Zari Dastani, Marie-France Hivert, Nicholas J. Timpson, Xin Yuan, Leo-Pekka Lyytikäinen, Toshiko Tanaka, Andrew P. Morris, Aaron Isaacs, Kurt Lohman, Lu Qi, Stavroula Kanoni, James S. Pankow, Jonna Grimsby, Ching-Ti Liu, Vincent E. Mooser, John Chambers, Nicholas J. Wareham, Claudia Langenberg, Rune Frants, Thomas W. Winkler, Bruce M. Psaty, Ida Chen, Jorma Viikari, Mika Kähönen, Peter P. Pramstaller, David M. Evans, Beate St. Pourcain, Andrew R. Wood, Stefania Bandinelli, Josephine M. Egan, Stefan Böhringer, Diana van Heemst, Marja-Liisa Nuotio, Melissa Garcia, Margot Haun, Panos Deloukas, David J. Couper, Jose C. Florez, MAGIC Consortium, James G. Wilson, Xiuqing Guo, Toby Johnson, Tanya M. Teslovich, Ingrid Meulenbelt, George V. Dedoussis, Frank B. Hu, Yongmei Liu, Bernhard Paulweber, Timothy D. Spector, Timothy M. Frayling, George Davey Smith, David S. Siscovick, Florian Kronenberg, Cornelia van Duijn, Dawn M. Waterworth, James B. Meigs, J. Brent Richards, Benjamin F. Voight, Laura J. Scott, Valgerdur Steinthorsdottir, Christian Dina, Ryan P. Welch, Eleftheria Zeggini, Cornelia Huth, Yurii S. Aulchenko, Gudmar Thorleifsson, Laura J. McCulloch, Teresa Ferreira, Soumya Raychaudhuri, Amanda J. Bennett, Roza Blagieva, Eric Boerwinkle, Kristina Bengtsson Boström, Suzannah Bumpstead, Noël P. Burtt, Peter S. Chines, Todd Green, Christopher J. Groves, Candace Guiducci, Samy Hadjadj, Neelam Hassanali, Bo Isomaa, Anne U. Jackson, Augustine Kong, Man Li, Cecilia M. Lindgren, Michel Marre, Thomas Meitinger, Mario A. Morken, Katharine R. Owen, Felicity Payne, Christine Proenca, Inga Prokopenko, Michael Roden, Michael J. Sampson, Richa Saxena, Thomas Sparsø, Heather M. Stringham, Barbara Thorand, Rob M. van Dam, G. Bragi Walters, Michael N. Weedon, Stephane Cauchi, Albert Hofman, Andrew D. Morris, Eric J. G. Sijbrands, Lincoln D. Stein, Andre Uitterlinden, Richard M. Watanabe, Mark J. Daly, Leif Groop, Unnur Thorsteinsdottir, David Altshuler, Reedik Mägi, Denis Rybin, Anuj Goel, Erik Ingelsson, Jian’an Luan, Amanda Elliott, Praveen Sethupathy, Yavuz Ariyurek, Sven Bergmann, Amélie Bonnefond, Eric Brunner, Robert Clarke, Lachlan J. M. Coin, Ian N. M. Day, Eco J. C. de Geus, Jerome Delplanque, Nita G. Forouhi, Mark O. Goodarzi, Rhian Gwilliam, Naomi Hammond, Caroline Hayward, Simon C. Heath, Aroon D. Hingorani, Jennie Hui, Mika Kivimaki, Seppo Koskinen, Kirsten Ohm Kyvik, Debbie A. Lawlor, Yun Li, Massimo Mangino, María Teresa Martínez-Larrad, Jarred B. McAteer, Christa Meisinger, David Meyre, Braxton D. Mitchell, Silvia Naitza, Matthew J. Neville, Cristian Pattaro, Daniel Pearson, John F. Peden, Irene Pichler, Simon C. Potter, Samuli Ripatti, Fernando Rivadeneira, Olov Rolandsson, Manjinder Sandhu, Paul Scheet, Stephen J. Sharp, Gunnar Sigurðsson, Angela Silveira, Laila Simpson, Ulla Sovio, Amy Swift, Dhiraj Varma, Veronique Vitart, Peter J. Wagner, Hugh Watkins, Gonneke Willemsen, Jing Hua Zhao, Ingrid B Borecki, Patrik K. E. Magnusson, David M. Nathan, Gordon H Williams, Joachim Spranger, Fredrik Karpe, Manuel Serrano-Ríos, Lyle J Palmer, Alan F Wright, Michael Stumvoll, Jerome I Rotter, Dorret I Boomsma, Antonio Cao, Angelo Scuteri, David Schlessinger, Aimo Ruokonen, Marjo-Riitta Jarvelin, Leena Peltonen, Kiran Musunuru, Christopher J. O’Donnell, James P. Pirruccello, Gina M. Peloso, Sally L. Ricketts, Xiaohui Li, Jong-Young Lee, Kyunga Kim, Joshua D. Smith, Robert Y. L. Zee, John B. Whitfield, Ida Surakka, Juha Sinisalo, Juha Saharinen, Lynda M. Rose, Mark Rieder, Alex N. Parker, Christopher J. O’Donnell, David Masson, Nicholas G. Martin, Fabio Marroni, Robert Luben, Marja-Liisa Lokki, Guillaume Lettre, Edward G. Lakatta, Inke R. König, Kay-Tee Khaw, Lee M. Kaplan, A. Cecile W. Janssens, G. Kees Hovingh, Nicholas D. Hastie, Elena Gonzalez, Nelson B. Freimer, Jeanette Erdmann, Kenechi G. Ejebe, Serkalem Demissie, S. Matthijs Boekholdt, Mark Seielstad, Tien Y. Wong, Stacey B. Gabriel, Vilmundur Gudnason, Muredach P. Reilly, Heribert Schunkert, L. Adrienne Cupples
Circulating levels of adiponectin, a hormone produced predominantly by adipocytes, are highly heritable and are inversely associated with type 2 diabetes mellitus (T2D) and other metabolic traits. We conducted a meta-analysis of genome-wide association studies in 39,883 individuals of European ancestry to identify genes associated with metabolic disease. We identified 8 novel loci associated with adiponectin levels and confirmed 2 previously reported loci (P = 4.5×10−8- 1.2 ×10−43). Using a novel method to combine data across ethnicities (N = 4,232 African Americans, N = 1,776 Asians, and N = 29,347 Europeans), we identified two additional novel loci. Expression analyses of 436 human adipocyte samples revealed that mRNA levels of 18 genes at candidate regions were associated with adiponectin concentrations after accounting for multiple testing (p<3×10−4). We next developed a multi-SNP genotypic risk score to test the association of adiponectin decreasing risk alleles on metabolic traits and diseases using consortia-level meta-analytic data. This risk score was associated with increased risk of T2D (p = 4.3×10−3, n = 22,044), increased triglycerides (p = 2.6×10−14, n = 93,440), increased waist-to-hip ratio (p = 1.8×10−5, n = 77,167), increased glucose two hours post oral glucose tolerance testing (p = 4.4×10−3, n = 15,234), increased fasting insulin (p = 0.015, n = 48,238), but with lower in HDL- cholesterol concentrations (p = 4.5×10−13, n = 96,748) and decreased BMI (p = 1.4×10−4, n = 121,335). These findings identify novel genetic determinants of adiponectin levels, which, taken together, influence risk of T2D and markers of insulin resistance. © 2012, Dastani et al.
Department of Epidemiology, Biostatistics,and Occupational Health, Jewish General Hospital, Lady Davis Institute, McGill University, Montreal, Canada; Department of Medicine, Université de Sherbrooke, Sherbrooke, Canada; General Medicine Division, Massachusetts General Hospital, Boston, MA, United States; MRC CAiTE Centre and School of Social and Community Medicine, University of Bristol, Bristol, United Kingdom; Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom; Genetics of Complex Traits, Peninsula Medical School, University of Exeter, Exeter, United Kingdom; Genetics, GlaxoSmithKline, King of Prussia, Pennsylvania, United States; MRC Epidemiology Unit, Institute of Metabolic Science, Addenbrooke’s Hospital, Cambridge, United Kingdom; Department of Human Genetics, Leiden University Medical Center, Leiden, Netherlands; Department of Epidemiology and Preventive Medicine, Regensburg University Medical Center, Regensburg, Germany; Departments of Medicine and Public Health, Weill Cornell Medical College, New York, NY, United States; Department of Clinical Chemistry, University of Tampere and Tampere University Hospital, Tampere, Finland; Center for Statistical Genetics, Department of Biostatistics, University of Michigan, Ann Arbor, MI, United States; Clinical Research Branch, National Institute on Aging, Baltimore, MD, United States; Department of Twin Research and Genetic Epidemiology, King’s College London, London, United Kingdom; Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, United Kingdom; Genetic Epidemiology Unit, Department of Epidemiology, Erasmus Medical Center, Rotterdam, Netherlands; Centre for Medical Systems Biology, Leiden, Netherlands; Section of Molecular Epidemiology, Leiden University Medical Center and The Netherlands Genomics Initiative, The Netherlands Consortium for Healthy Aging, Leiden, Netherlands; Division of Genetic Epidemiology, Innsbruck Medical University, Innsbruck, Austria; Wake Forest University School of Medicine, Winston-Salem, NC, United States; Harvard School of Public Health, Boston, MA, United States; Division of Epidemiology and Community Health, University of Minnesota, Minneapolis, MN, United States; Medical Statistics and Bioinformatics, Leiden University Medical Center, Leiden, Netherlands; Department of Genetics, University of North Carolina, Chapel Hill, NC, United States; Cardiovascular Research Institute, Morehouse School of Medicine, Atlanta, GA, United States; Department of Preventive Medicine, Feinberg School of Medicine, Northwestern University, Chicago, IL, United States; Lady Davis Institute for Medical Research, Department of Oncology, McGill University, Montreal, Canada; Department of Human Genetics McGill University, Montreal, Canada; Department of Medicine, Harvard Medical School, Boston, MA, United States; Department of Biostatistics, Boston University School of Public Health, Boston, MA, United States; Cardiology, Ealing Hospital National Health Service (NHS) Trust, London, United Kingdom; Department of Internal Medicine, University of Lausanne, Lausanne, Switzerland; Department of Medical Genetics, University of Lausanne, Lausanne, Switzerland; Epidemiology and Biostatistics, Imperial College London, London, United Kingdom; Deptartment of Clinical Genetics and Department of Epidemiology, Erasmus Medical Center, Rotterdam, Netherlands; Cardiovascular Health Research Unit, Departments of Medicine and Epidemiology, University of Washington, Seattle, WA, United States; Group Health Research Institute, Group Health Cooperative, Seattle, WA, United States; Departments of Pathology and Biochemistry, University of Vermont, Burlington, VT, United States; Cardiovascular Health Research Unit, University of Washington, Seattle, WA, United States; Medical Genetics Research Institute, Cedars Sinai Medical Center, Los Angeles, CA, United States; Department of Medicine, University of Turku and Turku University Hospital, Turku, Finland; Department of Clinical Physiology, University of Tampere and Tampere University Hospital, Tampere, Finland; Center for Biomedicine, European Academy Bozen/Bolzano (EURAC), Bolzano, Italy; Department of Neurology, General Central Hospital, Bolzano, Italy; Department of Neurology, University of Lübeck, Lübeck, Germany; School of Social and community medicine, University of Bristol, Bristol, United Kingdom; British Heart Foundation Glasgow Cardiovascular Research Centre, University of Glasgow, Glasgow, United Kingdom; Geriatric Unit, Azienda Sanitaria Firenze (ASF), Florence, Italy; Laboratory of Clinical Investigation, National Institute of Aging, Baltimore, MD, United States; Gerontology and Geriatrics, Leiden University Medical Center, Leiden, Netherlands; First Department of Internal Medicine, St. Johann Spital, Paracelsus Private Medical University Salzburg, Salzburg, Austria; Public Health Genomics Unit, FIMM, Department of Chronic Disease Prevention, University of Helsinki, National Institute for Health and Welfare and Institute for Molecular Medicine Finland, Helsinki, Finland; Population Studies Unit, Department of Chronic Disease Prevention, National Institute for Health and Welfare, Turku, Finland; Intramural Research Program, Laboratory of Epidemiology, Demography and Biometry, National Institute on Aging, National Institutes of Health, Bethesda, MD, United States; Division of General Internal Medicine, Women’s Health Clinical Research Center, University of California San Francisco, San Francisco, CA, United States; Institute of Epidemiology, Helmholtz Zentrum München, German Research Center for Environmental Health, Munich, Germany; Institute of Medical Informatics, Biometry and Epidemiology, Ludwig-Maximilians-Universität, Munich, Germany; Klinikum Großhadern, Munich, Germany; Harokopio University, Athens, Greece; Collaborative Studies Coordinating Center, Department of Biostatistics, University of North Carolina at Chapel Hill, Chapel Hill, NC, United States; School of Medicine, Federal University of Rio Grande do Sul, Porto Alegre, Brazil; Department of Epidemiology, University of North Carolina at Chapel Hill, Chapel Hill, NC, United States; Department of Rheumatology and Department of Clinical Epidemiology, Leiden, Netherlands; Department of Nutrition, University of North Carolina, Chapel Hill, NC, United States; Office of Population Studies Foundation, University of San Carlos, Cebu City, Philippines; Department of Physiology and Biophysics, University of Mississippi Medical Center, Jackson, MS, United States; Department of Medicine, University of Mississippi Medical Center, Jackson, MS, United States; Medical Genetics Institute, Cedars-Sinai Medical Center, Los Angeles, CA, United States; University Institute of Social and Preventative Medicine, Centre Hospitalier Universitaire Vaudois (CHUV) and University of Lausanne, Lausanne, Switzerland; Swiss Institute of Bioinformatics, Lausanne, Switzerland; Metabolic Research Laboratories, Institute of Metabolic Science, University of Cambridge, Addenbrooke’s Hospital, Cambridge, United Kingdom; Department of Family and Preventive Medicine, University of California San Diego, La Jolla, CA, United States; Brigham and Women’s Hospital, Boston, MA, United States; Jackson Heart Study Coordinating Center, Jackson State University, Jackson, MS, United States; Baylor College of Medicine and Methodist DeBakey Heart and Vascular Center, Houston, TX, United States; Molecular Epidemiology, Leiden University Medical Center, Leiden, Netherlands; Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku and the Department of Clinical Physiology, Turku University Hospital, Turku, Finland; Program in Medical and Population Genetics, Broad Institute, Cambridge, MA, United States; Center for Human Genetic Research, Massachusetts General Hospital, Boston, MA, United States; Diabetes Research Center, Diabetes Unit, Massachusetts General Hospital, Boston, MA, United States; Chronic Disease Epidemiology and Prevention Unit, Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, Finland; Diabetes Prevention Unit, Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, Finland; Unit of General Practice, Helsinki University Central Hospital, Helsinki, Finland; Folkhalsan Research Centre, Helsinki, Finland; Vaasa Central Hospital, Vaasa, Finland; Department of General Practice and Primary Health Care, University of Helsinki, Helsinki, Finland; University of Washington, Seattle, WA, United States; National Heart, Lung and Blood Institute’s Framingham Heart Study, Framingham, MA, United States; Departments of Medicine, Human Genetics, Epidemiology and Biostatistics, Lady Davis Institute, Jewish General Hospital, McGill University, Montreal, Canada; Broad Institute of Harvard and Massachusetts Institute of Technology (MIT), Cambridge, 02142, MA, United States; Center for Human Genetic Research, Massachusetts General Hospital, 185 Cambridge Street, Boston, 02114, MA, United States; Department of Medicine, Harvard Medical School, Boston, 02115, MA, United States; Department of Biostatistics, University of Michigan, Ann Arbor, 48109-2029, MI, United States; deCODE Genetics, 101 Reykjavik, Iceland; Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, OX3 7BN, United Kingdom; 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Hagedorn Research Institute, Gentofte, DK-2820, Denmark; Centre Hospitalier Universitaire de Poitiers, Endocrinologie Diabetologie, CIC INSERM 0801, INSERM U927, Université de Poitiers, UFR, Médecine Pharmacie, Poitiers Cedex, 86021, France; Institute for Clinical Diabetology, German Diabetes Center, Leibniz Center for Diabetes Research at Heinrich Heine University Dü sseldorf, Dü sseldorf, 40225, Germany; Folkhä lsan Research Center, Helsinki, FIN-00014, Finland; Malmska Municipal Health Center and Hospital, Jakobstad, 68601, Finland; Diabetes Research and Wellness Foundation Human Islet Isolation Facility and Oxford Islet Transplant Programme, University of Oxford, Old Road, Headington, Oxford, OX3 7LJ, United Kingdom; Research Centre for Prevention and Health, Glostrup University Hospital, Glostrup, DK-2600, Denmark; University of Copenhagen, Copenhagen, 2200, Denmark; Department of Epidemiology, Johns Hopkins University, Baltimore, 21287, MD, United States; Department of Medicine, and Welch Center for Prevention, Epidemiology, and Clinical Research, Johns Hopkins University, Baltimore, 21287, MD, United States; 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Institut interrégional pour la Santé (IRSA), La Riche, F-37521, France; Department of Medicine, Helsinki University Hospital, University of Helsinki, Helsinki, FIN-00290, Finland; Department of Internal Medicine, University Medical Center Utrecht, Utrecht, 3584 CG, Netherlands; Molecular Genetics, Medical Biology Section, Department of Pathology and Medical Biology, University Medical Center Groningen and University of Groningen, Groningen, 9700 RB, Netherlands; Department of Genetics, University Medical Center Groningen and University of Groningen, Groningen, 9713 EX, Netherlands; Department of Physiology and Biophysics, University of Southern California School of Medicine, Los Angeles, 90033, CA, United States; Department of Genetics and Pathology, Rudbeck Laboratory, Uppsala University, Uppsala, S-751 85, Sweden; University of Southern Denmark, Odense, DK-5230, Denmark; Centre for Diabetes, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London, E1 2AT, United Kingdom; Department of Medicine, The Hospital of Levanger, Levanger, N-7600, Norway; Department of Genetics, University of North Carolina, Chapel Hill, 27599, NC, United States; Institute of Genetic Medicine, European Academy Bozen/Bolzano (EURAC), Viale Druso 1, Bolzano, 39100, Italy; Croatian Centre for Global Health, University of Split, Soltanska 2, Split, 21000, Croatia; Institute for Clinical Medical Research, University Hospital Sestre Milosrdnice, Vinogradska 29, Zagreb, 10000, Croatia; Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, FIN-00300, Finland; Diabetes Research Group, Institute of Cellular Medicine, Newcastle University, Framlington Place, Newcastle upon Tyne, NE2 4HH, United Kingdom; Department of Preventitive Medicine, Keck Medical School, University of Southern California, Los Angeles, 90089-9001, CA, United States; Division of Epidemiology and Community Health, University of Minnesota, Minneapolis, 55454, MN, United States; Department of Biomedical Science, Panum, University of Copenhagen, Copenhagen, 2200, Denmark; 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Centre for Genetic Epidemiology and Biostatistics, University of Western Australia, Perth, Australia; Istituto di Neurogenetica e Neurofarmacologia (INN), Consiglio Nazionale delle Ricerche, c/o Cittadella Universitaria di Monserrato, Monserrato, Cagliari, Italy; Western Australian Sleep Disorders Research Institute, Queen Elizabeth Medical Centre II, Perth, Australia; Department of Endocrinology, Diabetes and Nutrition, Charite-Universitaetsmedizin Berlin, Berlin, Germany; Department of Clinical Nutrition, German Institute of Human Nutrition Potsdam-Rehbruecke, Nuthetal, Germany; Division of Endocrinology, Diabetes, and Hypertension, Brigham and Women’s Hospital, Harvard Medical School, Boston, MA, United States; Department of Human Genetics, Leiden University Medical Centre, Leiden, Netherlands; Department of Cardiovascular Research, Istituto di Ricerche Farmacologiche Mario Negri, Milan, Italy; Institut National de la Santé et de la Recherche Médicale, Institut National de la Recherche Agronomique, Université Paris 13, Bobigny Cedex, France; Department of Medicine III, Division Prevention and Care of Diabetes, University of Dresden, Dresden, Germany; Center for Human Nutrition, University of Texas Southwestern Medical Center, Dallas, TX, United States; Department of Genetics and Pathology, Rudbeck Laboratory, Uppsala University, Uppsala, Sweden; Centre Hospitalier Universitaire, de Poitiers, Endocrinologie Diabetologie, CIC INSERM 0802, INSERM U927, Université de Poitiers, Uniteé de Formation et de Recherche, Médecine Pharmacie, Poitiers, France; Department of Public Health and Clinical Medicine, Section for Nutritional Research, Umeå University, Umeå, Sweden; Department of Clinical Sciences, Obstetrics and Gynecology, University of Oulu, Finland; Centre National de Génotypage, Institut de génomique/Commissariat aàl’énergie atomique, Evry Cedex, France; INSERM U872, Paris Cedex, France; Institute for Clinical Diabetology, German Diabetes Center, Leibniz Center for Diabetes Research at Heinrich Heine University Düsseldorf, Düsseldorf, Germany; Institute of Genetic Medicine, European Academy Bozen/Bolzano (EURAC), Viale Druso, Bolzano, Italy; Institute of the University Lübeck, Lübeck, Germany; Department of Pulmonary Physiology, Sir Charles Gairdner Hospital, Perth, Australia; Busselton Population Medical Research Foundation, Sir Charles Gairdner Hospital, Perth, Australia; Heart Institute of Western Australia, Sir Charles Gairdner Hospital, Nedlands, WA, Australia; School of Medicine and Pharmacology, University of Western Australia, Nedlands, WA, Australia; Folkhalsan Research Centre, Helsinki, Finland; Malmska Municipal Health Care Center and Hospital, Jakobstad, Finland; Nuffield Department of Surgery, University of Oxford, Oxford, United Kingdom; Research Centre for Prevention and Health, Glostrup University Hospital, Glostrup, Denmark; University of Copenhagen, Copenhagen, Denmark; National Institute for Health and Welfare, Unit of Population Studies, Turku, Finland; Institute of Health Sciences and Biocenter Oulu, University of Oulu, Oulu, Finland; Department of Public Health, University of Helsinki, Helsinki, Finland; National Institute for Health and Welfare, Unit for Child and Adolescent Mental Health, Helsinki, Finland; Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland; Department of Internal Medicine and Biocenter Oulu, Oulu, Finland; Diabetes Genetics, Institute of Biomedical and Clinical Science, Peninsula College of Medicine and Dentistry, University of Exeter, Exeter, United Kingdom; National Institute for Health and Welfare, Unit of Living Conditions, Health and Wellbeing, Helsinki, Finland; Interdisciplinary Centre for Clinical Research, University of Leipzig, Leipzig, Germany; The Danish Twin Registry, Epidemiology, Institute of Public Health, University of Southern Denmark, Odense, Denmark; Department of Clinical Sciences, Diabetes and Endocrinology, Lund University, University Hospital Malmö, Malmö, Sweden; Gladstone Institute of Cardiovascular Disease, University of California, San Francisco, CA, United States; Diabetes Research Center, Diabetes Unit, Massachusetts General Hospital, Boston, MA, United States; Department of Medicine, Harvard Medical School, Boston, MA, United States; Division of Cardiology, University of Ottawa Heart Institute, Ottawa, ON, Canada; Oxford National Institute for Health Research, Biomedical Research Centre, Churchill Hospital, Oxford, United Kingdom; Department of Clinical Genetics, Erasmus Medical College, Rotterdam, Netherlands; Biomedical Research Institute, University of Dundee, Ninewells Hospital and Medical School, Dundee, United Kingdom; Department of Geriatric Medicine and Metabolic Disease, Second University of Naples, Naples, Italy; National Institute for Health and Welfare, Unit of Public Health Genomics, Helsinki, Finland; Department of Medical Genetics, University of Helsinki, Helsinki, Finland; Department of Medical Statistics, Epidemiology and Medical Informatics, Andrija Stampar School of Public Health, Medical School, University of Zagreb, Rockefellerova, Zagreb, Croatia; Department of Clinical Genetics, VU University and Medical Center, Amsterdam, Netherlands; Department of Obstetrics and Gynaecology, Oulu University Hospital, Oulu, Finland; Departments of Medicine, Epidemiology and Health Services, University of Washington, Seattle, WA, United States; Group Health Research Institute, Group Health Cooperative, Seattle, WA, United States; Institute of Biometrics and Epidemiology, German Diabetes Centre, Leibniz Centre at Heinrich Heine University Düsseldorf, Düsseldorf, Germany; Department of Biostatistics, University of Washington, Seattle, WA, United States; Department of Internal Medicine, Erasmus Medical College, Rotterdam, Netherlands; Department of Metabolic Diseases, Heinrich Heine University Duüsseldorf, Duüsseldorf, Germany; Department of Public Health and Clinical Medicine, Section for Family Medicine, Umeå University, Umeaå, Sweden; School of Public Health, Department of General Practice, University of Aarhus, Aarhus, Denmark; Department of Public Health and Primary Care, Strangeways Research Laboratory, University of Cambridge, Cambridge, United Kingdom; MRC Epidemiology Resource Centre, University of Southampton, Southampton General Hospital, Southampton, United Kingdom; Department of Epidemiology, University of Texas, M.D. Anderson Cancer Center, Houston, TX, United States; Leibniz-Institut für Arterioskleroseforschung an der Universität Münster, Muünster, Germany; Atherosclerosis Research Unit, Department of Medicine, Karolinska Institutet, Stockholm, Sweden; Laboratory of Neurogenetics, National Institute on Aging, Bethesda, MD, United States; Department of Epidemiology, University of Washington, Seattle, WA, United States; Seattle Epidemiologic Research and Information Center, Department of Veterans Affairs Office of Research and Development, Seattle, WA, United States; Department of Medical Sciences, Uppsala University, Uppsala, Sweden; Medstar Research Institute, Baltimore, MD, United States; Clinical Research Branch, National Institute on Aging, Baltimore, MD, United States; Institut interrégional pour la santé (IRSA), La Riche, France; Coordination Centre for Clinical Trials, University of Leipzig, Leipzig, Germany; Department of Medicine, Helsinki University Hospital, University of Helsinki, Helsinki, Finland; Department of Internal Medicine, Leiden University Medical Centre, Leiden, Netherlands; Research Unit, Cardiovascular Genetics, Nancy University Henri Poincaré, Nancy, France; EMGO Institute for Health and Care Research, Department of Psychiatry, VU University Medical Center, Amsterdam, Netherlands; Department of Internal Medicine, Centre Hospitalier Universitaire Vaudois, Lausanne, Switzerland; Genomic Medicine, Imperial College London, Hammersmith Hospital, London, United Kingdom; Epidemiology and Public Health, Queen’s University Belfast, Belfast, United Kingdom; Medical Products Agency, Uppsala, Sweden; UOC Geriatria, Istituto Nazionale Ricovero e cura per Anziani (INRCA) IRCCS, Rome, Italy; National Institute for Health and Welfare, Unit of Chronic Disease Epidemiology and Prevention, Helsinki, Finland; Departments of Nutrition and Epidemiology, Harvard School of Public Health, Boston, MA, United States; Channing Laboratory, Brigham and Women’s Hospital and Harvard Medical School, Boston, MA, United States; Genetic Epidemiology and Clinical Research Group, Department of Public Health and Clinical Medicine, Section for Medicine, Umeå University Hospital, Ume, Sweden; London School of Hygiene and Tropical Medicine, London, United Kingdom; Department of Medicine, School of Medicine, Johns Hopkins University, Baltimore, MD, United States; The Welch Center for Prevention, Epidemiology, and Clinical Research, School of Medicine and Bloomberg School of Public Health, Johns Hopkins University, Baltimore, MD, United States; Division of Epidemiology and Community Health, School of Public Health, University of Minnesota, Minneapolis, MN, United States; Department of Endocrinology and Diabetes, Norfolk and Norwich University Hospital National Health Service Trust, Norwich, United Kingdom; Department of Medicine, University of Kuopio and Kuopio University Hospital, Kuopio, Finland; University of Southern Denmark, Odense, Denmark; Institute of Biomedical Science, University of Copenhagen, Copenhagen, Denmark; Department of Neurology, General Central Hospital, Bolzano, Italy; Department of Neurology, University of Lübeck, Lübeck, Germany; Institute of Medical Informatics, Biometry and Epidemiology, Ludwig-Maximilians-Universität, Munich, Germany; Klinikum Grosshadern, Munich, Germany; School of Medicine, University of Split, Split, Croatia; Gen-Info Ltd, Zagreb, Croatia; Department of Physiology and Biophysics, Keck School of Medicine, University of Southern California, Los Angeles, CA, United States; Department of Medicine, Division of Endocrinology, Keck School of Medicine, University of Southern California, Los Angeles, CA, United States; Department of Genetics, University of North Carolina, Chapel Hill, NC, United States; National Institute for Health and Welfare, Unit of Diabetes Prevention, Helsinki, Finland; South Ostrobothnia Central Hospital, Seinajoki, Finland; Departments of Medicine and Epidemiology, University of Washington, Seattle, WA, United States; Longitudinal Studies Section, Clinical Research Branch, National Institute on Aging, NIH, Baltimore, MD, United States; University of Iceland, Reykjavík, Iceland; Lab of Cardiovascular Sciences, National Institute on Aging, National Institutes of Health, Baltimore, MD, United States; Department of Clinical Sciences/Clinical Chemistry, University of Oulu, Oulu, Finland; National Institute of Health and Welfare, Oulu, Finland; Department of Preventive Medicine, Keck School of Medicine, University of Southern California, Los Angeles, CA, United States; MRC-Health Protection Agency Centre for Environment and Health, Imperial College London, London, United Kingdom; Center for Statistical Genetics, Department of Biostatistics, University of Michigan, Ann Arbor, 48109, MI, United States; Center for Human Genetic Research, Massachusetts General Hospital, Boston, 02114, MA, United States; Cardiovascular Research Center, Massachusetts General Hospital, Boston, 02114, MA, United States; Department of Medicine, Harvard Medical School, Boston, 02115, MA, United States; Broad Institute, Cambridge, 02142, MA, United States; Johns Hopkins University School of Medicine, Baltimore, 21287, MD, United States; Icelandic Heart Association, Heart Preventive Clinic and Research Institute, 201 Kopavogur, Iceland; University of Iceland, 101 Reykjavik, Iceland; Cardiovascular Institute, University of Pennsylvania School of Medicine, Philadelphia, 19104, PA, United States; Institute for Translational Medicine and Therapeutics, University of Pennsylvania School of Medicine, Philadelphia, 19104, PA, United States; Division of Molecular Medicine, Department of Medicine, Columbia University, New York, 10032, NY, United States; Institute for Molecular Medicine Finland FIMM, University of Helsinki, Helsinki, FI-00014, Finland; National Institute for Health and Welfare, P.O. Box 104, Helsinki, FI-00251, Finland; Division of Preventive Medicine, Brigham and Women’s Hospital, Boston, 02215, MA, United States; Robarts Research Institute, University of Western Ontario, London, N6A 5K8, ON, Canada; Department of Vascular Medicine, Academic Medical Centre at the University of Amsterdam, Amsterdam, 1105 AZ, Netherlands; Department of Epidemiology, Erasmus University Medical Center, P.O. Box 2040, Rotterdam, 3000 CA, Netherlands; Department of Biostatistics, Boston University School of Public Health, Boston, 02118, MA, United States; National Heart, Lung and Blood Institute’s Framingham Heart Study, Framingham, 01702, MA, United States; Human Genetics Center, University of Texas Health Science Center at Houston, Houston, 77030, TX, United States; Department of Public Health and Primary Care, Strangeways Research Laboratory, University of Cambridge, Cambridge, CB1 8RN, United Kingdom; Cardiovascular Health Research Unit and Department of Medicine, University of Washington, Seattle, 98101, WA, United States; deCODE Genetics, 101 Reykjavik, Iceland; Division of Statistical Genomics in the Center for Genome Sciences, Washington University School of Medicine, St Louis, 63108, MO, United States; Department of Epidemiology and Public Health, Imperial College London, London, W2 1PG, United Kingdom; Department of Clinical Sciences, Lund University, MalmÖ, SE-20502, Sweden; Clinical Pharmacology and Barts and the London Genome Centre, William Harvey Research Institute, Barts and the London School of Medicine, Queen Mary University of London, London, EC1M 6BQ, United Kingdom; Medical Genetics Institute, Cedars-Sinai Medical Center, Los Angeles, 90048, CA, United States; Center for Genome Science, National Institute of Health, Seoul, 122-701, South Korea; Interdisciplinary Program in Bioinformatics, College of Natural Sciences, Seoul National University, Seoul, 151-742, South Korea; Department of Statistics, College of Natural Sciences, Seoul National University, Seoul, 151-742, South Korea; Department of Statistics, Sookmyung WomenÖs University, Seoul, 140-742, South Korea; Centre for Molecular Epidemiology, National University of Singapore, Singapore, 117597, Singapore; Genome Institute of Singapore, Singapore, 138672, Singapore; Department of Genetics, University of North Carolina, Chapel Hill, 27599, NC, United States; Department of Genome Sciences, University of Washington, Seattle, 98195, WA, United States; Genetics Division, GlaxoSmithKline R and D, King of Prussia, 19406, PA, United States; MRC Epidemiology Unit, Institute of Metabolic Science, AddenbrookeÖs Hospital, Cambridge, CB2 0QQ, United Kingdom; Division of Genetic Epidemiology, Department of Medical Genetics, Molecular and Clinical Pharmacology, Innsbruck Medical University, Schoepfstrasse 41, Innsbruck, A-6020, Austria; Institut für Medizinische Biometrie und Statistik, Universität zu Lübeck, Lübeck, 23562, Germany; MRC Human Genetics Unit, Institute of Genetics and Molecular Medicine, Edinburgh, EH4 2XU, United Kingdom; Netherlands Genomics Initiative (NGI)-sponsored Netherlands Consortium for Healthy Aging (NCHA), Center of Medical Systems Biology (CMSB), RC Leiden, 2300, Netherlands; Centre for Population Health Sciences, University of Edinburgh, Edinburgh, EH8 9AG, United Kingdom; Department of Biological Psychology, VU University Amsterdam, Van der Boechorststraat 1, Amsterdam, 1081 BT, Netherlands; Institute of Epidemiology, Helmholtz Zentrum Munchen - German Research Center for Environmental Health, Neuherberg, 85764, Germany; Genetic Epidemiology Unit, Queensland Institute of Medical Research, PO Royal Brisbane Hospital, Queensland, 4029, Australia; Department of Internal Medicine, Centre Hospitalier Universitaire Vaudois, Lausanne, 1011, Switzerland; Department of Internal Medicine, Erasmus University Medical Center, PO Box 2040, CA Rotterdam, 3000, Netherlands; Istituto di Neurogenetica e Neurofarmacologia (INN), Consiglio Nazionale delle Ricerche, c/o Cittadella Universitaria di Monserrato, Monserrato, Cagliari, 09042, Italy; Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, FI-00271, Finland; Department of Health Sciences, University of Leicester, Leicester, LE1 6TP, United Kingdom; Clinical Research Branch, National Institute on Aging, National Institutes of Health, Baltimore, 21225, MD, United States; Medstar Research Institute, Baltimore, 21218, MD, United States; Department of Twin Research and Genetic Epidemiology, King’s College London, London, SE1 7EH, United Kingdom; Wellcome Trust Sanger Institute, Hinxton, Cambridge, CB10 1SA, United Kingdom; Department of Psychiatry, EMGO Institute, Neuroscience Campus Amsterdam, VU University Medical Center, MB Amsterdam, 1007, Netherlands; Division of Cardiology, Department of Medicine, Helsinki University Central Hospital (HUCH), Helsinki, FI-00029, Finland; Unita Operativa Geriatria, Istituto Nazionale Ricovero e Cura Anziani (INRCA), Istituto Ricovero e Cura a Carattere Scientifico (IRCCS), Via Cassia 1167, Rome, 00189, Italy; Hammersmith Hospital, National Heart and Lung Institute, Imperial College London, London, W12 0NN, United Kingdom; Gerontology Research Center, National Institute on Aging, 5600 Nathan Shock Drive, Baltimore, 21224, MD, United States; Department of Health Research and Policy, Stanford University, Stanford, 94305, CA, United States; Department of Clinical Chemistry, University of Oulu, Oulu, FI-90220, Finland; The John and Jennifer Ruddy Canadian Cardiovascular Genetics Centre, University of Ottawa, Ottawa, K1Y 4W7, Canada; Departments of Medicine, Epidemiology, and Health Services, University of Washington, Group Health Research Institute, Group Health Cooperative, Seattle, 98101, WA, United States; Institute of Genetic Medicine, European Academy Bozen/Bolzano (EURAC), Viale Druso 1, Bolzano, 39100, Italy; Institute of the University of Lübeck, Germany; Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, SE-17177, Sweden; Amgen, Thousand Oaks, 91320, CA, United States; Genetic and Molecular Epidemiology Laboratory, McMaster University, Hamilton, L8N3Z5, ON, Canada; Department of Clinical Genetics, Erasmus University Medical Center, CA Rotterdam, 3000, Netherlands; Institut fur Humangenetik, Helmholtz Zentrum Munchen, Deutsches Forschungszentrum fur Umwelt und Gesundheit, Neuherberg, 85764, Germany; Institute of Human Genetics, Klinikum rechts der Isar, Technische Universität München, Muenchen, 81675, Germany; Wellcome Trust Centre for Human Genetics, University of Oxford, Roosevelt Drive, Oxford, OX3 7BN, United Kingdom; Oxford Centre for Diabetes, Endocrinology and Medicine, University of Oxford, Churchill Hospital, Oxford, OX3 7LJ, United Kingdom; Oxford NIHR Biomedical Research Centre, Churchill Hospital, Oxford, OX3 7LJ, United Kingdom; Avon Longitudinal Study of Parents and Children, University of Bristol, Bristol, BS8 2BN, United Kingdom; Institute of Applied Genomics, via Linussio 51, Udine, 33100, Italy; Cardiovascular Epidemiology and Genetics, Institut Municipal d’Investigacio Medica, Barcelona, 08003, Spain; Transplantation Laboratory, Haartman Institute, University of Helsinki, Helsinki, FI-00014, Finland; Montreal Heart Institute (Research Center), Université de Montréal, Montréal, H1T 1C8, QC, Canada; Laboratory of Epidemiology, Demography, and Biometry, National Institute on Aging, National Institutes of Health, Bethesda, 20892, MD, United States; Science Center, Tampere University Hospital, Tampere, FI-33521, Finland; Institute of Regional Health Research and the Danish Twin Registry, Institute of Public Health, University of Southern Denmark, JBWinsløws Vej 9B, Odense, DK-5000, Denmark; Department of Public Health, University of Helsinki, Helsinki, FI-00014, Finland; Massachusetts General Hospital Weight Center, Boston, 02114, MA, United States; Department of Genetics and Pathology, Rudbeck Laboratory, University of Uppsala, Uppsala, SE-75185, Sweden; Department of Epidemiology and Biostatistics, Imperial College London, St Mary’s Campus, Norfolk Place, London, W2 1PG, United Kingdom; Department of Public Health Science and General Practice, University of Oulu, Oulu, FI-90220, Finland; Klinik und Poliklinik für Innere Medizin II, Universität Regensburg, Regensburg, 93053, Germany; Department of Epidemiology and Preventive Medicine Regensburg University Medical Center Franz-Josef-Strauss-Allee 11, Regensburg, 93053, Germany; Department of Biomedical Engineering and Computational Science, Aalto University School of Science and Technology, Aalto, FI-00076, Finland; LIGHT Research Institute, University of Leeds, Leeds, LS2 9JT, United Kingdom; Department of Medicine, Helsinki University Hospital, Helsinki, FI-00029, Finland; Department of Psychiatry, Center for Neurobehavioral Genetics, The Jane and Terry Semel Institute for Neuroscience and Human Behavior, David Geffen School of Medicine, University of California, Los Angeles, 90095, CA, United States; Clinical Research Branch, National Institute on Aging, National Institutes of Health, Baltimore, 21225, MD, United States; Medizinische Klinik II, Universität zu Lübeck, Lübeck, 23538, Germany; MRC-HPA Centre for Environment and Health, Imperial College London, London, W2 1PG, United Kingdom; BHF Glasgow Cardiovascular Research Centre, University of Glasgow, 126 University Place, Glasgow, G12 8TA, United Kingdom; Division of Cardiovascular Epidemiology, Institute of Environmental Medicine, Karolinska Institutet, Stockholm, SE-17177, Sweden; National Human Genome Research Institute, National Institutes of Health, Bethesda, 20892, MD, United States; Departments of Vascular Medicine and Cardiology, Academic Medical Centre, Amsterdam, 1105 AZ, Netherlands; Department of Physiology and Biophysics, University of Southern California, Los Angeles, 90033, CA, United States; Geriatric Unit, Azienda Sanitaria Firenze (ASF), Florence, 50125, Italy; Department of Medicine, Baylor College of Medicine, Houston, 77030, TX, United States; Department of Medicine, Stanford University School of Medicine, Stanford, 94305, CA, United States; Singapore Eye Research Institute, National University of Singapore, Singapore, 168751, Singapore; Departments of Medicine/Epidemiology and Public Health, Yong Loo Lin School of Medicine, National University of Singapore, Singapore, 117597, Singapore; Office of Population Studies Foundation, University of San Carlos, Cebu City, 6000, Philippines; Department of Anthropology, Northwestern University, Evanston, 60208, IL, United States; Department of Nutrition, Carolina Population Center, University of North Carolina, Chapel Hill, 27516, NC, United States; Department of Medicine, University of Mississippi Medical Center, Jackson, 39126, MS, United States; Children’s Hospital Oakland Research Institute, Oakland, 94609, CA, United States; Department of Cardiovascular Epidemiology and Population Genetics, Centro Nacional de Investigaciones Cardiovasculares, Madrid, 28029, Spain; Nutrition and Genomics Laboratory, Jean Mayer United States Department of Agriculture Human Nutrition Research Center on Aging, Tufts University, Boston, 02111, MA, United States; Clinical Pharmacology and Barts and The London Genome Centre, William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, Charterhouse Square, London, EC1M 6BQ, United Kingdom; Sage Bionetworks, Seattle, 98109, WA, United States; Division of Community Health Sciences, St George’s University of London, London, SW17 0RE, United Kingdom; Department of Cardiovascular Sciences, University of Leicester, NIHR Biomedical Research Unit in Cardiovascular Disease, Glenfield Hospital, Leicester, LE3 9QP, United Kingdom
