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Here, using clustered protocadherin (\n                    <jats:italic>cPcdh<\/jats:italic>\n                    ) and\n                    <jats:italic>\u03b2\u2010globin<\/jats:italic>\n                    genes as model systems, we showed that\n                    <jats:italic>WAVE2<\/jats:italic>\n                    , a member of the WASP family, regulates chromatin organization by maintaining heterochromatin dynamics. Specifically, by CRISPR DNA\u2010fragment editing, in conjunction with integrated analyses of ChIP\u2010seq, MeDIP\u2010seq, ATAC\u2010seq, QHR\u20104C, and RNA\u2010seq, we showed that deposition of H3K9me3, a key heterochromatin mark, is significantly decreased at the\n                    <jats:italic>cPcdh<\/jats:italic>\n                    locus upon\n                    <jats:italic>WAVE2<\/jats:italic>\n                    deletion, concurrent with aberrant accumulation of CTCF\/cohesin complex at promoter regions and spatial reorganization of chromatin architecture around the nucleolus. In addition,\n                    <jats:italic>REST\/NRSF<\/jats:italic>\n                    exerts a similar heterochromatin\u2010dependent effect on the\n                    <jats:italic>cPcdh<\/jats:italic>\n                    locus. Finally, genetic and genomic data showed that\n                    <jats:italic>WAVE2<\/jats:italic>\n                    regulates\n                    <jats:italic>\u03b2\u2010globin<\/jats:italic>\n                    gene expression by maintaining heterochromatin status. Together, our data suggested that\n                    <jats:italic>WAVE2<\/jats:italic>\n                    and\n                    <jats:italic>REST\/NRSF<\/jats:italic>\n                    regulate developmental expression of clustered genes in a heterochromatin\u2010dependent manner.\n                  <\/jats:p>","DOI":"10.1096\/fj.202601513r","type":"journal-article","created":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T05:07:20Z","timestamp":1787116040000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["<scp>WAVE2<\/scp>\n                    and\n                    <scp>REST<\/scp>\n                    \/\n                    <scp>NRSF<\/scp>\n                    Regulate Clustered Gene Expression by Maintaining Heterochromatin Organization"],"prefix":"10.1096","volume":"40","author":[{"given":"Leyang","family":"Wang","sequence":"first","affiliation":[{"name":"Center for Comparative Biomedicine, Key Laboratory of Systems Biomedicine (MOE) Institute of Systems Biomedicine, Shanghai Jiao Tong University  Shanghai China"},{"name":"Shanghai Key Laboratory of Gene Editing and Cell\u2010Based Immunotherapy for Hematological Diseases, State Key Laboratory of Medical Genomics Ruijin Hospital, Shanghai Jiao Tong University School of Medicine  Shanghai China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuanxiao","family":"Tang","sequence":"additional","affiliation":[{"name":"Center for Comparative Biomedicine, Key Laboratory of Systems Biomedicine (MOE) Institute of Systems Biomedicine, Shanghai Jiao Tong University  Shanghai China"},{"name":"Shanghai Key Laboratory of Gene Editing and Cell\u2010Based Immunotherapy for Hematological Diseases, State Key Laboratory of Medical Genomics Ruijin Hospital, Shanghai Jiao Tong University School of Medicine  Shanghai 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