},{"key":"10.1016\/j.devcel.2020.02.001_bib13","doi-asserted-by":"crossref","first-page":"1093","DOI":"10.1242\/dev.113.4.1093","article-title":"The transcription factor TTF-1 is expressed at the onset of thyroid and lung morphogenesis and in restricted regions of the foetal brain","volume":"113","author":"Lazzaro","year":"1991","journal-title":"Development"},{"key":"10.1016\/j.devcel.2020.02.001_bib14","doi-asserted-by":"crossref","first-page":"1149","DOI":"10.1016\/j.cell.2017.07.028","article-title":"Anatomically and functionally distinct lung mesenchymal populations marked by Lgr5 and Lgr6","volume":"170","author":"Lee","year":"2017","journal-title":"Cell"},{"key":"10.1016\/j.devcel.2020.02.001_bib15","doi-asserted-by":"crossref","first-page":"297","DOI":"10.1016\/j.devcel.2018.01.008","article-title":"The strength of mechanical forces determines the differentiation of alveolar epithelial cells","volume":"44","author":"Li","year":"2018","journal-title":"Dev. Cell"},{"key":"10.1016\/j.devcel.2020.02.001_bib16","doi-asserted-by":"crossref","first-page":"3302","DOI":"10.1158\/0008-5472.CAN-15-0240","article-title":"Itgbl1 is a Runx2 transcriptional target and promotes breast cancer bone metastasis by activating the TGF\u03b2 signaling pathway","volume":"75","author":"Li","year":"2015","journal-title":"Cancer Res"},{"key":"10.1016\/j.devcel.2020.02.001_bib41","doi-asserted-by":"crossref","first-page":"923","DOI":"10.1093\/bioinformatics\/btt656","article-title":"featureCounts: an efficient general purpose program for assigning sequence reads to genomic features","volume":"30","author":"Liao","year":"2014","journal-title":"Bioinformatics"},{"key":"10.1016\/j.devcel.2020.02.001_bib42","doi-asserted-by":"crossref","first-page":"550","DOI":"10.1186\/s13059-014-0550-8","article-title":"Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2","volume":"15","author":"Love","year":"2014","journal-title":"Genome Biol"},{"key":"10.1016\/j.devcel.2020.02.001_bib17","doi-asserted-by":"crossref","first-page":"2157","DOI":"10.1242\/dev.01795","article-title":"FGF10 expression identifies parabronchial smooth muscle cell progenitors and is required for their entry into the smooth muscle cell lineage","volume":"132","author":"Mailleux","year":"2005","journal-title":"Development"},{"key":"10.1016\/j.devcel.2020.02.001_bib18","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1038\/nature07005","article-title":"The\u00a0branching programme of mouse lung development","volume":"453","author":"Metzger","year":"2008","journal-title":"Nature"},{"key":"10.1016\/j.devcel.2020.02.001_bib19","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1016\/j.devcel.2009.12.010","article-title":"Preparing for the first breath: genetic and cellular mechanisms in lung development","volume":"18","author":"Morrisey","year":"2010","journal-title":"Dev. Cell"},{"key":"10.1016\/j.devcel.2020.02.001_bib20","first-page":"219","article-title":"A Gobal double-fluorescent cre reporter mouse","volume":"224","author":"Muzumdar","year":"2006","journal-title":"Genesis"},{"key":"10.1016\/j.devcel.2020.02.001_bib21","first-page":"4328","article-title":"Microfluidic chest cavities reveal that transmural pressure controls the rate of lung development","volume":"144","author":"Nelson","year":"2017","journal-title":"Development"},{"key":"10.1016\/j.devcel.2020.02.001_bib22","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1002\/(SICI)1526-968X(200002)26:2<145::AID-GENE14>3.0.CO;2-C","article-title":"Col2a1-directed expression of Cre recombinase in differentiating chondrocytes in transgenic mice","volume":"26","author":"Ovchinnikov","year":"2000","journal-title":"Genesis"},{"key":"10.1016\/j.devcel.2020.02.001_bib23","doi-asserted-by":"crossref","first-page":"589","DOI":"10.1038\/nature12358","article-title":"Coordination of heart and lung co-development by a multipotent cardiopulmonary progenitor","volume":"500","author":"Peng","year":"2013","journal-title":"Nature"},{"key":"10.1016\/j.devcel.2020.02.001_bib24","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1186\/1471-2164-15-48","article-title":"Identification of mechanosensitive genes during skeletal development: alteration of genes associated with cytoskeletal rearrangement and cell signalling pathways","volume":"15","author":"Rolfe","year":"2014","journal-title":"BMC Genomics"},{"key":"10.1016\/j.devcel.2020.02.001_bib25","doi-asserted-by":"crossref","first-page":"2296","DOI":"10.1210\/en.2005-1020","article-title":"Runx2 is a target of mechanical unloading to alter osteoblastic activity and bone formation in\u00a0vivo","volume":"147","author":"Salingcarnboriboon","year":"2006","journal-title":"Endocrinology"},{"key":"10.1016\/j.devcel.2020.02.001_bib26","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1165\/ajrcmb.23.1.3926","article-title":"Spontaneous peristaltic airway contractions propel lung liquid through the bronchial tree of intact and fetal lung explants","volume":"23","author":"Schittny","year":"2000","journal-title":"Am. J. Respir. Cell Mol. Biol."},{"key":"10.1016\/j.devcel.2020.02.001_bib39","doi-asserted-by":"crossref","first-page":"671","DOI":"10.1038\/nmeth.2089","article-title":"NIH Image to ImageJ: 25 years of image analysis","volume":"9","author":"Schneider","year":"2012","journal-title":"Nat Methods"},{"key":"10.1016\/j.devcel.2020.02.001_bib27","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1242\/dev.088500","article-title":"Spatial mapping and quantification of developmental branching morphogenesis","volume":"140","author":"Short","year":"2013","journal-title":"Development"},{"key":"10.1016\/j.devcel.2020.02.001_bib28","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1126\/scitranslmed.aam7486","article-title":"Itgbl1 modulates integrin activity to promote cartilage formation and protect against arthritis","volume":"10","author":"Song","year":"2018","journal-title":"Sci. Transl. Med."},{"key":"10.1016\/j.devcel.2020.02.001_bib29","doi-asserted-by":"crossref","first-page":"361","DOI":"10.1016\/S1569-9048(03)00159-9","article-title":"Ontogeny of airway smooth muscle: structure, innervation, myogenesis and function in the fetal lung","volume":"137","author":"Sparrow","year":"2003","journal-title":"Respir. Physiol. Neurobiol."},{"key":"10.1016\/j.devcel.2020.02.001_bib30","first-page":"764","article-title":"BMP signaling and cellular dynamics during regeneration of airway epithelium from basal progenitors","volume":"143","author":"Tadokoro","year":"2016","journal-title":"Development"},{"key":"10.1016\/j.devcel.2020.02.001_bib31","doi-asserted-by":"crossref","first-page":"4409","DOI":"10.1172\/JCI58097","article-title":"Parabronchial smooth muscle constitutes an airway epithelial stem cell niche in the mouse lung after injury","volume":"121","author":"Volckaert","year":"2011","journal-title":"J.\u00a0Clin. Invest."},{"key":"10.1016\/j.devcel.2020.02.001_bib32","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1038\/6016","article-title":"Mice lacking link protein develop dwarfism and craniofacial abnormalities","volume":"21","author":"Watanabe","year":"1999","journal-title":"Nat. Genet."},{"key":"10.1016\/j.devcel.2020.02.001_bib33","doi-asserted-by":"crossref","first-page":"2815","DOI":"10.1038\/s41467-018-05043-5","article-title":"The potassium channel KCNJ13 is essential for smooth muscle cytoskeletal organization during mouse tracheal tubulogenesis","volume":"9","author":"Yin","year":"2018","journal-title":"Nat. Commun."},{"key":"10.1016\/j.devcel.2020.02.001_bib34","doi-asserted-by":"crossref","first-page":"952","DOI":"10.1101\/gad.1174704","article-title":"Runx2 and Runx3 are essential for chondrocyte maturation, and Runx2 regulates limb growth through induction of Indian hedgehog","volume":"18","author":"Yoshida","year":"2004","journal-title":"Genes Dev"},{"key":"10.1016\/j.devcel.2020.02.001_bib35","doi-asserted-by":"crossref","first-page":"1134","DOI":"10.1016\/j.cell.2017.07.034","article-title":"Distinct mesenchymal lineages and niches promote epithelial self-renewal and Myofibrogenesis in the lung","volume":"170","author":"Zepp","year":"2017","journal-title":"Cell"},{"key":"10.1016\/j.devcel.2020.02.001_bib36","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1186\/1741-7007-11-111","article-title":"Spatial-temporal targeting of lung-specific mesenchyme by a Tbx4 enhancer","volume":"11","author":"Zhang","year":"2013","journal-title":"BMC Biol."},{"key":"10.1016\/j.devcel.2020.02.001_bib37","doi-asserted-by":"crossref","first-page":"5522","DOI":"10.1074\/jbc.M109.062836","article-title":"Myosin light chain kinase is necessary for tonic airway smooth muscle contraction","volume":"285","author":"Zhang","year":"2010","journal-title":"J.\u00a0Biol. Chem."},{"key":"10.1016\/j.devcel.2020.02.001_bib38","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1016\/j.abb.2013.12.015","article-title":"Myocardin and smooth muscle differentiation","volume":"543","author":"Zheng","year":"2014","journal-title":"Arch. Biochem. Biophys."}],"container-title":["Developmental Cell"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S1534580720300708?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S1534580720300708?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,5,12]],"date-time":"2026-05-12T20:19:04Z","timestamp":1778617144000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S1534580720300708"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,4]]},"references-count":46,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2020,4]]}},"alternative-id":["S1534580720300708"],"URL":"https:\/\/doi.org\/10.1016\/j.devcel.2020.02.001","relation":{},"ISSN":["1534-5807"],"issn-type":[{"value":"1534-5807","type":"print"}],"subject":[],"published":{"date-parts":[[2020,4]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Smooth Muscle Differentiation Is Essential for Airway Size, Tracheal Cartilage Segmentation, but Dispensable for Epithelial Branching","name":"articletitle","label":"Article Title"},{"value":"Developmental Cell","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.devcel.2020.02.001","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2020 Elsevier Inc.","name":"copyright","label":"Copyright"}]}}