{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,12]],"date-time":"2026-05-12T18:15:28Z","timestamp":1778609728841,"version":"3.51.4"},"reference-count":55,"publisher":"Ovid Technologies (Wolters Kluwer Health)","issue":"5","content-domain":{"domain":["www.ahajournals.org"],"crossmark-restriction":true},"short-container-title":["Stroke"],"published-print":{"date-parts":[[2026,5]]},"abstract":"<jats:sec>\n                    <jats:title>BACKGROUND:<\/jats:title>\n                    <jats:p>\n                      Moyamoya disease (MMD) has a strong genetic basis, with the rare\n                      <jats:italic toggle=\"yes\">RNF213<\/jats:italic>\n                      p.Arg4810Lys variant (rs112735431) representing a major risk factor, while the broader genetic architecture and disease-relevant vascular cell types remain incompletely understood.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>METHODS:<\/jats:title>\n                    <jats:p>We conducted a genome-wide association study in Japanese individuals (n=47\u2009656; 401 MMD cases and 47\u2009255 controls). Population-level features at MMD risk loci were examined by regional allele frequency and haplotype analyses. We performed single-nucleus RNA-seq of superficial temporal arteries from patients with MMD (n=3). Cell type\u2013specific enrichment of genome-wide association study signals was assessed using the Single-Cell Disease Relevance Score. Endothelial signatures were validated by integration with publicly available single-cell data sets from controls (n=5) and immunohistochemistry for candidate markers (n=1).<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>RESULTS:<\/jats:title>\n                    <jats:p>\n                      Beyond rs112735431, we identified a genome-wide significant signal in the\n                      <jats:italic toggle=\"yes\">HDAC9-TWIST1<\/jats:italic>\n                      region (rs12530920;\n                      <jats:italic toggle=\"yes\">P<\/jats:italic>\n                      =3.3\u00d710\n                      <jats:sup>\u221214<\/jats:sup>\n                      ; odds ratio, 1.77). Conditional analysis on rs112735431 revealed a protective\n                      <jats:italic toggle=\"yes\">RNF213<\/jats:italic>\n                      missense variant, p.Asp1331Gly (rs8074015;\n                      <jats:italic toggle=\"yes\">P<\/jats:italic>\n                      =3.7\u00d710\n                      <jats:sup>\u2212<\/jats:sup>\n                      <jats:sup>9<\/jats:sup>\n                      ; odds ratio, 0.53), whose minor allele was mutually exclusive with rs112735431-A on haplotypes. Population analysis revealed geographic variation and extended haplotype structure of the rs112735431-A allele in Japan. Single-nucleus RNA-seq identified a mesenchymal-like endothelial cell (MEC) population with selective\n                      <jats:italic toggle=\"yes\">FN1<\/jats:italic>\n                      expression. Genome-wide association study\u2013prioritized disease genes were strongly enriched in MECs. MECs showed mesenchymal pathway activation with a regulatory program distinct from canonical endothelial states. The proportion of MECs was markedly increased in MMD (72% versus 28% in controls), and fibronectin 1 (FN1) expression in endothelial regions was confirmed by immunohistochemistry.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>CONCLUSIONS:<\/jats:title>\n                    <jats:p>\n                      Our findings identify a protective\n                      <jats:italic toggle=\"yes\">RNF213<\/jats:italic>\n                      p.Asp1331Gly variant (rs8074015) that is mutually exclusive with the known rs112735431-A allele. Genetic risk converges on an MEC state markedly expanded in MMD.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1161\/strokeaha.125.053747","type":"journal-article","created":{"date-parts":[[2026,3,2]],"date-time":"2026-03-02T10:01:03Z","timestamp":1772445663000},"page":"1336-1348","update-policy":"https:\/\/doi.org\/10.1161\/crossmarkpolicy","source":"Crossref","is-referenced-by-count":0,"title":["Integrative GWAS and snRNA-seq Reveal a Mesenchymal-Like Endothelial Signature in Moyamoya Disease"],"prefix":"10.1161","volume":"57","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-4043-0593","authenticated-orcid":true,"given":"Yudai","family":"Hirano","sequence":"first","affiliation":[{"name":"Faculty of Medicine, Department of Neurosurgery (Y.H., S.M., H. Imai, H.H., H.O., S.O., S.T., Y. Sakai, D.S., T.T., S.H., S. Koizumi, S.D., Y. Shinya, S. Kiyofuji, H.N., N.S.), The University of Tokyo, Japan."},{"name":"Department of Genome Informatics, Graduate School of Medicine (Y.H., K.S., S.N., Y. Shirai, Y.M., Y.O.), The University of Tokyo, Japan."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6369-3391","authenticated-orcid":true,"given":"Satoru","family":"Miyawaki","sequence":"additional","affiliation":[{"name":"Faculty of Medicine, Department of Neurosurgery (Y.H., S.M., H. Imai, H.H., H.O., S.O., S.T., Y. Sakai, D.S., T.T., S.H., S. Koizumi, S.D., Y. Shinya, S. Kiyofuji, H.N., N.S.), The University of Tokyo, Japan."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4536-1761","authenticated-orcid":true,"given":"Kyuto","family":"Sonehara","sequence":"additional","affiliation":[{"name":"Department of Genome Informatics, Graduate School of Medicine (Y.H., K.S., S.N., Y. Shirai, Y.M., Y.O.), The University of Tokyo, Japan."},{"name":"Department of Statistical Genetics, Graduate School of Medicine, The University of Osaka, Suita, Japan (K.S., S.N., Y. Shirai, Y.O.)."},{"name":"Laboratory for Systems Genetics, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan (K.S., S.N., Y. Shirai, Y.O.)."},{"name":"Wellcome Sanger Institute, Wellcome Genome Campus, Cambridge, United Kingdom (K.S.)."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7486-3146","authenticated-orcid":true,"given":"Shinichi","family":"Namba","sequence":"additional","affiliation":[{"name":"Department of Genome Informatics, Graduate School of Medicine (Y.H., K.S., S.N., Y. Shirai, Y.M., Y.O.), The University of Tokyo, Japan."},{"name":"Department of Statistical Genetics, Graduate School of Medicine, The University of Osaka, Suita, Japan (K.S., S.N., Y. Shirai, Y.O.)."},{"name":"Laboratory for Systems Genetics, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan (K.S., S.N., Y. Shirai, Y.O.)."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0626-3306","authenticated-orcid":true,"given":"Hirotaka","family":"Inoue","sequence":"additional","affiliation":[{"name":"Department of Preventive Medicine, Graduate School of Medicine (H. Inoue, D.K., S.I.), The University of Tokyo, Japan."},{"name":"Department of Neurosurgery, Graduate School of Medical Sciences, Kumamoto University, Japan (H. Inoue)."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0518-9020","authenticated-orcid":true,"given":"Yuya","family":"Shirai","sequence":"additional","affiliation":[{"name":"Department of Genome Informatics, Graduate School of Medicine (Y.H., K.S., S.N., Y. Shirai, Y.M., Y.O.), The University of Tokyo, Japan."},{"name":"Department of Statistical Genetics, Graduate School of Medicine, The University of Osaka, Suita, Japan (K.S., S.N., Y. Shirai, Y.O.)."},{"name":"Laboratory for Systems Genetics, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan (K.S., S.N., Y. Shirai, Y.O.)."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Hideaki","family":"Imai","sequence":"additional","affiliation":[{"name":"Faculty of Medicine, Department of Neurosurgery (Y.H., S.M., H. Imai, H.H., H.O., S.O., S.T., Y. Sakai, D.S., T.T., S.H., S. Koizumi, S.D., Y. Shinya, S. Kiyofuji, H.N., N.S.), The University of Tokyo, Japan."},{"name":"Department of Neurosurgery, Tokyo Shinjuku Medical Center, Japan (H. Imai, S.D.)."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Hiroki","family":"Hongo","sequence":"additional","affiliation":[{"name":"Faculty of Medicine, Department of Neurosurgery (Y.H., S.M., H. Imai, H.H., H.O., S.O., S.T., Y. Sakai, D.S., T.T., S.H., S. Koizumi, S.D., Y. Shinya, S. Kiyofuji, H.N., N.S.), The University of Tokyo, Japan."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Masahiro","family":"Shimizu","sequence":"additional","affiliation":[{"name":"Department of Neurosurgery, Kanto Neurosurgical Hospital, Kumagaya, Japan (M.S.)."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Hideaki","family":"Ono","sequence":"additional","affiliation":[{"name":"Faculty of Medicine, Department of Neurosurgery (Y.H., S.M., H. Imai, H.H., H.O., S.O., S.T., Y. Sakai, D.S., T.T., S.H., S. Koizumi, S.D., Y. Shinya, S. Kiyofuji, H.N., N.S.), The University of Tokyo, Japan."},{"name":"Department of Neurosurgery, Fuji Brain Institute and Hospital, Fujinomiya, Japan (H.O., S. Kiyofuji)."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0009-0008-4498-6410","authenticated-orcid":true,"given":"Yumie","family":"Mizoro","sequence":"additional","affiliation":[{"name":"Department of Genome Informatics, Graduate School of Medicine (Y.H., K.S., S.N., Y. Shirai, Y.M., Y.O.), The University of Tokyo, Japan."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"name":"the BioBank Japan Project;","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Koichi","family":"Matsuda","sequence":"additional","affiliation":[{"name":"Laboratory of Genome Technology, Human Genome Center, Institute of Medical Science (K.M.), The University of Tokyo, Japan."},{"name":"Department of Computational Biology and Medical Sciences, Laboratory of Clinical Genome Sequencing, Graduate School of Frontier Sciences (K.M.), The University of Tokyo, Japan."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2919-2224","authenticated-orcid":true,"given":"Shotaro","family":"Ogawa","sequence":"additional","affiliation":[{"name":"Faculty of Medicine, Department of Neurosurgery (Y.H., S.M., H. Imai, H.H., H.O., S.O., S.T., Y. Sakai, D.S., T.T., S.H., S. Koizumi, S.D., Y. Shinya, S. Kiyofuji, H.N., N.S.), The University of Tokyo, Japan."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8447-8287","authenticated-orcid":true,"given":"Seiei","family":"Torazawa","sequence":"additional","affiliation":[{"name":"Faculty of Medicine, Department of Neurosurgery (Y.H., S.M., H. Imai, H.H., H.O., S.O., S.T., Y. Sakai, D.S., T.T., S.H., S. Koizumi, S.D., Y. Shinya, S. Kiyofuji, H.N., N.S.), The University of Tokyo, Japan."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yu","family":"Sakai","sequence":"additional","affiliation":[{"name":"Faculty of Medicine, Department of Neurosurgery (Y.H., S.M., H. Imai, H.H., H.O., S.O., S.T., Y. Sakai, D.S., T.T., S.H., S. Koizumi, S.D., Y. Shinya, S. Kiyofuji, H.N., N.S.), The University of Tokyo, Japan."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Daisuke","family":"Sato","sequence":"additional","affiliation":[{"name":"Faculty of Medicine, Department of Neurosurgery (Y.H., S.M., H. Imai, H.H., H.O., S.O., S.T., Y. Sakai, D.S., T.T., S.H., S. Koizumi, S.D., Y. Shinya, S. Kiyofuji, H.N., N.S.), The University of Tokyo, Japan."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Takahiro","family":"Tsuchiya","sequence":"additional","affiliation":[{"name":"Faculty of Medicine, Department of Neurosurgery (Y.H., S.M., H. Imai, H.H., H.O., S.O., S.T., Y. Sakai, D.S., T.T., S.H., S. Koizumi, S.D., Y. Shinya, S. Kiyofuji, H.N., N.S.), The University of Tokyo, Japan."},{"name":"Department of Neurosurgery, Teraoka Memorial Hospital, Fukuyama, Japan (T.T., Y. Shinya, A.T.)."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"So","family":"Hirata","sequence":"additional","affiliation":[{"name":"Faculty of Medicine, Department of Neurosurgery (Y.H., S.M., H. Imai, H.H., H.O., S.O., S.T., Y. Sakai, D.S., T.T., S.H., S. Koizumi, S.D., Y. Shinya, S. Kiyofuji, H.N., N.S.), The University of Tokyo, Japan."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Satoshi","family":"Koizumi","sequence":"additional","affiliation":[{"name":"Faculty of Medicine, Department of Neurosurgery (Y.H., S.M., H. Imai, H.H., H.O., S.O., S.T., Y. Sakai, D.S., T.T., S.H., S. Koizumi, S.D., Y. Shinya, S. Kiyofuji, H.N., N.S.), The University of Tokyo, Japan."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Shogo","family":"Dofuku","sequence":"additional","affiliation":[{"name":"Faculty of Medicine, Department of Neurosurgery (Y.H., S.M., H. Imai, H.H., H.O., S.O., S.T., Y. Sakai, D.S., T.T., S.H., S. Koizumi, S.D., Y. Shinya, S. Kiyofuji, H.N., N.S.), The University of Tokyo, Japan."},{"name":"Department of Neurosurgery, Tokyo Shinjuku Medical Center, Japan (H. Imai, S.D.)."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0097-5804","authenticated-orcid":true,"given":"Yuki","family":"Shinya","sequence":"additional","affiliation":[{"name":"Faculty of Medicine, Department of Neurosurgery (Y.H., S.M., H. Imai, H.H., H.O., S.O., S.T., Y. Sakai, D.S., T.T., S.H., S. Koizumi, S.D., Y. Shinya, S. Kiyofuji, H.N., N.S.), The University of Tokyo, Japan."},{"name":"Department of Neurosurgery, Teraoka Memorial Hospital, Fukuyama, Japan (T.T., Y. Shinya, A.T.)."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8780-5773","authenticated-orcid":true,"given":"Satoshi","family":"Kiyofuji","sequence":"additional","affiliation":[{"name":"Faculty of Medicine, Department of Neurosurgery (Y.H., S.M., H. Imai, H.H., H.O., S.O., S.T., Y. Sakai, D.S., T.T., S.H., S. Koizumi, S.D., Y. Shinya, S. Kiyofuji, H.N., N.S.), The University of Tokyo, Japan."},{"name":"Department of Neurosurgery, Fuji Brain Institute and Hospital, Fujinomiya, Japan (H.O., S. Kiyofuji)."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0038-728X","authenticated-orcid":true,"given":"Daisuke","family":"Komura","sequence":"additional","affiliation":[{"name":"Department of Preventive Medicine, Graduate School of Medicine (H. Inoue, D.K., S.I.), The University of Tokyo, Japan."}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Akira","family":"Teraoka","sequence":"additional","affiliation":[{"name":"Department of Neurosurgery, Teraoka Memorial Hospital, Fukuyama, Japan (T.T., Y. 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