{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,3]],"date-time":"2026-03-03T16:03:50Z","timestamp":1772553830493,"version":"3.50.1"},"reference-count":139,"publisher":"MDPI AG","issue":"5","license":[{"start":{"date-parts":[[2026,3,3]],"date-time":"2026-03-03T00:00:00Z","timestamp":1772496000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000024","name":"CIHR","doi-asserted-by":"publisher","award":["186183"],"award-info":[{"award-number":["186183"]}],"id":[{"id":"10.13039\/501100000024","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Cells"],"abstract":"<jats:p>The development of the esophagus and trachea following the septation of the anterior foregut is a highly regulated process involving bidirectional communication between the endoderm and mesoderm. Signaling pathways such as the Bone Morphogenetic Protein family, Wnt\/\u03b2-catenin, Sonic Hedgehog, and Fibroblast Growth Factor family mediate this complex crosstalk to induce the dorsal-ventral patterning of the anterior foregut as well as lineage specification. Even though the mechanisms are not fully understood, dysregulation of signaling pathways may lead to congenital malformations such as tracheomalacia, laryngeal\u2013tracheal clefts and multiple types of esophageal atresia with\/without tracheoesophageal fistula (EA\/TEF). Human induced pluripotent stem cells (iPSCs) provide a robust in vitro platform to monitor the normal and abnormal development of esophagus and trachea and to understand the roles of the endoderm and mesoderm during anterior foregut development. Recent studies have demonstrated that direct differentiation of iPSCs into epithelial and mesenchymal lineages can recapitulate the key stages of foregut development. In this regard, in the current paper, we review the signaling pathways involved in the development of organs deriving from the anterior foregut as well as the roles of the endoderm and mesoderm revealed by previous studies. Furthermore, we discuss the use of iPSCs as a valuable model for investigating the bidirectional communications between the endoderm and mesoderm, which can broaden our knowledge and understanding of the critical mechanisms leading to normal and abnormal development of the esophagus and trachea.<\/jats:p>","DOI":"10.3390\/cells15050448","type":"journal-article","created":{"date-parts":[[2026,3,3]],"date-time":"2026-03-03T12:48:56Z","timestamp":1772542136000},"page":"448","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Induced Pluripotent Stem Cells as a Tool to Decipher the Normal and Abnormal Development of the Esophagus and Trachea from Normal Morphogenesis to Esophageal Atresia, Tracheomalacia, and Laryngo\u2013Tracheal Clefts"],"prefix":"10.3390","volume":"15","author":[{"given":"Yuxuan","family":"Zhang","sequence":"first","affiliation":[{"name":"Esophageal Development and Engineering Laboratory, CHU Sainte-Justine Research Centre, Universit\u00e9 de Montr\u00e9al, 3175 Chemin de la C\u00f4te-Sainte-Catherine, Montr\u00e9al, QC H3T 1C5, Canada"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Anu","family":"David","sequence":"additional","affiliation":[{"name":"Esophageal Development and Engineering Laboratory, CHU Sainte-Justine Research Centre, Universit\u00e9 de Montr\u00e9al, 3175 Chemin de la C\u00f4te-Sainte-Catherine, Montr\u00e9al, QC H3T 1C5, Canada"},{"name":"Department of Pediatrics, Universit\u00e9 de Montr\u00e9al, 3175 Chemin de la C\u00f4te-Sainte-Catherine, Montr\u00e9al, QC H3T 1C5, Canada"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Alireza","family":"Nemati","sequence":"additional","affiliation":[{"name":"Esophageal Development and Engineering Laboratory, CHU Sainte-Justine Research Centre, Universit\u00e9 de Montr\u00e9al, 3175 Chemin de la C\u00f4te-Sainte-Catherine, Montr\u00e9al, QC H3T 1C5, Canada"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2998-8935","authenticated-orcid":false,"given":"Christophe","family":"Faure","sequence":"additional","affiliation":[{"name":"Esophageal Development and Engineering Laboratory, CHU Sainte-Justine Research Centre, Universit\u00e9 de Montr\u00e9al, 3175 Chemin de la C\u00f4te-Sainte-Catherine, Montr\u00e9al, QC H3T 1C5, Canada"},{"name":"Department of Pediatrics, Universit\u00e9 de Montr\u00e9al, 3175 Chemin de la C\u00f4te-Sainte-Catherine, Montr\u00e9al, QC H3T 1C5, Canada"},{"name":"Esophageal Atresia Clinic and Division of Pediatric Gastroenterology Hepatology and Nutrition, CHU Sainte-Justine, 3175 Chemin de la C\u00f4te-Sainte-Catherine, Montr\u00e9al, QC H3T 1C5, Canada"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2026,3,3]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"dev193839","DOI":"10.1242\/dev.193839","article-title":"The development and stem cells of the esophagus","volume":"148","author":"Zhang","year":"2021","journal-title":"Development"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"1611","DOI":"10.1242\/dev.02310","article-title":"Regulation of early lung morphogenesis: Questions, facts and controversies","volume":"133","author":"Cardoso","year":"2006","journal-title":"Development"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"4158","DOI":"10.1038\/s41467-020-17968-x","article-title":"Single cell transcriptomics identifies a signaling network coordinating endoderm and mesoderm diversification during foregut organogenesis","volume":"11","author":"Han","year":"2020","journal-title":"Nat. Commun."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"501","DOI":"10.1016\/j.stem.2018.08.008","article-title":"Esophageal Organoids from Human Pluripotent Stem Cells Delineate Sox2 Functions during Esophageal Specification","volume":"23","author":"Trisno","year":"2018","journal-title":"Cell Stem Cell"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1002\/wdev.179","article-title":"The initial establishment and epithelial morphogenesis of the esophagus: A new model of tracheal\u2013esophageal separation and transition of simple columnar into stratified squamous epithelium in the developing esophagus","volume":"4","author":"Que","year":"2015","journal-title":"WIREs Dev. Biol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1016\/S0344-0338(88)80086-4","article-title":"The Morphology of Ciliogenesis in the Developing Fetal Human Respiratory Epithelium","volume":"183","author":"Moscoso","year":"1988","journal-title":"Pathol.-Res. Pract."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1038\/s41577-020-00477-9","article-title":"Regulation of immune responses by the airway epithelial cell landscape","volume":"21","author":"Hewitt","year":"2021","journal-title":"Nat. Rev. Immunol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1016\/j.devcel.2020.09.024","article-title":"Less Is More: Rare Pulmonary Neuroendocrine Cells Function as Critical Sensors in Lung","volume":"55","author":"Xu","year":"2020","journal-title":"Dev. Cell"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1164\/rccm.200502-203WS","article-title":"The Mysterious Pulmonary Brush Cell","volume":"172","author":"Reid","year":"2005","journal-title":"Am. J. Respir. Crit. Care Med."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1038\/s41586-018-0393-7","article-title":"A revised airway epithelial hierarchy includes CFTR-expressing ionocytes","volume":"560","author":"Montoro","year":"2018","journal-title":"Nature"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"S174","DOI":"10.1164\/ajrccm.157.5.rsaa-1","article-title":"The Development of Large and Small Airways","volume":"157","author":"Jeffery","year":"1998","journal-title":"Am. J. Respir. Crit. Care Med."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1016\/j.devcel.2020.01.033","article-title":"In Vitro and In Vivo Development of the Human Airway at Single-Cell Resolution","volume":"53","author":"Miller","year":"2020","journal-title":"Dev. Cell"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"668","DOI":"10.1002\/ca.23688","article-title":"Development pattern of tracheal cartilage in human embryos","volume":"34","author":"Fockens","year":"2021","journal-title":"Clin. Anat."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1146\/annurev.cellbio.15.1.393","article-title":"Vertebrate Endoderm Development","volume":"15","author":"Wells","year":"1999","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"421","DOI":"10.1177\/000348948409300501","article-title":"Respiratory and Alimentary Relations in Staged Human Embryos:New Embryological Data and Congenital Anomalies","volume":"93","year":"1984","journal-title":"Ann. Otol. Rhinol. Laryngol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"290","DOI":"10.1016\/j.devcel.2009.06.005","article-title":"Wnt2\/2b and \u03b2-Catenin Signaling Are Necessary and Sufficient to Specify Lung Progenitors in the Foregut","volume":"17","author":"Goss","year":"2009","journal-title":"Dev. Cell"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"66","DOI":"10.1016\/j.celrep.2016.05.060","article-title":"A Retinoic Acid-Hedgehog Cascade Coordinates Mesoderm-Inducing Signals and Endoderm Competence during Lung Specification","volume":"16","author":"Rankin","year":"2016","journal-title":"Cell Rep."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"971","DOI":"10.1242\/dev.053694","article-title":"Signaling through BMP receptors promotes respiratory identity in the foregut via repression of Sox2","volume":"138","author":"Domyan","year":"2011","journal-title":"Development"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"474","DOI":"10.1016\/0012-1606(86)90183-1","article-title":"Fetal gut mesenchyme induces differentiation of cultured intestinal endodermal and crypt cells","volume":"113","author":"Kedinger","year":"1986","journal-title":"Dev. Biol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1046\/j.1432-0436.1999.6510013.x","article-title":"In vitro analysis of mesenchymal influences on the differentiation of stomach epithelial cells of the chicken embryo","volume":"65","author":"Koike","year":"1999","journal-title":"Differentiation"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"dmm049541","DOI":"10.1242\/dmm.049541","article-title":"iPSCs derived from esophageal atresia patients reveal SOX2 dysregulation at the anterior foregut stage","volume":"15","author":"Raad","year":"2022","journal-title":"Dis. Models Mech."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"2521","DOI":"10.1242\/dev.003855","article-title":"Multiple dose-dependent roles for Sox2 in the patterning and differentiation of anterior foregut endoderm","volume":"134","author":"Que","year":"2007","journal-title":"Development"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1413","DOI":"10.1093\/hmg\/ddl064","article-title":"Mutations in SOX2 cause anophthalmia-esophageal-genital (AEG) syndrome","volume":"15","author":"Williamson","year":"2006","journal-title":"Hum. Mol. Genet."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"dmm046573","DOI":"10.1242\/dmm.046573","article-title":"Disruption of a hedgehog-foxf1-rspo2 signaling axis leads to tracheomalacia and a loss of sox9+ tracheal chondrocytes","volume":"14","author":"Nasr","year":"2020","journal-title":"Dis. Model. Mech."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1016\/j.ydbio.2021.05.015","article-title":"Developmental basis of trachea-esophageal birth defects","volume":"477","author":"Edwards","year":"2021","journal-title":"Dev. Biol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1314","DOI":"10.1097\/00005537-200308000-00010","article-title":"Complete Laryngotracheoesophageal Cleft: Complicated Management Issues","volume":"113","author":"Chitkara","year":"2003","journal-title":"Laryngoscope"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"26","DOI":"10.1038\/s41572-019-0077-0","article-title":"Oesophageal atresia","volume":"5","author":"Singendonk","year":"2019","journal-title":"Nat. Rev. Dis. Primers"},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Kova\u010devi\u0107, T., Poli\u0107, B., Marki\u0107, J., Ardali\u0107 \u010catipovi\u0107, T., Bucat, M., Mikuli\u0107, S., \u017duvan, L., Pogoreli\u0107, Z., Despot, R., and \u017ditko, V. (2025). Epidemiology and Treatment Outcomes in Neonates with Esophageal Atresia: A 30-Year Population-Based Study. Healthcare, 13.","DOI":"10.3390\/healthcare13040418"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"675","DOI":"10.1016\/j.devcel.2019.11.002","article-title":"Isl1 Regulation of Nkx2.1 in the Early Foregut Epithelium Is Required for Trachea-Esophageal Separation and Lung Lobation","volume":"51","author":"Kim","year":"2019","journal-title":"Dev. Cell"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"1899","DOI":"10.1242\/dev.034629","article-title":"Multiple roles for Sox2 in the developing and adult mouse trachea","volume":"136","author":"Que","year":"2009","journal-title":"Development"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"bio048728","DOI":"10.1242\/bio.048728","article-title":"The absence of SOX2 in the anterior foregut alters the esophagus into trachea and bronchi in both epithelial and mesenchymal components","volume":"9","author":"Teramoto","year":"2020","journal-title":"Biol. Open"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"4005","DOI":"10.1242\/dev.126.18.4005","article-title":"Bmp signaling regulates proximal-distal differentiation of endoderm in mouse lung development","volume":"126","author":"Weaver","year":"1999","journal-title":"Development"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1016\/j.ydbio.2014.02.008","article-title":"BMP antagonism by Noggin is required in presumptive notochord cells for mammalian foregut morphogenesis","volume":"391","author":"Fausett","year":"2014","journal-title":"Dev. Biol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"4159","DOI":"10.1038\/s41467-020-17969-w","article-title":"Bidirectional Wnt signaling between endoderm and mesoderm confers tracheal identity in mouse and human cells","volume":"11","author":"Kishimoto","year":"2020","journal-title":"Nat. Commun."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1016\/j.ydbio.2017.11.018","article-title":"Timing is everything: Reiterative Wnt, BMP and RA signaling regulate developmental competence during endoderm organogenesis","volume":"434","author":"Rankin","year":"2018","journal-title":"Dev. Biol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"4867","DOI":"10.1242\/dev.124.23.4867","article-title":"Fibroblast Growth Factor 10 (FGF10) and branching morphogenesis in the embryonic mouse lung","volume":"124","author":"Bellusci","year":"1997","journal-title":"Development"},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Wang, J.H., Deimling, S.J., D\u2019Alessandro, N.E., Zhao, L., Possmayer, F., and Drysdale, T.A. (2011). Retinoic acid is a key regulatory switch determining the difference between lung and thyroid fates in Xenopus laevis. BMC Dev. Biol., 11.","DOI":"10.1186\/1471-213X-11-75"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"2699","DOI":"10.1038\/s41596-022-00733-3","article-title":"Directed differentiation of human pluripotent stem cells into diverse organ-specific mesenchyme of the digestive and respiratory systems","volume":"17","author":"Kishimoto","year":"2022","journal-title":"Nat. Protoc."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1016\/j.stem.2016.12.007","article-title":"Human iPSC-Derived Cerebral Organoids Model Cellular Features of Lissencephaly and Reveal Prolonged Mitosis of Outer Radial Glia","volume":"20","author":"Bershteyn","year":"2017","journal-title":"Cell Stem Cell"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"445","DOI":"10.1242\/dev.32.2.445","article-title":"Investigation on the origin of the definitive endoderm in the rat embryo","volume":"32","year":"1974","journal-title":"Development"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"891","DOI":"10.1242\/dev.113.3.891","article-title":"Clonal analysis of epiblast fate during germ layer formation in the mouse embryo","volume":"113","author":"Lawson","year":"1991","journal-title":"Development"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1016\/j.stem.2013.12.007","article-title":"Efficient Endoderm Induction from Human Pluripotent Stem Cells by Logically Directing Signals Controlling Lineage Bifurcations","volume":"14","author":"Loh","year":"2014","journal-title":"Cell Stem Cell"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"516","DOI":"10.1016\/j.stem.2018.08.009","article-title":"3D Modeling of Esophageal Development using Human PSC-Derived Basal Progenitors Reveals a Critical Role for Notch Signaling","volume":"23","author":"Zhang","year":"2018","journal-title":"Cell Stem Cell"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"16806","DOI":"10.1073\/pnas.0603916103","article-title":"Wnt and TGF-\u03b2 signaling are required for the induction of an in vitro model of primitive streak formation using embryonic stem cells","volume":"103","author":"Gadue","year":"2006","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"451","DOI":"10.1016\/j.cell.2016.06.011","article-title":"Mapping the Pairwise Choices Leading from Pluripotency to Human Bone, Heart, and Other Mesoderm Cell Types","volume":"166","author":"Loh","year":"2016","journal-title":"Cell"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"aag0053","DOI":"10.1126\/science.aag0053","article-title":"An interactive three-dimensional digital atlas and quantitative database of human development","volume":"354","author":"Hagoort","year":"2016","journal-title":"Science"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"665","DOI":"10.1016\/j.devcel.2019.11.003","article-title":"Endosome-Mediated Epithelial Remodeling Downstream of Hedgehog-Gli Is Required for Tracheoesophageal Separation","volume":"51","author":"Nasr","year":"2019","journal-title":"Dev. Cell"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"2487","DOI":"10.1016\/j.devcel.2025.04.026","article-title":"Disrupted endosomal trafficking of the Vangl-Celsr polarity complex underlies congenital anomalies in Xenopus trachea-esophageal morphogenesis","volume":"60","author":"Edwards","year":"2025","journal-title":"Dev. Cell"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"10643","DOI":"10.1038\/s41467-025-65644-9","article-title":"Convergent flow-mediated mesenchymal force drives embryonic foregut constriction and splitting","volume":"16","author":"Yan","year":"2025","journal-title":"Nat. Commun."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1038\/s41586-018-0394-6","article-title":"A single-cell atlas of the airway epithelium reveals the CFTR-rich pulmonary ionocyte","volume":"560","author":"Plasschaert","year":"2018","journal-title":"Nature"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1016\/j.celrep.2015.06.011","article-title":"Clonal Dynamics Reveal Two Distinct Populations of Basal Cells in Slow-Turnover Airway Epithelium","volume":"12","author":"Watson","year":"2015","journal-title":"Cell Rep."},{"key":"ref_52","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":"ref_53","doi-asserted-by":"crossref","first-page":"122","DOI":"10.1186\/s12931-022-02042-5","article-title":"Roles of airway basal stem cells in lung homeostasis and regenerative medicine","volume":"23","author":"Wu","year":"2022","journal-title":"Respir. Res."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"4841","DOI":"10.1016\/j.cell.2022.11.005","article-title":"A human fetal lung cell atlas uncovers proximal-distal gradients of differentiation and key regulators of epithelial fates","volume":"185","author":"He","year":"2022","journal-title":"Cell"},{"key":"ref_55","first-page":"351","article-title":"A topographic atlas defines developmental origins of cell heterogeneity in the human embryonic lung","volume":"25","author":"Sountoulidis","year":"2023","journal-title":"Nat. Cell Biol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"1359","DOI":"10.1016\/j.devcel.2024.12.030","article-title":"A spatiotemporal and machine-learning platform facilitates the manufacturing of hPSC-derived esophageal mucosa","volume":"60","author":"Yang","year":"2025","journal-title":"Dev. Cell"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1038\/nature18633","article-title":"Resolving early mesoderm diversification through single-cell expression profiling","volume":"535","author":"Scialdone","year":"2016","journal-title":"Nature"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"e190374","DOI":"10.1172\/JCI190374","article-title":"SOX2 regulates foregut squamous epithelial homeostasis and is lost during Barrett\u2019s esophagus development","volume":"135","author":"Jin","year":"2025","journal-title":"J. Clin. Investig."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"e55526","DOI":"10.7554\/eLife.55526","article-title":"Delineating the early transcriptional specification of the mammalian trachea and esophagus","volume":"9","author":"Kuwahara","year":"2020","journal-title":"eLife"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"1636","DOI":"10.1172\/JCI71545","article-title":"SOX2 and p63 colocalize at genetic loci in squamous cell carcinomas","volume":"124","author":"Watanabe","year":"2014","journal-title":"J. Clin. Investig."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1165\/rcmb.2013-0419OC","article-title":"Sox2 Regulates the Emergence of Lung Basal Cells by Directly Activating the Transcription of Trp63","volume":"51","author":"Ochieng","year":"2014","journal-title":"Am. J. Respir. Cell Mol. Biol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"296","DOI":"10.1016\/j.ydbio.2008.02.035","article-title":"Sox2 is important for two crucial processes in lung development: Branching morphogenesis and epithelial cell differentiation","volume":"317","author":"Gontan","year":"2008","journal-title":"Dev. Biol."},{"key":"ref_63","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":"Domyan","year":"2009","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"110510","DOI":"10.1016\/j.celrep.2022.110510","article-title":"Tracheal separation is driven by NKX2-1-mediated repression of Efnb2 and regulation of endodermal cell sorting","volume":"38","author":"Lewis","year":"2022","journal-title":"Cell Rep."},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Tagne, J.-B., Gupta, S., Gower, A.C., Shen, S.S., Varma, S., Lakshminarayanan, M., Cao, Y., Spira, A., Volkert, T.L., and Ramirez, M.I. (2012). Genome-Wide Analyses of Nkx2-1 Binding to Transcriptional Target Genes Uncover Novel Regulatory Patterns Conserved in Lung Development and Tumors. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0029907"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"60","DOI":"10.1006\/dbio.1999.9234","article-title":"Defects in Tracheoesophageal and Lung Morphogenesis inNkx2.1(\u2212\/\u2212) Mouse Embryos","volume":"209","author":"Minoo","year":"1999","journal-title":"Dev. Biol."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"bvae163.1917","DOI":"10.1210\/jendso\/bvae163.1917","article-title":"7775 Clinical Features of NKX2-1 Gene-Related Diseases","volume":"8","author":"Fukui","year":"2024","journal-title":"J. Endocr. Soc."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1242\/dev.128.2.155","article-title":"The forkhead transcription factor Foxf1 is required for differentiation of extra-embryonic and lateral plate mesoderm","volume":"128","author":"Mahlapuu","year":"2001","journal-title":"Development"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"2397","DOI":"10.1242\/dev.128.12.2397","article-title":"Haploinsufficiency of the forkhead gene Foxf1, a target for sonic hedgehog signaling, causes lung and foregut malformations","volume":"128","author":"Mahlapuu","year":"2001","journal-title":"Development"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"833","DOI":"10.1242\/dev.02252","article-title":"Foxf1 and Foxf2 control murine gut development by limiting mesenchymal Wnt signaling and promoting extracellular matrix production","volume":"133","author":"Ormestad","year":"2006","journal-title":"Development"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1007\/s00383-007-2036-1","article-title":"Visualizing expression patterns of Shh and Foxf1 genes in the foregut and lung buds by optical projection tomography","volume":"24","author":"Sato","year":"2008","journal-title":"Pediatr. Surg. Int."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1016\/j.ydbio.2018.08.011","article-title":"FOXF1 transcription factor promotes lung morphogenesis by inducing cellular proliferation in fetal lung mesenchyme","volume":"443","author":"Ustiyan","year":"2018","journal-title":"Dev. Biol."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1038\/s41374-021-00534-4","article-title":"Forkhead box F1 induces columnar phenotype and epithelial-to-mesenchymal transition in esophageal squamous cells to initiate Barrett\u2019s like metaplasia","volume":"101","author":"De","year":"2021","journal-title":"Lab. Investig."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"1580","DOI":"10.1101\/gad.10.13.1580","article-title":"Bone morphogenetic proteins: Multifunctional regulators of vertebrate development","volume":"10","author":"Hogan","year":"1996","journal-title":"Genes. Dev."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"1693","DOI":"10.1242\/dev.122.6.1693","article-title":"Evidence from normal expression and targeted misexpression that Bone Morphogenetic Protein-4 (Bmp-4) plays a role in mouse embryonic lung morphogenesis","volume":"122","author":"Bellusci","year":"1996","journal-title":"Development"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"746","DOI":"10.1002\/dvdy.21075","article-title":"Aberrant Bmp signaling and notochord delamination in the pathogenesis of esophageal atresia","volume":"236","author":"Li","year":"2007","journal-title":"Dev. Dyn."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1016\/j.ydbio.2008.07.021","article-title":"Bmp4 is required for tracheal formation: A novel mouse model for tracheal agenesis","volume":"322","author":"Li","year":"2008","journal-title":"Dev. Biol."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"2105","DOI":"10.1101\/gad.9.17.2105","article-title":"Bone morphogenetic protein-4 is required for mesoderm formation and patterning in the mouse","volume":"9","author":"Winnier","year":"1995","journal-title":"Genes. Dev."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"847","DOI":"10.1038\/nmeth.3016","article-title":"A method to recapitulate early embryonic spatial patterning in human embryonic stem cells","volume":"11","author":"Warmflash","year":"2014","journal-title":"Nat. Methods"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1016\/j.ydbio.2023.04.007","article-title":"Disruption of BMP4 signaling is associated with laryngeal birth defects in a mouse model","volume":"500","author":"Burra","year":"2023","journal-title":"Dev. Biol."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"L1033","DOI":"10.1152\/ajplung.00277.2004","article-title":"Smad1 expression and function during mouse embryonic lung branching morphogenesis","volume":"288","author":"Chen","year":"2005","journal-title":"Am. J. Physiol.-Lung Cell. Mol. Physiol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"422","DOI":"10.1111\/j.1432-0436.2006.00096.x","article-title":"Morphogenesis of the trachea and esophagus: Current players and new roles for noggin and Bmps","volume":"74","author":"Que","year":"2006","journal-title":"Differentiation"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"599","DOI":"10.1016\/S0092-8674(00)80133-6","article-title":"The Spemann Organizer Signal noggin Binds and Inactivates Bone Morphogenetic Protein 4","volume":"86","author":"Zimmerman","year":"1996","journal-title":"Cell"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/j.semcdb.2016.12.008","article-title":"Development and stem cells of the esophagus","volume":"66","author":"Zhang","year":"2017","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"4171","DOI":"10.1242\/dev.056077","article-title":"BMP signaling in the development of the mouse esophagus and forestomach","volume":"137","author":"Rodriguez","year":"2010","journal-title":"Development"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1038\/s41392-021-00762-6","article-title":"Wnt\/\u03b2-catenin signalling: Function, biological mechanisms, and therapeutic opportunities","volume":"7","author":"Liu","year":"2022","journal-title":"Signal Transduct. Target. Ther."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"844","DOI":"10.1016\/j.stem.2017.03.001","article-title":"Efficient Derivation of Functional Human Airway Epithelium from Pluripotent Stem Cells via Temporal Regulation of Wnt Signaling","volume":"20","author":"McCauley","year":"2017","journal-title":"Cell Stem Cell"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"472","DOI":"10.1016\/j.stem.2017.08.014","article-title":"Differentiation of human pluripotent stem cells into functional lung alveolar epithelial cells","volume":"21","author":"Jacob","year":"2017","journal-title":"Cell Stem Cell"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1016\/j.ydbio.2015.06.009","article-title":"Endodermal Wnt signaling is required for tracheal cartilage formation","volume":"405","author":"Snowball","year":"2015","journal-title":"Dev. Biol."},{"key":"ref_90","doi-asserted-by":"crossref","unstructured":"Woo, J., Miletich, I., Kim, B.-M., Sharpe, P.T., and Shivdasani, R.A. (2011). Barx1-Mediated Inhibition of Wnt Signaling in the Mouse Thoracic Foregut Controls Tracheo-Esophageal Septation and Epithelial Differentiation. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0022493"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"913","DOI":"10.1016\/0092-8674(93)90535-X","article-title":"The TGF\u03b2 homolog dpp and the segment polarity gene hedgehog are required for propagation of a morphogenetic wave in the Drosophila retina","volume":"75","author":"Heberlein","year":"1993","journal-title":"Cell"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1038\/1717","article-title":"Sonic hedgehog is essential to foregut development","volume":"20","author":"Litingtung","year":"1998","journal-title":"Nat. Genet."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"e77055","DOI":"10.7554\/eLife.77055","article-title":"Hedgehog regulation of epithelial cell state and morphogenesis in the larynx","volume":"11","author":"Ramachandran","year":"2022","journal-title":"eLife"},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1002\/dvdy.20105","article-title":"Role of Sonic hedgehog in patterning of tracheal\u2013bronchial cartilage and the peripheral lung","volume":"231","author":"Miller","year":"2004","journal-title":"Dev. Dyn."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"514","DOI":"10.1002\/dvdy.22192","article-title":"Regulation of Sox9 by Sonic Hedgehog (Shh) is essential for patterning and formation of tracheal cartilage","volume":"239","author":"Park","year":"2010","journal-title":"Dev. Dyn."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"589","DOI":"10.1016\/j.devcel.2020.04.016","article-title":"Hedgehog Signaling Regulates Epithelial Morphogenesis to Position the Ventral Embryonic Midline","volume":"53","author":"Arraf","year":"2020","journal-title":"Dev. Cell"},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"1142","DOI":"10.1002\/humu.21328","article-title":"Molecular analysis expands the spectrum of phenotypes associated with GLI3 mutations","volume":"31","author":"Johnston","year":"2010","journal-title":"Hum. Mutat."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"5703","DOI":"10.1096\/fj.201701581R","article-title":"Loss of sonic hedgehog gene leads to muscle development disorder and megaesophagus in mice","volume":"32","author":"Jia","year":"2018","journal-title":"FASEB J."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1242\/dev.124.1.53","article-title":"Involvement of Sonic hedgehog (Shh) in mouse embryonic lung growth and morphogenesis","volume":"124","author":"Bellusci","year":"1997","journal-title":"Development"},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"3731","DOI":"10.1242\/dev.096560","article-title":"Localized Fgf10 expression is not required for lung branching morphogenesis but prevents differentiation of epithelial progenitors","volume":"140","author":"Volckaert","year":"2013","journal-title":"Development"},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"dev171496","DOI":"10.1242\/dev.171496","article-title":"Wnt\/Fgf crosstalk is required for the specification of basal cells in the mouse trachea","volume":"146","author":"Hou","year":"2019","journal-title":"Development"},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1002\/stem.249","article-title":"FGF2 Specifies hESC-Derived Definitive Endoderm into Foregut\/Midgut Cell Lineages in a Concentration-Dependent Manner","volume":"28","author":"Ameri","year":"2009","journal-title":"Stem Cells"},{"key":"ref_103","doi-asserted-by":"crossref","unstructured":"Baran, K., Skrzynska, K., Czyrek, A.A., Wittek, A., Krowarsch, D., Szlachcic, A., Zakrzewska, M., and Chudzian, J. (2025). Fibroblast Growth Factors in Lung Development and Regeneration: Mechanisms and Therapeutic Potential. Cells, 14.","DOI":"10.3390\/cells14161256"},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1242\/dev.051680","article-title":"FGF10 controls the patterning of the tracheal cartilage rings via Shh","volume":"138","author":"Sala","year":"2011","journal-title":"Development"},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"1999","DOI":"10.1002\/dvdy.22032","article-title":"Conditional gene inactivation reveals roles for Fgf10 and Fgfr2 in establishing a normal pattern of epithelial branching in the mouse lung","volume":"238","author":"Abler","year":"2009","journal-title":"Dev. Dyn."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1016\/j.gep.2015.12.003","article-title":"Expression of FGFs during early mouse tongue development","volume":"20","author":"Du","year":"2016","journal-title":"Gene Expr. Patterns"},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"537","DOI":"10.1016\/j.jpedsurg.2003.12.004","article-title":"Aberrant fibroblast growth factor receptor 2 signalling in esophageal atresia with tracheoesophageal fistula","volume":"39","author":"Spilde","year":"2004","journal-title":"J. Pediatr. Surg."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"322","DOI":"10.1016\/j.devcel.2015.10.006","article-title":"FGF-Regulated ETV Transcription Factors Control FGF-SHH Feedback Loop in Lung Branching","volume":"35","author":"Herriges","year":"2015","journal-title":"Dev. Cell"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"477","DOI":"10.1002\/dvdy.22504","article-title":"Patterning and plasticity in development of the respiratory lineage","volume":"240","author":"Domyan","year":"2011","journal-title":"Dev. Dyn."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"10","DOI":"10.4161\/cc.9.1.10272","article-title":"Wnt signaling and specification of the respiratory endoderm","volume":"9","author":"Goss","year":"2010","journal-title":"Cell Cycle"},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"200","DOI":"10.1359\/jbmr.090806","article-title":"Wnt inhibitors Dkk1 and Sost are downstream targets of BMP signaling through the type IA receptor (BMPRIA) in osteoblasts","volume":"25","author":"Kamiya","year":"2010","journal-title":"J. Bone Miner. Res."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"1627","DOI":"10.1242\/dev.127.8.1627","article-title":"Transdifferentiation of esophageal smooth to skeletal muscle is myogenic bHLH factor-dependent","volume":"127","author":"Kablar","year":"2000","journal-title":"Development"},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"68","DOI":"10.1038\/s41525-022-00338-5","article-title":"SMAD6-deficiency in human genetic disorders","volume":"7","author":"Luyckx","year":"2022","journal-title":"npj Genom. Med."},{"key":"ref_114","doi-asserted-by":"crossref","unstructured":"Yoshida, T., Matsuda, M., and Hirashima, T. (2020). Incoherent Feedforward Regulation via Sox9 and ERK Underpins Mouse Tracheal Cartilage Development. Front. Cell Dev. Biol., 8.","DOI":"10.3389\/fcell.2020.585640"},{"key":"ref_115","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":"ref_116","doi-asserted-by":"crossref","first-page":"342","DOI":"10.1002\/dvdy.24234","article-title":"Wnt and FGF mediated epithelial-mesenchymal crosstalk during lung development","volume":"244","author":"Volckaert","year":"2015","journal-title":"Dev. Dyn."},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"3197","DOI":"10.1242\/dev.110254","article-title":"Crucial requirement of ERK\/MAPK signaling in respiratory tract development","volume":"141","author":"Boucherat","year":"2014","journal-title":"Development"},{"key":"ref_118","doi-asserted-by":"crossref","unstructured":"Raslan, A.A., Oh, Y.J., Jin, Y.R., and Yoon, J.K. (2022). R-Spondin2, a Positive Canonical WNT Signaling Regulator, Controls the Expansion and Differentiation of Distal Lung Epithelial Stem\/Progenitor Cells in Mice. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23063089"},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"420","DOI":"10.1089\/thy.2019.0828","article-title":"Enhanced Canonical Wnt Signaling During Early Zebrafish Development Perturbs the Interaction of Cardiac Mesoderm and Pharyngeal Endoderm and Causes Thyroid Specification Defects","volume":"31","author":"Vandernoot","year":"2020","journal-title":"Thyroid\u00ae"},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"2207","DOI":"10.1242\/dev.001230","article-title":"Repression of Wnt\/\u03b2-catenin signaling in the anterior endoderm is essential for liver and pancreas development","volume":"134","author":"McLin","year":"2007","journal-title":"Development"},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"1670","DOI":"10.1101\/gad.10.13.1670","article-title":"Hepatic specification of the gut endoderm in vitro: Cell signaling and transcriptional control","volume":"10","author":"Gualdi","year":"1996","journal-title":"Genes Dev."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"36","DOI":"10.1016\/j.stem.2021.10.010","article-title":"Functional human gastrointestinal organoids can be engineered from three primary germ layers derived separately from pluripotent stem cells","volume":"29","author":"Eicher","year":"2022","journal-title":"Cell Stem Cell"},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"2277","DOI":"10.1172\/JCI89950","article-title":"Prospective isolation of NKX2-1-expressing human lung progenitors derived from pluripotent stem cells","volume":"127","author":"Hawkins","year":"2017","journal-title":"J. Clin. Investig."},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"653","DOI":"10.1007\/s13770-024-00628-2","article-title":"Lung Organoids: Systematic Review of Recent Advancements and its Future Perspectives","volume":"21","author":"Thangam","year":"2024","journal-title":"Tissue Eng. Regen. Med."},{"key":"ref_125","doi-asserted-by":"crossref","unstructured":"Luce, E., and Duclos-Vallee, J.-C. (2025). Stem Cells and Organoids: A Paradigm Shift in Preclinical Models Toward Personalized Medicine. Pharmaceuticals, 18.","DOI":"10.3390\/ph18070992"},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"1375","DOI":"10.1007\/s12015-024-10723-5","article-title":"Replacing Animal Testing with Stem Cell-Organoids: Advantages and Limitations","volume":"20","author":"Park","year":"2024","journal-title":"Stem Cell Rev. Rep."},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"84","DOI":"10.1038\/nbt.2754","article-title":"Efficient generation of lung and airway epithelial cells from human pluripotent stem cells","volume":"32","author":"Huang","year":"2013","journal-title":"Nat. Biotechnol."},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"102711","DOI":"10.1016\/j.scr.2022.102711","article-title":"Generation of three induced pluripotent stem cells lines from patients with esophageal atresia\/tracheoesophageal fistula type C","volume":"60","author":"Raad","year":"2022","journal-title":"Stem Cell Res."},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"100197","DOI":"10.1016\/j.crtox.2024.100197","article-title":"iPSC-derived and Patient-Derived Organoids: Applications and challenges in scalability and reproducibility as pre-clinical models","volume":"7","author":"Heinzelmann","year":"2024","journal-title":"Curr. Res. Toxicol."},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"e399","DOI":"10.1002\/wdev.399","article-title":"Human pluripotent stem cell-derived lung organoids: Potential applications in development and disease modeling","volume":"10","author":"Tian","year":"2021","journal-title":"WIREs Dev. Biol."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"1534","DOI":"10.1038\/nbt1163","article-title":"Efficient differentiation of human embryonic stem cells to definitive endoderm","volume":"23","author":"Agulnick","year":"2005","journal-title":"Nat. Biotechnol."},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"4295","DOI":"10.1016\/j.cell.2025.05.041","article-title":"Co-development of mesoderm and endoderm enables organotypic vascularization in lung and gut organoids","volume":"188","author":"Miao","year":"2025","journal-title":"Cell"},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1016\/j.diff.2016.04.002","article-title":"Redefining definitive endoderm subtypes by robust induction of human induced pluripotent stem cells","volume":"92","author":"Matsuno","year":"2016","journal-title":"Differentiation"},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1016\/j.stem.2020.09.017","article-title":"Derivation of Airway Basal Stem Cells from Human Pluripotent Stem Cells","volume":"28","author":"Hawkins","year":"2021","journal-title":"Cell Stem Cell"},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"e32839","DOI":"10.7554\/eLife.32839","article-title":"Micropattern differentiation of mouse pluripotent stem cells recapitulates embryo regionalized cell fate patterning","volume":"7","author":"Morgani","year":"2018","journal-title":"eLife"},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s11626-014-9801-y","article-title":"Improved efficiency of definitive endoderm induction from human induced pluripotent stem cells in feeder and serum-free culture system","volume":"51","author":"Ninomiya","year":"2015","journal-title":"Vitr. Cell. Dev. Biol.-Anim."},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"dev201023","DOI":"10.1242\/dev.201023","article-title":"Mesodermal FGF and BMP govern the sequential stages of zebrafish thyroid specification","volume":"150","author":"Haerlingen","year":"2023","journal-title":"Development"},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"1662","DOI":"10.1126\/science.1188302","article-title":"Reconstituting Organ-Level Lung Functions on a Chip","volume":"328","author":"Huh","year":"2010","journal-title":"Science"},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1038\/nmeth.3697","article-title":"Small airway-on-a-chip enables analysis of human lung inflammation and drug responses in vitro","volume":"13","author":"Benam","year":"2016","journal-title":"Nat. Methods"}],"container-title":["Cells"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4409\/15\/5\/448\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,3,3]],"date-time":"2026-03-03T13:12:24Z","timestamp":1772543544000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4409\/15\/5\/448"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,3,3]]},"references-count":139,"journal-issue":{"issue":"5","published-online":{"date-parts":[[2026,3]]}},"alternative-id":["cells15050448"],"URL":"https:\/\/doi.org\/10.3390\/cells15050448","relation":{},"ISSN":["2073-4409"],"issn-type":[{"value":"2073-4409","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,3,3]]}}}