Koyama, Satoshi https://orcid.org/0000-0002-9286-0360
Wang, Ying https://orcid.org/0000-0001-7808-6279
Paruchuri, Kaavya https://orcid.org/0000-0003-1228-1674
Uddin, Md Mesbah https://orcid.org/0000-0003-1846-0411
Cho, So Mi J. https://orcid.org/0000-0003-2460-3335
Urbut, Sarah M. https://orcid.org/0000-0002-1135-9647
Sui, Yang
Fahed, Akl C.
Kamineni, Himani https://orcid.org/0009-0005-0863-5064
Haidermota, Sara
Hornsby, Whitney E.
Green, Robert C. https://orcid.org/0000-0001-8472-0424
Daly, Mark J. https://orcid.org/0000-0002-0949-8752
Neale, Benjamin M. https://orcid.org/0000-0003-1513-6077
Ellinor, Patrick T. https://orcid.org/0000-0002-2067-0533
Smoller, Jordan W. https://orcid.org/0000-0002-0381-6334
Slaugenhaupt, Susan A. https://orcid.org/0000-0001-7958-6494
Lebo, Matthew S. https://orcid.org/0000-0002-9733-5207
Karlson, Elizabeth W.
Martin, Alicia R. https://orcid.org/0000-0003-0241-3522
Natarajan, Pradeep https://orcid.org/0000-0001-8402-7435
Funding for this research was provided by:
U.S. Department of Health & Human Services | NIH | National Human Genome Research Institute (U01HG011719)
U.S. Department of Health & Human Services | NIH | National Human Genome Research Institute (R01HG009922)
U.S. Department of Health & Human Services | NIH | National Human Genome Research Institute (U01HG008685)
U.S. Department of Health & Human Services | NIH | National Human Genome Research Institute (U01HG008685)
U.S. Department of Health & Human Services | NIH | National Human Genome Research Institute (U01HG011719)
U.S. Department of Health & Human Services | NIH | National Heart, Lung, and Blood Institute (R01HL127564)
U.S. Department of Health & Human Services | NIH | National Heart, Lung, and Blood Institute (K99HL169733)
U.S. Department of Health & Human Services | NIH | National Heart, Lung, and Blood Institute (K99HL177340)
U.S. Department of Health & Human Services | NIH | National Heart, Lung, and Blood Institute (K08HL161448)
U.S. Department of Health & Human Services | NIH | National Heart, Lung, and Blood Institute (R01HL143295)
U.S. Department of Health & Human Services | NIH | National Heart, Lung, and Blood Institute (R01HL143295)
U.S. Department of Health & Human Services | NIH | National Heart, Lung, and Blood Institute (OT2HL161841)
MEXT | Japan Society for the Promotion of Science (202160643)
Uehara Memorial Foundation
Massachusetts General Hospital (MGH Executive Committee for Research Fund for Medical Discovery)
American Heart Association (AHA career Development Award)
Burroughs Wellcome Fund
TOPMed fellowship
U.S. Department of Health & Human Services | National Institutes of Health (U01TR003201)
U.S. Department of Health & Human Services | National Institutes of Health (OT2OD026553)
U.S. Department of Health & Human Services | National Institutes of Health (U01TR003201)
U.S. Department of Health & Human Services | National Institutes of Health (OT2OD002750)
U.S. Department of Health & Human Services | National Institutes of Health (OT2OD026553)
U.S. Department of Health & Human Services | National Institutes of Health (U01HG008685)
U.S. Department of Health & Human Services | National Institutes of Health (P30AR070253)
U.S. Department of Health & Human Services | NIH | National Institute of Mental Health (R37MH107649)
U.S. Department of Health & Human Services | NIH | National Institute of Mental Health (R01MH118233)
U.S. Department of Health & Human Services | NIH | National Institute of Mental Health (R00MH117229)
European Union’s Horizon 2020 research and innovation program
Article History
Received: 27 September 2024
Accepted: 23 October 2025
First Online: 26 November 2025
Competing interests
: K.P. reports research grants, paid to her institution, from Allelica, Apple, Amgen, AstraZeneca, Boston Scientific, Genentech/Roche, and Ionis. Y.S. reports serving as a consultant for Arboretum Lifesciences. A.C.F. reports being co-founder of Goodpath and Avigena, and serving as scientific advisor to MyOme, HeartFlow, Aditum Bio, and Arboretum Health, all unrelated to the present work. R.C.G. receives compensation for advising the following companies: Allelica, Fabric, Genomic Life and Juniper Genomics; and is co-founder of Genome Medical and Nurture Genomics. M.J.D. is a founder of Maze Therapeutics and is a member of the scientific advisory board for Neumora Therapeutics, Inc. (formerly known as RBNC Therapeutics). B.M.N. is a member of the scientific advisory board at Deep Genomics and Neumora Therapeutics, Inc. J.W.S. is a member of the Scientific Advisory Board of Sensorium Therapeutics (with equity), and has received grant support from Biogen, Inc. J.W.S. is PI of a collaborative study of the genetics of depression and bipolar disorder sponsored by 23andMe for which 23andMe provides analysis time as in-kind support but no payments. P.N. reports research grants from Allelica, Amgen, Apple, Boston Scientific, Cleerly, Genentech/Roche, Ionis, Novartis, and Silence Therapeutics, personal fees from AIRNA, Allelica, Apple, AstraZeneca, Bain Capital, Blackstone Life Sciences, Bristol Myers Squibb, Creative Education Concepts, CRISPR Therapeutics, Eli Lilly & Co, Esperion Therapeutics, Foresite Capital, Foresite Labs, Genentech/Roche, GV, HeartFlow, Incyte, Magnet Biomedicine, Merck, Novartis, Novo Nordisk, TenSixteen Bio, and Tourmaline Bio, equity in Bolt, Candela, Mercury, MyOme, Parameter Health, Preciseli, and TenSixteen Bio, royalties from Recora for intensive cardiac rehabilitation, and spousal employment at Vertex Pharmaceuticals, all unrelated to the present work. The remaining authors declare no competing interests.