{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,11]],"date-time":"2026-09-11T09:06:46Z","timestamp":1789117606111,"version":"build-2803163510"},"reference-count":80,"publisher":"The Company of Biologists","issue":"17","license":[{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/www.biologists.com\/user-licence-1-2\/"}],"funder":[{"name":"National Cancer Institute","award":["R01 CA272903"],"award-info":[{"award-number":["R01 CA272903"]}]},{"name":"National Institute of Child Health and Human Development","award":["P01 HD093363"],"award-info":[{"award-number":["P01 HD093363"]}]},{"name":"Shipley Foundation"},{"DOI":"10.13039\/100008796","name":"Allen Foundation","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100008796","id-type":"DOI","asserted-by":"crossref"}]},{"name":"L.B. Research and Education Foundation"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,9,1]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>Esophageal atresia and tracheoesophageal fistula (EA\/TEF) are congenital malformations of the foregut. We identified two distinct variants in the BMP\/TGF\u03b2 repressor SMAD6 in two individuals exhibiting EA\/TEF. We investigated the function of SMAD6 in tracheoesophageal development using two orthogonal approaches in Xenopus embryos, both resulting in foregut malformations similar to those observed in EA\/TEF. We then used human pluripotent stem cell-derived foregut epithelium and mesenchyme to explore the separate roles of SMAD6 in these germ layers. CRISPR-mediated disruption of human SMAD6 caused an increase in BMP signaling in foregut epithelium and mesenchyme consistent with its role as a BMP repressor. Loss of SMAD6 caused patterning defects in both tissue types; SMAD6\u2212\/\u2212 endoderm showed increased expression of distal gut tube markers and SMAD6\u2212\/\u2212 mesenchyme showed increased expression of ventral and posterior markers, including markers of cardiac and liver mesenchyme lineages. Furthermore, SMAD6\u2212\/\u2212\u00a0mesenchyme had decreased ability to form CD31-positive endothelial cells. Our results demonstrate that SMAD6 is required for foregut development and that rare variants in this gene are likely causative for foregut malformations in EA\/TEF.<\/jats:p>","DOI":"10.1242\/dev.205236","type":"journal-article","created":{"date-parts":[[2026,8,14]],"date-time":"2026-08-14T11:31:55Z","timestamp":1786707115000},"source":"Crossref","is-referenced-by-count":0,"title":["SMAD6 is required for normal foregut development in humans and frogs"],"prefix":"10.1242","volume":"153","author":[{"given":"Vivien","family":"Pagan","sequence":"first","affiliation":[{"name":"Center for Stem Cell and Organoid Medicine (CuSTOM), Cincinnati Children's Hospital Medical Center 1 , Cincinnati, OH 45229 ,","place":["USA"]},{"name":"Cincinnati Children's Hospital Medical Center 2 Division of Developmental Biology , , Cincinnati, OH 45229 ,","place":["USA"]},{"name":"Cincinnati Children's Hospital Medical Center 3 Development, Stem Cells, and Regenerative Medicine Graduate Program , , Cincinnati, OH 45229 ,","place":["USA"]},{"name":"University of Cincinnati College of Medicine 4 Medical Scientist Training Program , , Cincinnati, OH 45267 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Scott A.","family":"Rankin","sequence":"additional","affiliation":[{"name":"Center for Stem Cell and Organoid Medicine (CuSTOM), Cincinnati Children's Hospital Medical Center 1 , Cincinnati, OH 45229 ,","place":["USA"]},{"name":"Cincinnati Children's Hospital Medical Center 2 Division of Developmental Biology , , Cincinnati, OH 45229 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nicole A.","family":"Edwards","sequence":"additional","affiliation":[{"name":"Center for Stem Cell and Organoid Medicine (CuSTOM), Cincinnati Children's Hospital Medical Center 1 , Cincinnati, OH 45229 ,","place":["USA"]},{"name":"Cincinnati Children's Hospital Medical Center 2 Division of Developmental Biology , , Cincinnati, OH 45229 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Konrad","family":"Thorner","sequence":"additional","affiliation":[{"name":"Center for Stem Cell and Organoid Medicine (CuSTOM), Cincinnati Children's Hospital Medical Center 1 , Cincinnati, OH 45229 ,","place":["USA"]},{"name":"Cincinnati Children's Hospital Medical Center 2 Division of Developmental Biology , , Cincinnati, OH 45229 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shangqian","family":"Xie","sequence":"additional","affiliation":[{"name":"Columbia University Irving Medical Center 5 Department of Systems Biology , , New York, NY 10032 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3438-5685","authenticated-orcid":false,"given":"Wendy K.","family":"Chung","sequence":"additional","affiliation":[{"name":"Boston Children's Hospital, Harvard Medical School 6 Department of Pediatrics , , Boston, MA 02115 ,","place":["USA"]},{"name":"Columbia University Irving Medical Center 7 Departments of Pediatrics and Medicine , , New York, NY 10032 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yufeng","family":"Shen","sequence":"additional","affiliation":[{"name":"Columbia University Irving Medical Center 5 Department of Systems Biology , , New York, NY 10032 ,","place":["USA"]},{"name":"Columbia University Irving Medical Center 8 Department of Biomedical Informatics , , New York, NY 10032 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aaron M.","family":"Zorn","sequence":"additional","affiliation":[{"name":"Center for Stem Cell and Organoid Medicine (CuSTOM), Cincinnati Children's Hospital Medical Center 1 , Cincinnati, OH 45229 ,","place":["USA"]},{"name":"Cincinnati Children's Hospital Medical Center 2 Division of Developmental Biology , , Cincinnati, OH 45229 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4218-128X","authenticated-orcid":false,"given":"James M.","family":"Wells","sequence":"additional","affiliation":[{"name":"Center for Stem Cell and Organoid Medicine (CuSTOM), Cincinnati Children's Hospital Medical Center 1 , Cincinnati, OH 45229 ,","place":["USA"]},{"name":"Cincinnati Children's Hospital Medical Center 2 Division of Developmental Biology , , Cincinnati, OH 45229 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2026,9,10]]},"reference":[{"key":"2026091104163847800_DEV205236C1","doi-asserted-by":"publisher","first-page":"688","DOI":"10.1097\/MPG.0b013e3182a373dc","article-title":"Chromosome aberrations and gene mutations in patients with esophageal atresia","volume":"57","author":"Bednarczyk","year":"2013","journal-title":"J. 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