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However, a portion of CHD-associated 4q deletions without known CHD genes suggests unknown CHD genes within these intervals. Here, we have shown that knockdown of\n                    <jats:italic>SORBS2<\/jats:italic>\n                    , a 4q interval gene, disrupted sarcomeric integrity of cardiomyocytes and caused reduced cardiomyocyte number in human embryonic stem cell differentiation model. Molecular analyses revealed decreased expression of second heart field (SHF) marker genes and impaired NOTCH and SHH signaling in\n                    <jats:italic>SORBS2-<\/jats:italic>\n                    knockdown cells. Exogenous SHH rescued\n                    <jats:italic>SORBS2<\/jats:italic>\n                    knockdown-induced cardiomyocyte differentiation defects.\n                    <jats:italic>\n                      Sorbs2\n                      <jats:sup>-\/-<\/jats:sup>\n                    <\/jats:italic>\n                    mouse mutants had atrial septal hypoplasia\/aplasia or double atrial septum (DAS) derived from impaired posterior SHF with a similar expression alteration. Rare\n                    <jats:italic>SORBS2<\/jats:italic>\n                    variants were significantly enriched in a cohort of 300 CHD patients. Our findings indicate that\n                    <jats:italic>SORBS2<\/jats:italic>\n                    is a regulator of SHF development and its variants contribute to CHD pathogenesis. The presence of DAS in\n                    <jats:italic>\n                      Sorbs2\n                      <jats:sup>-\/-<\/jats:sup>\n                    <\/jats:italic>\n                    hearts reveals the first molecular etiology of this rare anomaly linked to paradoxical thromboembolism.\n                  <\/jats:p>","DOI":"10.7554\/elife.67481","type":"journal-article","created":{"date-parts":[[2021,6,7]],"date-time":"2021-06-07T20:17:13Z","timestamp":1623097033000},"update-policy":"https:\/\/doi.org\/10.7554\/elife.67481","source":"Crossref","is-referenced-by-count":18,"title":["SORBS2 is a genetic factor contributing to cardiac malformation of 4q deletion syndrome patients"],"prefix":"10.7554","volume":"10","author":[{"given":"Fei","family":"Liang","sequence":"first","affiliation":[{"name":"Neonatal Intensive Care Unit, Shanghai Pediatric Congenital Heart Disease Institute and Pediatric Translational Medicine Institute, Shanghai Children\u2019s Medical Center, Shanghai Jiao Tong University School of Medicine, Shanghai, China"},{"name":"Shanghai Pediatric Congenital Heart Disease Institute and Pediatric Translational Medicine Institute, Shanghai Children\u2019s Medical Center, Shanghai Jiao Tong University School of Medicine, Shanghai, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4376-1398","authenticated-orcid":true,"given":"Bo","family":"Wang","sequence":"additional","affiliation":[{"name":"Shanghai Key Laboratory of Clinical Molecular Diagnostics for Pediatrics, Pediatric Translational Medicine Institute, Shanghai Children\u2019s Medical Center, Shanghai Jiao Tong University School of Medicine, Shanghai, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Juan","family":"Geng","sequence":"additional","affiliation":[{"name":"Shanghai Key Laboratory of Clinical Molecular Diagnostics for Pediatrics, Pediatric Translational Medicine Institute, Shanghai Children\u2019s Medical 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