Loïc Yengo, Sailaja Vedantam, Eirini Marouli, Julia Sidorenko, Eric Bartell, Saori Sakaue, Marielisa Graff, Anders U. Eliasen, Yunxuan Jiang, Sridharan Raghavan, Jenkai Miao, Joshua D. Arias, Sarah E. Graham, Ronen E. Mukamel, Cassandra N. Spracklen, Xianyong Yin, Shyh-Huei Chen, Teresa Ferreira, Heather H. Highland, Yingjie Ji, Tugce Karaderi, Kuang Lin, Kreete Lüll, Deborah E. Malden, Carolina Medina-Gomez, Moara Machado, Amy Moore, Sina Rüeger, Xueling Sim, Scott Vrieze, Tarunveer S. Ahluwalia, Masato Akiyama, Matthew A. Allison, Marcus Alvarez, Mette K. Andersen, Alireza Ani, Vivek Appadurai, Liubov Arbeeva, Seema Bhaskar, Lawrence F. Bielak, Sailalitha Bollepalli, Lori L. Bonnycastle, Jette Bork-Jensen, Jonathan P. Bradfield, Yuki Bradford, Peter S. Braund, Jennifer A. Brody, Kristoffer S. Burgdorf, Brian E. Cade, Hui Cai, Qiuyin Cai, Archie Campbell, Marisa Cañadas-Garre, Eulalia Catamo, Jin-Fang Chai, Xiaoran Chai, Li-Ching Chang, Yi-Cheng Chang, Chien-Hsiun Chen, Alessandra Chesi, Seung Hoan Choi, Ren-Hua Chung, Massimiliano Cocca, Maria Pina Concas, Christian Couture, Gabriel Cuellar-Partida, Rebecca Danning, E. Warwick Daw, Frauke Degenhard, Graciela E. Delgado, Alessandro Delitala, Ayse Demirkan, Xuan Deng, Poornima Devineni, Alexander Dietl, Maria Dimitriou, Latchezar Dimitrov, Rajkumar Dorajoo, Arif B. Ekici, Jorgen E. Engmann, Zammy Fairhurst-Hunter, Aliki-Eleni Farmaki, Jessica D. Faul, Juan-Carlos Fernandez-Lopez, Lukas Forer, Margherita Francescatto, Sandra Freitag-Wolf, Christian Fuchsberger, Tessel E. Galesloot, Yan Gao, Zishan Gao, Frank Geller, Olga Giannakopoulou, Franco Giulianini, Anette P. Gjesing, Anuj Goel, Scott D. Gordon, Mathias Gorski, Jakob Grove, Xiuqing Guo, Stefan Gustafsson, Jeffrey Haessler, Thomas F. Hansen, Aki S. Havulinna, Simon J. Haworth, Jing He, Nancy Heard-Costa, Prashantha Hebbar, George Hindy, Yuk-Lam A. Ho, Edith Hofer, Elizabeth Holliday, Katrin Horn, Whitney E. Hornsby, Jouke-Jan Hottenga, Hongyan Huang, Jie Huang, Alicia Huerta-Chagoya, Jennifer E. Huffman, Yi-Jen Hung, Shaofeng Huo, Mi Yeong Hwang, Hiroyuki Iha, Daisuke D. Ikeda, Masato Isono, Anne U. Jackson, Susanne Jäger, Iris E. Jansen, Ingegerd Johansson, Jost B. Jonas, Anna Jonsson, Torben Jørgensen, Ioanna-Panagiota Kalafati, Masahiro Kanai, Stavroula Kanoni, Line L. Kårhus, Anuradhani Kasturiratne, Tomohiro Katsuya, Takahisa Kawaguchi, Rachel L. Kember, Katherine A. Kentistou, Han-Na Kim, Young Jin Kim, Marcus E. Kleber, Maria J. Knol, Azra Kurbasic, Marie Lauzon, Phuong Le, Rodney Lea, Jong-Young Lee, Hampton L. Leonard, Shengchao A. Li, Xiaohui Li, Xiaoyin Li, Jingjing Liang, Honghuang Lin, Shih-Yi Lin, Jun Liu, Xueping Liu, Ken Sin Lo, Jirong Long, Laura Lores-Motta, Jian’an Luan, Valeriya Lyssenko, Leo-Pekka Lyytikäinen, Anubha Mahajan, Vasiliki Mamakou, Massimo Mangino, Ani Manichaikul, Jonathan Marten, Manuel Mattheisen, Laven Mavarani, Aaron F. McDaid, Karina Meidtner, Tori L. Melendez, Josep M. Mercader, Yuri Milaneschi, Jason E. Miller, Iona Y. Millwood, Pashupati P. Mishra, Ruth E. Mitchell, Line T. Møllehave, Anna Morgan, Soeren Mucha, Matthias Munz, Masahiro Nakatochi, Christopher P. Nelson, Maria Nethander, Chu Won Nho, Aneta A. Nielsen, Ilja M. Nolte, Suraj S. Nongmaithem, Raymond Noordam, Ioanna Ntalla, Teresa Nutile, Anita Pandit, Paraskevi Christofidou, Katri Pärna, Marc Pauper, Eva R. B. Petersen, Liselotte V. Petersen, Niina Pitkänen, Ozren Polašek, Alaitz Poveda, Michael H. Preuss, Saiju Pyarajan, Laura M. Raffield, Hiromi Rakugi, Julia Ramirez, Asif Rasheed, Dennis Raven, Nigel W. Rayner, Carlos Riveros, Rebecca Rohde, Daniela Ruggiero, Sanni E. Ruotsalainen, Kathleen A. Ryan, Maria Sabater-Lleal, Richa Saxena, Markus Scholz, Anoop Sendamarai, Botong Shen, Jingchunzi Shi, Jae Hun Shin, Carlo Sidore, Colleen M. Sitlani, Roderick C. Slieker, Roelof A. J. Smit, Albert V. Smith, Jennifer A. Smith, Laura J. Smyth, Lorraine Southam, Valgerdur Steinthorsdottir, Liang Sun, Fumihiko Takeuchi, Divya Sri Priyanka Tallapragada, Kent D. Taylor, Bamidele O. Tayo, Catherine Tcheandjieu, Natalie Terzikhan, Paola Tesolin, Alexander Teumer, Elizabeth Theusch, Deborah J. Thompson, Gudmar Thorleifsson, Paul R. H. J. Timmers, Stella Trompet, Constance Turman, Simona Vaccargiu, Sander W. van der Laan, Peter J. van der Most, Jan B. van Klinken, Jessica van Setten, Shefali S. Verma, Niek Verweij, Yogasudha Veturi, Carol A. Wang, Chaolong Wang, Lihua Wang, Zhe Wang, Helen R. Warren, Wen Bin Wei, Ananda R. Wickremasinghe, Matthias Wielscher, Kerri L. Wiggins, Bendik S. Winsvold, Andrew Wong, Yang Wu, Matthias Wuttke, Rui Xia, Tian Xie, Ken Yamamoto, Jingyun Yang, Jie Yao, Hannah Young, Noha A. Yousri, Lei Yu, Lingyao Zeng, Weihua Zhang, Xinyuan Zhang, Jing-Hua Zhao, Wei Zhao, Wei Zhou, Martina E. Zimmermann, Magdalena Zoledziewska, Linda S. Adair, Hieab H. H. Adams, Carlos A. Aguilar-Salinas, Fahd Al-Mulla, Donna K. Arnett, Folkert W. Asselbergs, Bjørn Olav Åsvold, John Attia, Bernhard Banas, Stefania Bandinelli, David A. Bennett, Tobias Bergler, Dwaipayan Bharadwaj, Ginevra Biino, Hans Bisgaard, Eric Boerwinkle, Carsten A. Böger, Klaus Bønnelykke, Dorret I. Boomsma, Anders D. Børglum, Judith B. Borja, Claude Bouchard, Donald W. Bowden, Ivan Brandslund, Ben Brumpton, Julie E. Buring, Mark J. Caulfield, John C. Chambers, Giriraj R. Chandak, Stephen J. Chanock, Nish Chaturvedi, Yii-Der Ida Chen, Zhengming Chen, Ching-Yu Cheng, Ingrid E. Christophersen, Marina Ciullo, John W. Cole, Francis S. Collins, Richard S. Cooper, Miguel Cruz, Francesco Cucca, L. Adrienne Cupples, Michael J. Cutler, Scott M. Damrauer, Thomas M. Dantoft, Gert J. de Borst, Lisette C. P. G. M. de Groot, Philip L. De Jager, Dominique P. V. de Kleijn, H. Janaka de Silva, George V. Dedoussis, Anneke I. den Hollander, Shufa Du, Douglas F. Easton, Petra J. M. Elders, A. Heather Eliassen, Patrick T. Ellinor, Sölve Elmståhl, Jeanette Erdmann, Michele K. Evans, Diane Fatkin, Bjarke Feenstra, Mary F. Feitosa, Luigi Ferrucci, Ian Ford, Myriam Fornage, Andre Franke, Paul W. Franks, Barry I. Freedman, Paolo Gasparini, Christian Gieger, Giorgia Girotto, Michael E. Goddard, Yvonne M. Golightly, Clicerio Gonzalez-Villalpando, Penny Gordon-Larsen, Harald Grallert, Struan F. A. Grant, Niels Grarup, Lyn Griffiths, Vilmundur Gudnason, Christopher Haiman, Hakon Hakonarson, Torben Hansen, Catharina A. Hartman, Andrew T. Hattersley, Caroline Hayward, Susan R. Heckbert, Chew-Kiat Heng, Christian Hengstenberg, Alex W. Hewitt, Haretsugu Hishigaki, Carel B. Hoyng, Paul L. Huang, Wei Huang, Steven C. Hunt, Kristian Hveem, Elina Hyppönen, William G. Iacono, Sahoko Ichihara, M. Arfan Ikram, Carmen R. Isasi, Rebecca D. Jackson, Marjo-Riitta Jarvelin, Zi-Bing Jin, Karl-Heinz Jöckel, Peter K. Joshi, Pekka Jousilahti, J. Wouter Jukema, Mika Kähönen, Yoichiro Kamatani, Kui Dong Kang, Jaakko Kaprio, Sharon L. R. Kardia, Fredrik Karpe, Norihiro Kato, Frank Kee, Thorsten Kessler, Amit V. Khera, Chiea Chuen Khor, Lambertus A. L. M. Kiemeney, Bong-Jo Kim, Eung Kweon Kim, Hyung-Lae Kim, Paulus Kirchhof, Mika Kivimaki, Woon-Puay Koh, Heikki A. Koistinen, Genovefa D. Kolovou, Jaspal S. Kooner, Charles Kooperberg, Anna Köttgen, Peter Kovacs, Adriaan Kraaijeveld, Peter Kraft, Ronald M. Krauss, Meena Kumari, Zoltan Kutalik, Markku Laakso, Leslie A. Lange, Claudia Langenberg, Lenore J. Launer, Loic Le Marchand, Hyejin Lee, Nanette R. Lee, Terho Lehtimäki, Huaixing Li, Liming Li, Wolfgang Lieb, Xu Lin, Lars Lind, Allan Linneberg, Ching-Ti Liu, Jianjun Liu, Markus Loeffler, Barry London, Steven A. Lubitz, Stephen J. Lye, David A. Mackey, Reedik Mägi, Patrik K. E. Magnusson, Gregory M. Marcus, Pedro Marques Vidal, Nicholas G. Martin, Winfried März, Fumihiko Matsuda, Robert W. McGarrah, Matt McGue, Amy Jayne McKnight, Sarah E. Medland, Dan Mellström, Andres Metspalu, Braxton D. Mitchell, Paul Mitchell, Dennis O. Mook-Kanamori, Andrew D. Morris, Lorelei A. Mucci, Patricia B. Munroe, Mike A. Nalls, Saman Nazarian, Amanda E. Nelson, Matt J. Neville, Christopher Newton-Cheh, Christopher S. Nielsen, Markus M. Nöthen, Claes Ohlsson, Albertine J. Oldehinkel, Lorena Orozco, Katja Pahkala, Päivi Pajukanta, Colin N. A. Palmer, Esteban J. Parra, Cristian Pattaro, Oluf Pedersen, Craig E. Pennell, Brenda W. J. H. Penninx, Louis Perusse, Annette Peters, Patricia A. Peyser, David J. Porteous, Danielle Posthuma, Chris Power, Peter P. Pramstaller, Michael A. Province, Qibin Qi, Jia Qu, Daniel J. Rader, Olli T. Raitakari, Sarju Ralhan, Loukianos S. Rallidis, Dabeeru C. Rao, Susan Redline, Dermot F. Reilly, Alexander P. Reiner, Sang Youl Rhee, Paul M. Ridker, Michiel Rienstra, Samuli Ripatti, Marylyn D. Ritchie, Dan M. Roden, Frits R. Rosendaal, Jerome I. Rotter, Igor Rudan, Femke Rutters, Charumathi Sabanayagam, Danish Saleheen, Veikko Salomaa, Nilesh J. Samani, Dharambir K. Sanghera, Naveed Sattar, Börge Schmidt, Helena Schmidt, Reinhold Schmidt, Matthias B. Schulze, Heribert Schunkert, Laura J. Scott, Rodney J. Scott, Peter Sever, Eric J. Shiroma, M. Benjamin Shoemaker, Xiao-Ou Shu, Eleanor M. Simonsick, Mario Sims, Jai Rup Singh, Andrew B. Singleton, Moritz F. Sinner, J. Gustav Smith, Harold Snieder, Tim D. Spector, Meir J. Stampfer, Klaus J. Stark, David P. Strachan, Leen M. ‘t Hart, Yasuharu Tabara, Hua Tang, Jean-Claude Tardif, Thangavel A. Thanaraj, Nicholas J. Timpson, Anke Tönjes, Angelo Tremblay, Tiinamaija Tuomi, Jaakko Tuomilehto, Maria-Teresa Tusié-Luna, Andre G. Uitterlinden, Rob M. van Dam, Pim van der Harst, Nathalie Van der Velde, Cornelia M. van Duijn, Natasja M. van Schoor, Veronique Vitart, Uwe Völker, Peter Vollenweider, Henry Völzke, Niels H. Wacher-Rodarte, Mark Walker, Ya Xing Wang, Nicholas J. Wareham, Richard M. Watanabe, Hugh Watkins, David R. Weir, Thomas M. Werge, Elisabeth Widen, Lynne R. Wilkens, Gonneke Willemsen, Walter C. Willett, James F. Wilson, Tien-Yin Wong, Jeong-Taek Woo, Alan F. Wright, Jer-Yuarn Wu, Huichun Xu, Chittaranjan S. Yajnik, Mitsuhiro Yokota, Jian-Min Yuan, Eleftheria Zeggini, Babette S. Zemel, Wei Zheng, Xiaofeng Zhu, Joseph M. Zmuda, Alan B. Zonderman, John-Anker Zwart, Gabriel Cuellar Partida, Yan Sun, Damien Croteau-Chonka, Judith M. Vonk, Loic Le Marchand, Daniel I. Chasman, Yoon Shin Cho, Iris M. Heid, Mark I. McCarthy, Maggie C. Y. Ng, Christopher J. O’Donnell, Fernando Rivadeneira, Unnur Thorsteinsdottir, Yan V. Sun, E. Shyong Tai, Michael Boehnke, Panos Deloukas, Anne E. Justice, Cecilia M. Lindgren, Ruth J. F. Loos, Karen L. Mohlke, Kari E. North, Kari Stefansson, Robin G. Walters, Thomas W. Winkler, Kristin L. Young, Po-Ru Loh, Jian Yang, Tõnu Esko, Themistocles L. Assimes, Adam Auton, Goncalo R. Abecasis, Cristen J. Willer, Adam E. Locke, Sonja I. Berndt, Guillaume Lettre, Timothy M. Frayling, Yukinori Okada, Andrew R. Wood, Peter M. Visscher, Joel N. Hirschhorn
Common single-nucleotide polymorphisms (SNPs) are predicted to collectively explain 40–50% of phenotypic variation in human height, but identifying the specific variants and associated regions requires huge sample sizes1. Here, using data from a genome-wide association study of 5.4 million individuals of diverse ancestries, we show that 12,111 independent SNPs that are significantly associated with height account for nearly all of the common SNP-based heritability. These SNPs are clustered within 7,209 non-overlapping genomic segments with a mean size of around 90 kb, covering about 21% of the genome. The density of independent associations varies across the genome and the regions of increased density are enriched for biologically relevant genes. In out-of-sample estimation and prediction, the 12,111 SNPs (or all SNPs in the HapMap 3 panel2) account for 40% (45%) of phenotypic variance in populations of European ancestry but only around 10–20% (14–24%) in populations of other ancestries. Effect sizes, associated regions and gene prioritization are similar across ancestries, indicating that reduced prediction accuracy is likely to be explained by linkage disequilibrium and differences in allele frequency within associated regions. Finally, we show that the relevant biological pathways are detectable with smaller sample sizes than are needed to implicate causal genes and variants. Overall, this study provides a comprehensive map of specific genomic regions that contain the vast majority of common height-associated variants. Although this map is saturated for populations of European ancestry, further research is needed to achieve equivalent saturation in other ancestries. © 2022, The Author(s).
Institute for Molecular Bioscience, The University of Queensland, Brisbane, QLD, Australia; Division of Endocrinology, Boston Children’s Hospital, Boston, MA, United States; Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, United States; William Harvey Research Institute, Barts and the London School of Medicine and Dentistry, Queen Mary University of London, London, United Kingdom; Harvard Medical School, Boston, MA, United States; Laboratory for Statistical Analysis, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan; Department of Statistical Genetics, Osaka University Graduate School of Medicine, Osaka, Japan; Divisions of Genetics and Rheumatology, Brigham and Women’s Hospital and Department of Medicine, Harvard Medical School, Boston, MA, United States; Department of Epidemiology, Gillings School of Global Public Health, University of North Carolina at Chapel Hill, Chapel Hill, NC, United States; COPSAC, Copenhagen Prospective Studies on Asthma in Childhood, Herlev and Gentofte Hospital, University of Copenhagen, Copenhagen, Denmark; Section for Bioinformatics, Department of Health Technology, Technical University of Denmark, Copenhagen, Denmark; 23andMe, Sunnyvale, CA, United States; Department of Veterans Affairs, Eastern Colorado Healthcare System, Aurora, CO, United States; Division of Biomedical Informatics and Personalized Medicine, University of Colorado Anschutz Medical Campus, Aurora, CO, United States; Division of Cancer Epidemiology and Genetics, National Cancer Institute, Rockville, MD, United States; Department of Internal Medicine, Division of Cardiology, University of Michigan, Ann Arbor, MI, United States; Division of Genetics, Department of Medicine, Brigham and Women’s Hospital, Boston, MA, United States; Department of Medicine, Harvard Medical School, Boston, MA, United States; Department of Genetics, University of North Carolina at Chapel Hill, Chapel Hill, NC, United States; Department of Biostatistics and Epidemiology, School of Public Health and Health Sciences, University of Massachusetts, Amherst, MA, United States; Department of Biostatistics and Center for Statistical Genetics, University of Michigan School of Public Health, Ann Arbor, MI, United States; Department of Biostatistics and Data Science, Wake Forest School of Medicine, Winston-Salem, NC, United States; Big Data Institute, Li Ka Shing Centre for Health Information and Discovery, University of Oxford, Oxford, United Kingdom; Genetics of Complex Traits, College of Medicine and Health, University of Exeter, Exeter, United Kingdom; Center for Health Data Science, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark; Wellcome Centre for Human Genetics, Nuffield Department of Medicine, University of Oxford, Oxford, United Kingdom; Nuffield Department of Population Health, University of Oxford, Oxford, United Kingdom; Institute of Genomics, Estonian Genome Centre, University of Tartu, Tartu, Estonia; Department of Internal Medicine, Erasmus MC, University Medical Center Rotterdam, Rotterdam, Netherlands; Division of Biostatistics and Epidemiology, RTI International, Durham, NC, United States; Center for Primary Care and Public Health, University of Lausanne, Lausanne, Switzerland; Swiss Institute of Bioinformatics, Lausanne, Switzerland; Saw Swee Hock School of Public Health, National University of Singapore and National University Health System, Singapore, Singapore; Department of Psychology, University of Minnesota, Minneapolis, MN, United States; Steno Diabetes Center Copenhagen, Herlev, Denmark; Department of Biology, The Bioinformatics Center, University of Copenhagen, Copenhagen, Denmark; Department of Ophthalmology, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan; Department of Family Medicine, University of California, San Diego, La Jolla, CA, United States; Department of Human Genetics, David Geffen School of Medicine at UCLA, Los Angeles, CA, United States; Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark; Department of Epidemiology, University of Groningen, University Medical Center Groningen, Groningen, Netherlands; Department of Bioinformatics, Isfahan University of Medical Sciences, Isfahan, Iran; Institute of Biological Psychiatry, Mental Health Services, Copenhagen University Hospital, Copenhagen, Denmark; Thurston Arthritis Research Center, University of North Carolina at Chapel Hill, Chapel Hill, NC, United States; Genomic Research on Complex diseases (GRC-Group), CSIR-Centre for Cellular and Molecular Biology, Hyderabad, India; Department of Epidemiology, University of Michigan School of Public Health, Ann Arbor, MI, United States; Institute for Molecular Medicine Finland (FIMM), HiLIFE, University of Helsinki, Helsinki, Finland; Molecular Genetics Section, Center for Precision Health Research, National Human Genome Research Institute, National Institutes of Health, Bethesda, MD, United States; Center for Applied Genomics, Children’s Hospital of Philadelphia, Philadelphia, PA, United States; Quantinuum Research, Wayne, PA, United States; Department of Genetics, University of Pennsylvania, Philadelphia, PA, United States; Department of Cardiovascular Sciences, University of Leicester, Leicester, United Kingdom; NIHR Leicester Biomedical Research Centre, Glenfield Hospital, Leicester, United Kingdom; Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, WA, United States; Department of Clinical Immunology, Copenhagen University Hospital, Rigshospitalet, Copenhagen, Denmark; NovoNordic Center for Protein Research, Copenhagen University, Copenhagen, Denmark; Department of Medicine, Brigham and Women’s Hospital, Boston, MA, United States; Division of Epidemiology, Department of Medicine, Vanderbilt University Medical Center, Nashville, TN, United States; Centre for Genomic and Experimental Medicine, Institute of Genetics and Cancer, University of Edinburgh, Edinburgh, United Kingdom; Centre for Public Health, Queen’s University of Belfast, Belfast, United Kingdom; Institute for Maternal and Child Health – IRCCS, Burlo Garofolo, Trieste, Italy; Ocular Epidemiology, Singapore Eye Research Institute, Singapore National Eye Centre, Singapore, Singapore; Department of Ophthalmology, National University of Singapore and National University Health System, Singapore, Singapore; Institute of Biomedical Sciences, Academia Sinica, Taipei, Taiwan; Graduate Institute of Medical Genomics and Proteomics, Medical College, National Taiwan University, Taipei, Taiwan; Department of Internal Medicine, National Taiwan University Hospital, Taipei, Taiwan; Department of Pathology and Laboratory Medicine, University of Pennsylvania, Philadelphia, PA, United States; Center for Spatial and Functional Genomics, Division of Human Genetics, Children’s Hospital of Philadelphia, Philadelphia, PA, United States; Cardiovascular Disease Initiative, Broad Institute of MIT and Harvard, Cambridge, MA, United States; Institute of Population Health Sciences, National Health Research Institutes, Zhunan, Taiwan; Department of Kinesiology, Faculty of Medicine, Université Laval, Québec City, QC, Canada; Diamantina Institute, The University of Queensland, Brisbane, QLD, Australia; Division of Preventive Medicine, Brigham and Women’s Hospital, Boston, MA, United States; Division of Statistical Genomics, Department of Genetics, Washington University School of Medicine, St Louis, MO, United States; Institute of Clinical Molecular Biology, Christian-Albrechts University of Kiel, Kiel, Germany; Vth Department of Medicine, Medical Faculty Mannheim, Heidelberg University, Mannheim, Germany; Dipartimento di Scienze Mediche Chirurgiche e Sperimentali, Università degli Studi di Sassari, Sassari, Italy; Department of Epidemiology, Erasmus MC, University Medical Center Rotterdam, Rotterdam, Netherlands; Section of Statistical Multi-omics, Department of Clinical and Experimental Medicine, University of Surrey, Guildford, United Kingdom; Department of Biostatistics, Boston University School of Public Health, Boston, MA, United States; Center for Data and Computational Sciences, VA Boston Healthcare System, Boston, MA, United States; Department of Genetic Epidemiology, University of Regensburg, Regensburg, Germany; Department of Internal Medicine II, University Hospital Regensburg, Regensburg, Germany; Department of Nutrition and Dietetics, School of Health and Education, Harokopio University of Athens, Athens, Greece; Center for Precision Medicine, Wake Forest School of Medicine, Medical Center Boulevard, Winston-Salem, NC, United States; Genome Institute of Singapore, Agency for Science, Technology and Research, Singapore, Singapore; Health Services and Systems Research, Duke-NUS Medical School, Singapore, Singapore; Institute of Human Genetics, Universitätsklinikum Erlangen, Friedrich-Alexander-Universität Erlangen-Nürnberg, Erlangen, Germany; Institute of Cardiovascular Science, Faculty of Population Health, University College London, London, United Kingdom; Survey Research Center, Institute for Social Research, University of Michigan, Ann Arbor, MI, United States; Computational Genomics Department, National Institute of Genomic Medicine, Mexico City, Mexico; Institute of Genetic Epidemiology, Medical University of Innsbruck, Innsbruck, Austria; Department of Medicine, Surgery and Health Sciences, University of Trieste, Trieste, Italy; Institute of Medical Informatics and Statistics, Kiel University, Kiel, Germany; Eurac Research, Institute for Biomedicine, Affiliated Institute of the University of Lübeck, Bolzano, Italy; Radboud University Medical Center, Radboud Institute for Health Sciences, Department for Health Evidence, Nijmegen, Netherlands; Jackson Heart Study, Department of Medicine, University of Mississippi, Jackson, MS, United States; Nanjing University of Chinese Medicine, Nanjing, China; Research Unit of Molecular Epidemiology, Institute of Epidemiology, Helmholtz Zentrum München Research Center for Environmental Health, Neuherberg, Germany; Institute of Epidemiology, Helmholtz Zentrum München Research Center for Environmental Health, Neuherberg, Germany; Department of Epidemiology Research, Statens Serum Institut, Copenhagen, Denmark; Cardiovascular Medicine, Radcliffe Department of Medicine, University of Oxford, John Radcliffe Hospital, Oxford, United Kingdom; Genetic Epidemiology, QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia; Department of Biomedicine (Human Genetics) and iSEQ Center, Aarhus University, Aarhus, Denmark; The Lundbeck Foundation Initiative for Integrative Psychiatric Research, iPSYCH, Aarhus, Denmark; BiRC—Bioinformatics Research Centre, Aarhus University, Aarhus, Denmark; The Institute for Translational Genomics and Population Sciences, Department of Pediatrics, The Lundquist Institute for Biomedical Innovation at Harbor-UCLA Medical Center, Torrance, CA, United States; Department of Medical Sciences, Uppsala University, Uppsala, Sweden; Division of Public Health Sciences, Fred Hutchinson Cancer Research Center, Seattle, WA, United States; Danish Headache Center, Department of Neurology, Copenhagen University Hospital, Rigshospitalet, Copenhagen, Rigshospitalet, Denmark; Department of Public Health and Welfare, Finnish Institute for Health and Welfare, Helsinki, Finland; MRC Integrative Epidemiology Unit, University of Bristol, Bristol, United Kingdom; Bristol Dental School, University of Bristol, Bristol, United Kingdom; Department of Neurology, Boston University School of Medicine, Boston, MA, United States; Framingham Heart Study, Framingham, MA, United States; Department of Genetics and Bioinformatics, Dasman Diabetes Institute, Kuwait City, Kuwait; Department of Clinical Sciences in Malmö, Lund University, Malmö, Sweden; Veterans Affairs Boston Healthcare System, Boston, MA, United States; Clinical Division of Neurogeriatrics, Department of Neurology, Medical University of Graz, Graz, Austria; Institute for Medical Informatics, Statistics and Documentation, Medical University of Graz, Graz, Austria; School of Medicine and Public Health, University of Newcastle, Callaghan, NSW, Australia; Institute for Medical Informatics, Statistics and Epidemiology, University of Leipzig, Medical Faculty, Leipzig, Germany; LIFE Research Center for Civilization Diseases, University of Leipzig, Medical Faculty, Leipzig, Germany; Department of Biological Psychology, Behaviour and Movement Sciences, Vrije Universiteit Amsterdam, Amsterdam, Netherlands; Department of Epidemiology, Harvard T.H. Chan School of Public Health, Boston, MA, United States; School of Public Health and Emergency Management, Southern University of Science and Technology, Shenzhen, China; Institute for Global Health and Development, Peking University, Beijing, China; Programs in Metabolism and Medical and Population Genetics, Broad Institute of Harvard and MIT, Cambridge, MA, United States; Departamento de Medicina Genómica y Toxicología Ambiental, Instituto de Investigaciones Biomédicas, Universidad Nacional Autónoma de México Ciudad Universitaria, Mexico City, Mexico; Unidad de Biología Molecular y Medicina Genómica, Instituto Nacional de Ciencias Médicas y Nutrición, Mexico City, Mexico; Division of Endocrine and Metabolism, Tri-Service General Hospital Songshan Branch, Taipei, Taiwan; Shanghai Institute of Nutrition and Health, University of Chinese Academy of Sciences, Chinese Academy of Sciences, Shanghai, China; Division of Genome Science, Department of Precision Medicine, National Institute of Health, Cheongju, South Korea; Biomedical Technology Research Center, Tokushima Research Institute, Otsuka Pharmaceutical Co., Tokushima, Japan; Research Institute, National Center for Global Health and Medicine, Tokyo, Japan; Department of Molecular Epidemiology, German Institute of Human Nutrition Potsdam-Rehbruecke, Nuthetal, Germany; German Center for Diabetes Research (DZD), Neuherberg, Germany; Department of Complex Trait Genetics, Center for Neurogenomics and Cognitive Research, Amsterdam Neuroscience, Vrije Universiteit Amsterdam, Amsterdam, Netherlands; Department of Child and Adolescent Psychiatry and Pediatric Psychology, Section Complex Trait Genetics, Amsterdam Neuroscience, Vrije Universiteit Medical Center, Amsterdam, Netherlands; Department of Biobank Research, Umeå University, Umeå, Sweden; Department of Odontology, Umeå University, Umeå, Sweden; Institute of Molecular and Clinical Ophthalmology Basel, Basel, Switzerland; Privatpraxis Prof Jonas und Dr Panda-Jonas, Heidelberg, Germany; Department of Ophthalmology, Medical Faculty Mannheim, Heidelberg University, Mannheim, Germany; Beijing Institute of Ophthalmology, Beijing Tongren Eye Center, Beijing Tongren Hospital, Capital Medical University, Beijing Ophthalmology and Visual Sciences Key Laboratory, Beijing, China; Center for Clinical Research and Prevention, Copenhagen University Hospital - Bispebjerg and Frederiksberg, Copenhagen, Denmark; Department of Public Health, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark; Faculty of Medicine, University of Kelaniya, Ragama, Sri Lanka; Department of Geriatric and General Medicine, Osaka University Graduate School of Medicine, Suita, Japan; Center for Genomic Medicine, Kyoto University Graduate School of Medicine, Kyoto, Japan; Department of Psychiatry, University of Pennsylvania, Philadelphia, PA, United States; Centre for Global Health, Usher Institute, University of Edinburgh, Edinburgh, United Kingdom; Centre for Cardiovascular Sciences, Queens Medical Research Institute, University of Edinburgh, Edinburgh, United Kingdom; Medical Research Institute, Kangbuk Samsung Hospital, Sungkyunkwan University School of Medicine, Seoul, South Korea; Department of Clinical Research Design and Evaluation (SAIHST), Sungkyunkwan University, Seoul, South Korea; SYNLAB MVZ Humangenetik Mannheim, Mannheim, Germany; Department of Clinical Sciences, Genetic and Molecular Epidemiology Unit, Lund University, Malmö, Sweden; Department of Computer Science, University of Toronto, Toronto, ON, Canada; Department of Anthropology, University of Toronto at Mississauga, Mississauga, ON, Canada; Genomics Research Centre, Centre for Genomics and Personalised Health, School of Biomedical Sciences, Queensland University of Technology, Kelvin Grove, QLD, Australia; Oneomics, Soonchunhyang Mirai Medical Center, Bucheon-si, South Korea; Laboratory of Neurogenetics, National Institute on Aging, National Institutes of Health, Bethesda, MD, United States; Center for Alzheimer’s and Related Dementias, National Institutes of Health, Bethesda, MD, United States; Data Tecnica International, Glen Echo, MD, United States; Cancer Genomics Research Laboratory, Leidos Biomedical Research, Rockville, MD, United States; Department of Population and Quantitative Health Sciences, Case Western Reserve University, Cleveland, OH, United States; Department of Medicine, University of Massachusetts Chan Medical School, Worcester, MA, United States; Center for Geriatrics and Gerontology, Taichung Veterans General Hospital, Taichung, Taiwan; Montreal Heart Institute, Montreal, QC, Canada; Departments of Ophthalmology and Human Genetics, Radboud University Nijmegen Medical Center, Nijmegen, Netherlands; MRC Epidemiology Unit, University of Cambridge School of Clinical Medicine, Cambridge, United Kingdom; Department of Clinical Science, Center for Diabetes Research, University of Bergen, Bergen, Norway; Department of Clinical Sciences, Lund University Diabetes Centre, Malmö, Sweden; Department of Clinical Chemistry, Fimlab Laboratories, Tampere, Finland; Department of Clinical Chemistry, Finnish Cardiovascular Research Center - Tampere, Faculty of Medicine and Health Technology, Tampere University, Tampere, Finland; Department of Cardiology, Heart Center, Tampere University Hospital, Tampere, Finland; National and Kapodistrian University of Athens, Dromokaiteio Psychiatric Hospital, Athens, Greece; Department of Twin Research and Genetic Epidemiology, King’s College London, London, United Kingdom; NIHR Biomedical Research Centre at Guy’s and St Thomas’ Foundation Trust, London, United Kingdom; Center for Public Health Genomics, University of Virginia School of Medicine, Charlottesville, VA, United States; MRC Human Genetics Unit, Institute of Genetics and Cancer, University of Edinburgh, Western General Hospital, Edinburgh, United Kingdom; Department of Psychiatry and Department of Community Health and Epidemiology, Dalhousie University, Halifax, NS, Canada; Institute of Psychiatric Phenomics and Genomics (IPPG), University Hospital, LMU Munich, Munich, Germany; Institute for Medical Informatics, Biometry and Epidemiology, University Hospital Essen, Essen, Germany; Diabetes Unit, Massachusetts General Hospital, Boston, MA, United States; Center for Genomic Medicine, Massachusetts General Hospital, Boston, MA, United States; Department of Psychiatry, Amsterdam Public Health and Amsterdam Neuroscience, Amsterdam UMC and Vrije Universiteit, Amsterdam, Netherlands; Biomedical and Translational Informatics Institute, Geisinger, Danville, PA, United States; Department of Genetics, Institute for Biomedical Informatics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, United States; MRC Population Health Research Unit, Nuffield Department of Population Health, University of Oxford, Oxford, United Kingdom; Population Health Sciences, Bristol Medical School, University of Bristol, Bristol, United Kingdom; Institute for Cardiogenetics, University of Lübeck, DZHK (German Research Centre for Cardiovascular Research) partner site Hamburg/Lübeck/Kiel and University Heart Center Lübeck, Lübeck, Germany; Public Health Informatics Unit, Department of Integrated Health Sciences, Nagoya University Graduate School of Medicine, Nagoya, Japan; Centre for Bone and Arthritis Research, Department of Internal Medicine and Clinical Nutrition, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, Gothenburg, Sweden; Bioinformatics Core Facility, Sahlgrenska Academy, University of Gothenburg, Gothenburg, Sweden; Korea Institute of Science and Technology, Gangneung Institute of Natural Products, Gangneung, South Korea; Department of Clinical Biochemistry, Lillebaelt Hospital, Kolding, Denmark; Department of Human Genetics, Wellcome Sanger Institute, Hinxton, United Kingdom; Department of Internal Medicine, Section of Gerontology and Geriatrics, Leiden University Medical Center, Leiden, Netherlands; Institute of Genetics and Biophysics A. Buzzati-Traverso, CNR, Naples, Italy; Department of Clinical Biochemistry and Immunology, Hospital of Southern Jutland, Aabenraa, Denmark; The National Centre for Register-based Research, University of Aarhus, Aarhus, Denmark; Centre for Population Health Research, University of Turku and Turku University Hospital, Turku, Finland; Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku, Finland; Medical School, University of Split, Split, Croatia; Algebra University College, Zagreb, Croatia; The Charles Bronfman Institute for Personalized Medicine, Icahn School of Medicine at Mount Sinai, New York, NY, United States; Department of Environmental Medicine and Public Health, Icahn School of Medicine at Mount Sinai, New York, NY, United States; Aragon Institute of Engineering Research, University of Zaragoza, Zaragoza, Spain; Centro de Investigación Biomédica en Red en Bioingeniería, Biomateriales y Nanomedicina (CIBER-BBN), Madrid, Spain; Center for Non-Communicable Diseases, Karachi, Pakistan; Department of Psychiatry, Interdisciplinary Center Psychopathology and Emotion Regulation, University of Groningen, University Medical Center Groningen, Groningen, Netherlands; Institute of Translational Genomics, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, Germany; Oxford Centre for Diabetes, Endocrinology and Metabolism, Radcliffe Department of Medicine, University of Oxford, Churchill Hospital, Oxford, United Kingdom; Hunter Medical Research Institute, New Lambton Heights, NSW, Australia; School of Medicine and Public Health, College of Health, Medicine and Wellbeing, The University of Newcastle, New Lambton Heights, NSW, Australia; IRCCS Neuromed, Pozzilli, Italy; Department of Medicine, Division of Endocrinology, Diabetes and Nutrition, University of Maryland School of Medicine, Baltimore, MD, United States; Program for Personalized and Genomic Medicine, University of Maryland School of Medicine, Baltimore, MD, United States; Unit of Genomics of Complex Diseases, Sant Pau Biomedical Research Institute (IIB Sant Pau), Barcelona, Spain; Cardiovascular Medicine Unit, Department of Medicine, Karolinska Institutet, Center for Molecular Medicine, Stockholm, Sweden; Laboratory of Epidemiology and Population Sciences, National Institute on Aging, National Institutes of Health, Baltimore, MD, United States; Department of Biomedical Science, Hallym University, Chuncheon, South Korea; Istituto di Ricerca Genetica e Biomedica, Consiglio Nazionale delle Ricerche (CNR), Cagliari, Italy; Department of Cell and Chemical Biology, Leiden University Medical Center, Leiden, Netherlands; Epidemiology and Data Science, Amsterdam UMC, location Vrije Universiteit Amsterdam, Amsterdam, Netherlands; Amsterdam Cardiovascular Sciences, Amsterdam, Netherlands; Department of Clinical Epidemiology, Leiden University Medical Center, Leiden, Netherlands; Icelandic Heart Association, Kópavogur, Iceland; Wellcome Sanger Institute, Hinxton, United Kingdom; deCODE Genetics/Amgen, Reykjavik, Iceland; Mohn Nutrition Research Laboratory, Department of Clinical Science, University of Bergen, Bergen, Norway; Department of Public Health Sciences, Parkinson School of Health Sciences and Public Health, Loyola University Chicago, Maywood, IL, United States; VA Palo Alto Health Care System, Palo Alto, CA, United States; Department of Medicine, Stanford University School of Medicine, Stanford, CA, United States; Institute for Community Medicine, University Medicine Greifswald, Greifswald, Germany; DZHK (German Centre for Cardiovascular Research), partner site Greifswald, Greifswald, Germany; Cardiology Division, Department of Pediatrics, University of California, San Francisco, Oakland, CA, United States; Centre for Cancer Genetic Epidemiology, University of Cambridge, Cambridge, United Kingdom; Department of Public Health and Primary Care, University of Cambridge, Cambridge, United Kingdom; Department of Cardiology, Leiden University Medical Center, Leiden, Netherlands; Central Diagnostics Laboratory, Division Laboratories, Pharmacy and Biomedical Genetics, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands; Department of Human Genetics, Leiden University Medical Center, Leiden, Netherlands; Laboratory Genetic Metabolic Diseases, Department of Clinical Chemistry, Amsterdam UMC, University of Amsterdam, Amsterdam, Netherlands; Core Facility Metabolomics, Amsterdam UMC, University of Amsterdam, Amsterdam, Netherlands; Department of Cardiology, Division Heart and Lungs, University Medical Center Utrecht, Utrecht University, Utrecht, Netherlands; Department of Cardiology, University of Groningen, University Medical Center Groningen, Groningen, Netherlands; Department of Epidemiology and Biostatistics, School of Public Health, Tongji Medical College, Huazhong University of Science and Technology, Wuhan, China; NIHR Barts Cardiovascular Biomedical Research Centre, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London, United Kingdom; Department of Ophthalmology, Beijing Tongren Hospital, Capital Medical University, Beijing, China; Department of Epidemiology and Biostatistics, MRC-PHE Centre for Environment and Health, School of Public Health, Imperial College London, London, United Kingdom; Department of Dermatology, Medical University of Vienna, Vienna, Austria; Department of Research and Innovation, Division of Clinical Neuroscience, Oslo University Hospital, Oslo, Norway; Department of Neurology, Oslo University Hospital, Oslo, Norway; MRC Unit for Lifelong Health and Ageing at UCL, Institute of Cardiovascular Science, University College London, London, United Kingdom; Institute of Genetic Epidemiology, Faculty of Medicine and Medical Center, University of Freiburg, Freiburg, Germany; Department of Medicine IV – Nephrology and Primary Care, Faculty of Medicine and Medical Center, University of Freiburg, Freiburg, Germany; Institute of Molecular Medicine, McGovern Medical School, University of Texas Health Science Center at Houston, Houston, TX, United States; Department of Medical Biochemistry, Kurume University School of Medicine, Kurume, Japan; Rush Alzheimer’s Disease Center, Rush University Medical Center, Chicago, IL, United States; Department of Neurological Sciences, Rush University Medical Center, Chicago, IL, United States; Department of Genetic Medicine, Weill Cornell Medicine-Qatar, Doha, Qatar; Department of Computer and Systems Engineering, Alexandria University, Alexandria, Egypt; Department of Cardiology, German Heart Centre Munich, Technical University Munich, Munich, Germany; Department of Cardiology, Ealing Hospital, London North West University Healthcare NHS Trust, London, United Kingdom; Department of Epidemiology and Biostatistics, Imperial College London, London, United Kingdom; Cardiovascular Epidemiology Unit, Department of Public Health and Primary Care, University of Cambridge, Strangeways Research Laboratory, Cambridge, United Kingdom; Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, MI, United States; Analytic and Translational Genetics Unit, Massachusetts General Hospital, Boston, MA, United States; Stanley Center for Psychiatric Research, Broad Institute of MIT and Harvard, Cambridge, MA, United States; Department of Nutrition, Gillings School of Global Public Health, University of North Carolina at Chapel Hill, Chapel Hill, NC, United States; Carolina Population Center, University of North Carolina at Chapel Hill, Chapel Hill, NC, United States; Department of Clinical Genetics, Erasmus MC, Rotterdam, Netherlands; Department of Radiology and Nuclear Medicine, Erasmus MC, Rotterdam, Netherlands; Latin American Brain Health (BrainLat), Universidad Adolfo Ibáñez, Santiago, Chile; Unidad de Investigacion de Enfermedades Metabolicas and Direction of Nutrition, Instituto Nacional de Ciencias Medicas y Nutricion, Mexico City, Mexico; Escuela de Medicina y Ciencias de la Salud, Tecnologico de Monterrey, Monterrey, Mexico; Department of Epidemiology and Dean’s Office, College of Public Health, University of Kentucky, Lexington, KY, United States; Institute of Cardiovascular Science, Faculty of Population Health Sciences, University College London, London, United Kingdom; Health Data Research UK and Institute of Health Informatics, University College London, London, United Kingdom; KG Jebsen Center for Genetic Epidemiology, Department of Public Health and Nursing, Faculty of Medicine and Health Sciences, Norwegian University of Science and Technology, NTNU, Trondheim, Norway; HUNT Research Centre, Department of Public Health and Nursing, Norwegian University of Science and Technology, Levanger, Norway; Department of Endocrinology, Clinic of Medicine, St. Olavs Hospital, Trondheim University Hospital, Trondheim, Norway; Department of Nephrology, University Hospital Regensburg, Regensburg, Germany; Geriatric Unit, Azienda Toscana Centro, Florence, Italy; Systems Genomics Laboratory, School of Biotechnology, Jawaharlal Nehru University (JNU), New Delhi, India; Institute of Molecular Genetics, National Research Council of Italy, Pavia, Italy; Human Genetics Center and Department of Epidemiology, University of Texas Health Science Center at Houston, Houston, TX, United States; Department of Nephrology and Rheumatology, Kliniken Südostbayern, Regensburg, Germany; KfH Kidney Center Traunstein, Traunstein, Germany; Center for Genomics and Personalized Medicine (CGPM), Aarhus University, Aarhus, Denmark; Bioinformatics Research Centre, Aarhus University, Aarhus, Denmark; USC-Office of Population Studies Foundation, University of San Carlos, Cebu City, Philippines; Department of Nutrition and Dietetics, University of San Carlos, Cebu City, Philippines; Human Genomics Laboratory, Pennington Biomedical Research Center, Baton Rouge, LA, United States; Department of Biochemistry, Wake Forest School of Medicine, Medical Center Boulevard, Winston-Salem, NC, United States; Department of Clinical Biochemistry, Lillebaelt Hospital, Vejle, Denmark; Institute of Regional Health Research, University of Southern Denmark, Odense, Denmark; Clinic of Medicine, St. Olavs Hospital, Trondheim University Hospital, Trondheim, Norway; Lee Kong Chian School of Medicine, Nanyang Technological University, Singapore, Singapore; Imperial College Healthcare NHS Trust, Imperial College London, London, United Kingdom; Adjunct Faculty, JSS University Academy of Higher Education and Research (JSSAHER), JSS (Deemed to be) University, Mysuru, India; Ophthalmology and Visual Sciences Academic Clinical Program (Eye ACP), Duke-NUS Medical School, Singapore, Singapore; Department of Medical Genetics, Oslo University Hospital, Oslo, Norway; Department of Medical Research, Bærum Hospital, Vestre Viken Hospital Trust, Gjettum, Norway; Department of Neurology, Division of Vascular Neurology, University of Maryland School of Medicine, Baltimore, MD, United States; Baltimore Veterans Affairs Medical Center, Department of Neurology, Baltimore, MD, United States; Unidad de Investigación Médica en Bioquímica, Hospital de Especialidades, Centro Médico Nacional Siglo XXI, Instituto Mexicano del Seguro Social, Mexico City, Mexico; Dipartimento di Scienze Biomediche, Università degli Studi di Sassari, Sassari, Italy; Intermountain Heart Institute, Intermountain Medical Center, Murray, UT, United States; Department of Surgery, University of Pennsylvania, Philadelphia, PA, United States; Corporal Michael J. Crescenz VA Medical Center, Philadelphia, PA, United States; Department of Vascular Surgery, University Medical Center Utrecht, University of Utrecht, Utrecht, Netherlands; Department of Human Nutrition, Wageningen University, Wageningen, Netherlands; Center for Translational and Computational Neuroimmunology, Department of Neurology, Columbia University Medical Center, New York, NY, United States; Department of Oncology, University of Cambridge, Cambridge, United Kingdom; Department of General Practice, Amsterdam Public Health Institute, Amsterdam UMC, location VUmc, Amsterdam, Netherlands; Department of Nutrition, Harvard T.H. Chan School of Public Health, Boston, MA, United States; Cardiac Arrhythmia Service, Massachusetts General Hospital, Boston, MA, United States; Cardiovascular Research Center, Massachusetts General Hospital, Boston, MA, United States; Department of Clinical Sciences in Malmö, Division of Geriatric Medicine, Lund University, Malmö, Sweden; Molecular Cardiology Division, Victor Chang Cardiac Research Institute, Darlinghurst, NSW, Australia; Cardiology Department, St Vincent’s Hospital, Darlinghurst, NSW, Australia; Faculty of Medicine, UNSW Sydney, Kensington, NSW, Australia; Translational Gerontology Branch, National Institute on Aging, National Institutes of Health, Baltimore, MD, United States; Robertson Center for Biostatistics, University of Glasgow, Glasgow, United Kingdom; Human Genetics Center, School of Public Health, University of Texas Health Science Center at Houston, Houston, TX, United States; Department of Public Health and Clinical Medicine, Umeå University, Umeå, Sweden; Department of Internal Medicine, Wake Forest School of Medicine, Medical Center Boulevard, Winston-Salem, NC, United States; Faculty of Veterinary and Agricultural Science, University of Melbourne, Parkville, VIC, Australia; Agriculture Victoria Research, Department of Jobs, Precincts and Regions, Bundoora, VIC, Australia; Injury Prevention Research Center, University of North Carolina at Chapel Hill, Chapel Hill, NC, United States; Division of Physical Therapy, University of North Carolina at Chapel Hill, Chapel Hill, NC, United States; Centro de Investigacion en Salud Poblacional Instituto Nacional de Salud Publica and Centro de Estudios en Diabetes, Cuernavaca, Mexico; Division of Human Genetics, Children’s Hospital of Philadelphia, Philadelphia, PA, United States; Departments of Pediatrics and Genetics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, United States; Division of Endocrinology and Diabetes, Children’s Hospital of Philadelphia, Philadelphia, PA, United States; Faculty of Medicine, University of Iceland, Reykjavik, Iceland; Department of Preventive Medicine, Keck School of Medicine of USC, Los Angeles, CA, United States; Department of Pediatrics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, United States; Division of Pulmonary Medicine, Children’s Hospital of Philadelphia, Philadelphia, PA, United States; Institute of Biomedical and Clinical Science, University of Exeter Medical School, Exeter, United Kingdom; Cardiovascular Health Research Unit, Department of Epidemiology, University of Washington, Seattle, WA, United States; Department of Paediatrics, Yong Loo Lin School of Medicine, National University of Singapore, Singapore, Singapore; Khoo Teck Puat - National University Children’s Medical Institute, National University Health System, Singapore, Singapore; Department of Internal Medicine II, Division of Cardiology, Medical University of Vienna, Vienna, Austria; Menzies Research Institute Tasmania, University of Tasmania, Hobart, TAS, Australia; Centre for Eye Research Australia, Royal Victorian Eye and Ear Hospital, University of Melbourne, Melbourne, VIC, Australia; Lions Eye Institute, Centre for Ophthalmology and Vision Science, University of Western Australia, Perth, WA, Australia; Cardiology Division, Massachusetts General Hospital, Boston, MA, United States; Department of Genetics, Shanghai-MOST Key Laboratory of Heath and Disease Genomics, Chinese National Human Genome Center and Shanghai Industrial Technology Institute, Shanghai, China; Department of Internal Medicine, University of Utah, Salt Lake City, UT, United States; Australian Centre for Precision Health, Clinical and Health Sciences, University of South Australia, Adelaide, SA, Australia; South Australian Health and Medical Research Institute, Adelaide, SA, Australia; Department of Environmental and Preventive Medicine, Jichi Medical University School of Medicine, Shimotsuke, Japan; Department of Epidemiology and Population Health, Albert Einstein College of Medicine, Bronx, NY, United States; Division of Endocrinology, Diabetes and Metabolism, School of Medicine, Ohio State University, Columbus, OH, United States; Center for Life Course Health Research, Faculty of Medicine, University of Oulu, Oulu, Finland; Unit of Primary Health Care, Oulu University Hospital, OYS, Oulu, Finland; Department of Life Sciences, College of Health and Life Sciences, Brunel University London, Uxbridge, United Kingdom; The Eye Hospital, School of Ophthalmology and Optometry, Wenzhou Medical University, Wenzhou, China; Einthoven Laboratory for Experimental Vascular Medicine, LUMC, Leiden, Netherlands; Netherlands Heart Institute, Utrecht, Netherlands; Department of Clinical Physiology, Tampere University Hospital, Tampere, Finland; Department of Clinical Physiology, Finnish Cardiovascular Research Center - Tampere, Faculty of Medicine and Health Technology, Tampere University, Tampere, Finland; Laboratory of Complex Trait Genomics, Department of Computational Biology and Medical Sciences, Graduate School of Frontier Sciences, The University of Tokyo, Tokyo, Japan; Department of Ophthalmology, The Catholic University of Korea Incheon St. Mary’s Hospital, Incheon, South Korea; NIHR Oxford Biomedical Research Centre, Churchill Hospital, Oxford, United Kingdom; German Centre for Cardiovascular Research (DZHK), partner site Munich Heart Alliance, Munich, Germany; Radboud University Medical Center, Radboud Institute for Health Sciences, Department of Urology, Nijmegen, Netherlands; Corneal Dystrophy Research Institute, Yonsei University College of Medicine, Seoul, South Korea; Saevit Eye Hospital, Goyang, South Korea; Department of Biochemistry, College of Medicine, Ewha Womans University, Seoul, South Korea; Department of Cardiology, University Heart and Vascular Center UKE Hamburg, Hamburg, Germany; Institute of Cardiovascular Sciences, College of Medical and Dental Sciences, University of Birmingham, Birmingham, United Kingdom; German Center for Cardiovascular Research, partner site Hamburg/Kiel/Lübeck, Hamburg, Germany; Atrial Fibrillation NETwork, Münster, Germany; Department of Epidemiology and Public Health, UCL Institute of Epidemiology and Health Care, University College London, London, United Kingdom; Healthy Longevity Translational Research Programme, Yong Loo Lin School of Medicine, National University of Singapore, Singapore, Singapore; University of Helsinki and Department of Medicine, Helsinki University Hospital, Helsinki, Finland; Minerva Foundation Institute for Medical Research, Helsinki, Finland; Department of Preventive Cardiology, Lipoprotein Apheresis Unit and Lipid Disorders Clinic, Metropolitan Hospital, Athens, Greece; MRC-PHE Centre for Environment and Health, Imperial College London, London, United Kingdom; National Heart and Lung Institute, Imperial College London, London, United Kingdom; Medical Department III – Endocrinology, Nephrology, Rheumatology, University of Leipzig Medical Center, Leipzig, Germany; Institute for Social and Economic Research, University of Essex, Colchester, United Kingdom; Institute of Clinical Medicine, Internal Medicine, University of Eastern Finland and Kuopio University Hospital, Kuopio, Finland; Department of Medicine, University of Colorado at Denver, Aurora, CO, United States; Berlin Institute of Health at Charité – Universitätsmedizin Berlin, Berlin, Germany; Epidemiology Program, University of Hawaii Cancer Center, Honolulu, HI, United States; Department of Internal Medicine, Ewha Womans University School of Medicine, Seoul, South Korea; Department of Epidemiology and Biostatistics, Peking University Health Science Center, Beijing, China; Peking University Center for Public Health and Epidemic Preparedness and Response, Beijing, China; Institute of Epidemiology and Biobank Popgen, Kiel University, Kiel, Germany; Key Laboratory of Systems Health Science of Zhejiang Province, Hangzhou Institute for Advanced Study, University of Chinese Academy of Sciences, Chinese Academy of Sciences, Hangzhou, China; Department of Clinical Medicine, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark; Division of Cardiovascular Medicine and Abboud Cardiovascular Research Center, University of Iowa Hospitals and Clinics, Iowa City, IA, United States; Alliance for Human Development, Lunenfeld-Tanenbaum Research Institute, Sinai Health System, Toronto, ON, Canada; Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden; Division of Cardiology, University of California, San Francisco, San Francisco, CA, United States; Department of Medicine, Internal Medicine, Lausanne University Hospital, Lausanne, Switzerland; University of Lausanne, Lausanne, Switzerland; SYNLAB Academy, SYNLAB Holding Deutschland, Mannheim, Germany; Clinical Institute of Medical and Chemical Laboratory Diagnostics, Medical University of Graz, Graz, Austria; Department of Medicine, Division of Cardiology, Duke University School of Medicine, Durham, NC, United States; Duke Molecular Physiology Institute, Duke University School of Medicine, Durham, NC, United States; Psychiatric Genetics, QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia; Geriatric Medicine, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, Gothenburg, Sweden; Geriatrics Research and Education Clinical Center, Baltimore Veterans Administration Medical Center, Baltimore, MD, United States; Centre for Vision Research and Department of Ophthalmology, Westmead Millennium Institute of Medical Research, University of Sydney, Sydney, NSW, Australia; Department of Public Health and Primary Care, Leiden University Medical Center, Leiden, Netherlands; Usher Institute of Population Health Sciences and Informatics, University of Edinburgh, Edinburgh, United Kingdom; Electrophysiology Section, Division of Cardiovascular Medicine, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, United States; Department of Medicine, University of North Carolina at Chapel Hill, Chapel Hill, NC, United States; Department of Chronic Diseases, Norwegian Institute of Public Health, Oslo, Norway; Department of Pain Management and Research, Oslo University Hospital, Oslo, Norway; Institute of Human Genetics, School of Medicine and University Hospital Bonn, Bonn, Germany; Sahlgrenska University Hospital, Department of Drug Treatment, Gothenburg, Sweden; Laboratorio de Inmunogenómica y Enfermedades Metabólicas, Instituto Nacional de Medicina Genómica, CDMX, Mexico City, Mexico; Paavo Nurmi Centre, Sports and Exercise Medicine Unit, Department of Physical Activity and Health, University of Turku, Turku, Finland; Institute for Precision Health, David Geffen School of Medicine at UCLA, Los Angeles, CA, United States; Pat MacPherson Centre for Pharmacogenetics and Pharmacogenomics, Division of Population Health and Genomics, School of Medicine, University of Dundee, Ninewells Hospital and Medical School, Dundee, United Kingdom; Centre Nutrition, Santé et Société (NUTRISS), Institute of Nutrition and Functional Foods, Université Laval, Québec City, QC, Canada; IBE, LMU Munich, Neuherberg, Germany; Population, Policy and Practice, UCL Great Ormond Street Hospital Institute of Child Health, London, United Kingdom; Department of Medicine, University of Pennsylvania, Philadelphia, PA, United States; Department of Clinical Physiology and Nuclear Medicine, Turku University Hospital, Turku, Finland; Hero DMC Heart Institute, Dyanand Medical College, Ludhiana, India; Second Department of Cardiology, Medical School, National and Kapodistrian University of Athens, University General Hospital Attikon, Athens, Greece; Division of Biostatistics, Washington University School of Medicine, St Louis, MO, United States; Genetics, Merck Sharp & Dohme, Kenilworth, NJ, United States; Department of Epidemiology, University of Washington, Seattle, WA, United States; Department of Endocrinology and Metabolism, Kyung Hee University School of Medicine, Seoul, South Korea; Department of Public Health, Clinicum, Faculty of Medicine, University of Helsinki, Helsinki, Finland; Departments of Medicine, Pharmacology, and Biomedical Informatics, Vanderbilt University Medical Center, Nashville, TN, United States; Department of Epidemiology and Data Science, Amsterdam Public Health Institute, Amsterdam Cardiovascular Sciences Institute, Amsterdam UMC, location VUmc, Amsterdam, Netherlands; Department of Cardiology and Department of Medicine, Columbia University, New York, NY, United States; Department of Pediatrics, Section of Genetics, College of Medicine, University of Oklahoma Health Sciences Center, Oklahoma City, OK, United States; Department of Pharmaceutical Sciences, University of Oklahoma Health Sciences Center, Oklahoma City, OK, United States; Department of Physiology, University of Oklahoma Health Sciences Center, Oklahoma City, OK, United States; Oklahoma Center for Neuroscience, University of Oklahoma Health Sciences Center, Oklahoma City, OK, United States; Institute of Cardiovascular and Medical Sciences, University of Glasgow, Glasgow, United Kingdom; Gottfried Schatz Research Center (for Cell Signaling, Metabolism and Aging), Medical University of Graz, Graz, Austria; Institute of Nutritional Science, University of Potsdam, Nuthetal, Germany; Deutsches Herzzentrum München, Cardiology, Deutsches Zentrum für Herz- und Kreislaufforschung (DZHK) – Munich Heart Alliance, and Technische Universität München, München, Germany; School of Biomedical Science and Pharmacy, University of Newcastle, New Lambton Heights, NSW, Australia; Department of Medicine, Vanderbilt University Medical Center, Nashville, TN, United States; Central University of Punjab, Bathinda, India; Department of Medicine I, University Hospital, LMU Munich, Munich, Germany; Department of Cardiology, Clinical Sciences, Lund University and Skåne University Hospital, Lund, Sweden; The Wallenberg Laboratory, Department of Molecular and Clinical Medicine, Institute of Medicine, Gothenburg University and the Department of Cardiology, Sahlgrenska University Hospital, Gothenburg, Sweden; Wallenberg Center for Molecular Medicine and Lund University Diabetes Center, Lund University, Lund, Sweden; Population Health Research Institute, St George’s, University of London, London, United Kingdom; Molecular Epidemiology Section, Department of Biomedical Data Sciences, Leiden University Medical Center, Leiden, Netherlands; Department of Genetics, Stanford University School of Medicine, Stanford, CA, United States; Department of Medicine, Faculty of Medicine, Université de Montréal, Montreal, QC, Canada; Helsinki University Central Hospital, Research Program for Clinical and Molecular Metabolism, University of Helsinki, Helsinki, Finland; Folkhälsan Research Center, Helsinki, Finland; Department of Public Health, University of Helsinki, Helsinki, Finland; Diabetes Research Group, King Abdulaziz University, Jeddah, Saudi Arabia; Unidad de Biología Molecular y Medicina Genómica, Instituto de Investigaciones Biomédicas, UNAM, Mexico City, Mexico; Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán, Mexico City, Mexico; Yong Loo Lin School of Medicine, National University of Singapore and National University Health System, Singapore, Singapore; Milken Institute School of Public Health, The George Washington University, Washington, DC, United States; Department Geriatric Medicine, Amsterdam Public Health, Amsterdam UMC location University of Amsterdam, Amsterdam, Netherlands; Department of Epidemiology and Data Science, Amsterdam UMC, Amsterdam, Netherlands; Interfaculty Institute for Genetics and Functional Genomics, University Medicine Greifswald, Greifswald, Germany; Unidad de Investigación Médica en Epidemiología Clínica, Hospital de Especialidades, Centro Médico Nacional Siglo XXI, Instituto Mexicano del Seguro Social, Mexico City, Mexico; Institute of Cellular Medicine, Newcastle University, Newcastle upon Tyne, United Kingdom; Department of Population and Public Health Sciences, Keck School of Medicine of USC, Los Angeles, CA, United States; Department of Physiology and Neuroscience, Keck School of Medicine of USC, Los Angeles, CA, United States; USC Diabetes and Obesity Research Institute, Keck School of Medicine of USC, Los Angeles, CA, United States; Lundbeck Foundation Center for GeoGenetics, GLOBE Institute, University of Copenhagen, Copenhagen, Denmark; School of Chinese Medicine, China Medical University, Taichung, Taiwan; Diabetes Unit, KEM Hospital and Research Centre, Pune, India; Kurume University School of Medicine, Kurume, Japan; Division of Cancer Control and Population Sciences, UPMC Hillman Cancer Center, University of Pittsburgh, Pittsburgh, PA, United States; Department of Epidemiology, Graduate School of Public Health, University of Pittsburgh, Pittsburgh, PA, United States; TUM School of Medicine, Technical University of Munich and Klinikum Rechts der Isar, Munich, Germany; Institute of Clinical Medicine, Faculty of Medicine, University of Oslo, Oslo, Norway; Department of Population Health Sciences, Geisinger, Danville, PA, United States; Vanderbilt Genetics Institute, Division of Genetic Medicine, Vanderbilt University Medical Center, Nashville, TN, United States; Department of Medicine, Veterans Affairs Boston Healthcare System, Boston, MA, United States; Department of Epidemiology, Emory University Rollins School of Public Health, Atlanta, GA, United States; Atlanta VA Health Care System, Decatur, GA, United States; Princess Al-Jawhara Al-Brahim Centre of Excellence in Research of Hereditary Disorders (PACER-HD), King Abdulaziz University, Jeddah, Saudi Arabia; The Mindich Child Health and Development Institute, Icahn School of Medicine at Mount Sinai, New York, NY, United States; School of Life Sciences, Westlake University, Hangzhou, China; Westlake Laboratory of Life Sciences and Biomedicine, Hangzhou, China; Department of Human Genetics, University of Michigan, Ann Arbor, MI, United States; McDonnell Genome Institute and Department of Medicine, Washington University School of Medicine, St Louis, MO, United States; Laboratory of Statistical Immunology, Immunology Frontier Research Center (WPI-IFReC), Osaka, Japan; Integrated Frontier Research for Medical Science Division, Institute for Open and Transdisciplinary Research Initiatives, Osaka University, Osaka, Japan; Programs in Metabolism and Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, United States; Departments of Pediatrics and Genetics, Harvard Medical School, Boston, MA, United States; Department of Mathematics and Statistics, St Cloud State University, St Cloud, MN, United States; Genentech, South San Francisco, CA, United States; Laboratory for Systems Genetics, RIKEN Center for Integrative Medical Sciences, Kanagawa, Japan; Department of Genome Informatics, Graduate School of Medicine, The University of Tokyo, Tokyo, Japan
