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This mechanism activates discrete spatial patterns of epicardial gene expression, including\n                    <jats:italic toggle=\"yes\">vegfaa<\/jats:italic>\n                    ,\n                    <jats:italic toggle=\"yes\">loxl2a<\/jats:italic>\n                    , and\n                    <jats:italic toggle=\"yes\">col12a1b<\/jats:italic>\n                    . Through live and fixed imaging, we find that cardiomyocytes and endothelial cells exhibit coordinated expansion patterns through third-party epicardial signals that are required for both coronary development and myocardial expansion. Using\n                    <jats:italic toggle=\"yes\">\n                      cxcr4a\n                      <jats:sup>um20<\/jats:sup>\n                    <\/jats:italic>\n                    mutants lacking functional coronary vessels, we show that coronary vessels provide negative feedback on epicardial hypoxia, while positively responding to the same hypoxic cues that drive myocardial expansion. Disruption of this negative feedback leads to increased myocardial stiffness through dysregulated extracellular matrix crosslinking as observed in pathological conditions such as cardiomyopathies. These findings establish the role of regional epicardial hypoxia within a fundamental regulatory network that drives appropriate regional tissue growth with integrated vascular supply during cardiac morphogenesis.\n                  <\/jats:p>","DOI":"10.1073\/pnas.2536457123","type":"journal-article","created":{"date-parts":[[2026,6,22]],"date-time":"2026-06-22T19:02:29Z","timestamp":1782154949000},"update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":0,"title":["Hypoxia-mediated epicardial signaling coordinates coronary angiogenesis and myocardial expansion during zebrafish ventricle maturation"],"prefix":"10.1073","volume":"123","author":[{"ORCID":"https:\/\/orcid.org\/0009-0009-2443-777X","authenticated-orcid":false,"given":"Ku-Chi","family":"Tsao","sequence":"first","affiliation":[{"name":"Cardiovascular Research Institute, Weill Cornell Medicine","place":["New York"]},{"id":[{"id":"https:\/\/ror.org\/02r109517","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Cell and Developmental Biology, Weill Cornell Medicine","place":["New York"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0004-7186-0494","authenticated-orcid":false,"given":"Isaac","family":"Bakis","sequence":"additional","affiliation":[{"name":"Cardiovascular Research Institute, Weill Cornell Medicine","place":["New York"]},{"id":[{"id":"https:\/\/ror.org\/02r109517","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Cell and Developmental Biology, Weill Cornell Medicine","place":["New York"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0060-4835","authenticated-orcid":false,"given":"Shuofei","family":"Sun","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/05bnh6r87","id-type":"ROR","asserted-by":"publisher"}],"name":"Meinig School of Biomedical Engineering, Cornell University","place":["Ithaca"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maki","family":"Nakayama","sequence":"additional","affiliation":[{"name":"Cardiovascular Research Institute, Weill Cornell Medicine","place":["New York"]},{"id":[{"id":"https:\/\/ror.org\/02r109517","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Cell and Developmental Biology, Weill Cornell Medicine","place":["New York"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zachary","family":"Kalmanson","sequence":"additional","affiliation":[{"name":"Cardiovascular Research Institute, Weill Cornell Medicine","place":["New York"]},{"id":[{"id":"https:\/\/ror.org\/02r109517","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Cell and Developmental Biology, Weill Cornell Medicine","place":["New York"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Laila","family":"Abd Elmagid","sequence":"additional","affiliation":[{"name":"Cardiovascular Research Institute, Weill Cornell Medicine","place":["New York"]},{"id":[{"id":"https:\/\/ror.org\/02r109517","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Cell and Developmental Biology, Weill Cornell Medicine","place":["New York"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Joshua S.","family":"Shuster","sequence":"additional","affiliation":[{"name":"Cardiovascular Research Institute, Weill Cornell Medicine","place":["New York"]},{"id":[{"id":"https:\/\/ror.org\/02r109517","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Cell and Developmental Biology, Weill Cornell Medicine","place":["New York"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0952-8760","authenticated-orcid":false,"given":"Yu","family":"Xia","sequence":"additional","affiliation":[{"name":"Cardiovascular Research Institute, Weill Cornell Medicine","place":["New York"]},{"id":[{"id":"https:\/\/ror.org\/02r109517","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Cell and Developmental Biology, Weill Cornell Medicine","place":["New York"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2023-976X","authenticated-orcid":false,"given":"James V.","family":"Eichenbaum","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/03taz7m60","id-type":"ROR","asserted-by":"publisher"}],"name":"Alfred E. 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