Abstract
Obesity is heritable and predisposes to many diseases. To understand the genetic basis of obesity better, here we conduct a genome-wide association study and Metabochip meta-analysis of body mass index (BMI), a measure commonly used to define obesity and assess adiposity, in up to 339,224 individuals. This analysis identifies 97 BMI-associated loci (P < 5 × 10−8), 56 of which are novel. Five loci demonstrate clear evidence of several independent association signals, and many loci have significant effects on other metabolic phenotypes. The 97 loci account for ∼2.7% of BMI variation, and genome-wide estimates suggest that common variation accounts for >20% of BMI variation. Pathway analyses provide strong support for a role of the central nervous system in obesity susceptibility and implicate new genes and pathways, including those related to synaptic function, glutamate signalling, insulin secretion/action, energy metabolism, lipid biology and adipogenesis.
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The genetics of obesity: from discovery to biology
Genome-wide discovery of genetic loci that uncouple excess adiposity from its comorbidities
Obesity and the risk of cardiometabolic diseases
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Acknowledgements
A full list of acknowledgements can be found in the Supplementary Information.
Author information
Jong-Young Lee
Present address: Present address: Second Floor, B-dong, AICT Building, 145 Gwanggyo-ro, Yeongyong-gu, Suwon-si, Gyeonggi-do,443-270, South Korea.,
Adam E. Locke, Bratati Kahali, Sonja I. Berndt, Anne E. Justice and Tune H. Pers: These authors contributed equally to this work.
Kari E. North, Erik Ingelsson, Joel N. Hirschhorn, Ruth J. F. Loos and Elizabeth K. Speliotes: These authors jointly supervised this work.
Biocenter Oulu, University of Oulu, FI-90014 Oulu, Finland.
Authors and Affiliations
Department of Biostatistics, Center for Statistical Genetics, University of Michigan, Ann Arbor, 48109, Michigan, USA
Adam E. Locke, Jennifer L. Bragg-Gresham, Anne U. Jackson, Heather M. Stringham, Gonçalo R. Abecasis & Michael Boehnke
Department of Internal Medicine, Division of Gastroenterology, and Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, 48109, Michigan, USA
Bratati Kahali, Corey Powell & Elizabeth K. Speliotes
Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, 20892, Maryland, USA
Sonja I. Berndt, Zhaoming Wang, Kevin B. Jacobs & Stephen J. Chanock
Department of Epidemiology, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina 27599, USA.,
Anne E. Justice, Keri L. Monda & Kari E. North
Divisions of Endocrinology and Genetics and Center for Basic and Translational Obesity Research, Boston Children’s Hospital, Boston, 02115, Massachusetts, USA
Tune H. Pers, Sailaja Vedantam, Tonu Esko, Cameron D. Palmer, Rany M. Salem & Joel N. Hirschhorn
Broad Institute of the Massachusetts Institute of Technology and Harvard University, Cambridge, 02142, Massachusetts, USA
Tune H. Pers, Sailaja Vedantam, Tonu Esko, Cameron D. Palmer, Karol Estrada, Robert E. Handsaker, Sekar Kathiresan, Steven A. McCarroll, Rany M. Salem, Cecilia M. Lindgren & Joel N. Hirschhorn
Department of Genetics, Harvard Medical School, Boston, 02115, Massachusetts, USA
Tune H. Pers, Tonu Esko, Robert E. Handsaker, Steven A. McCarroll, Rany M. Salem & Joel N. Hirschhorn
Department of Systems Biology, Center for Biological Sequence Analysis, Technical University of Denmark, Lyngby 2800, Denmark.,
Tune H. Pers
MRC Epidemiology Unit, University of Cambridge School of Clinical Medicine, Institute of Metabolic Science, Cambridge Biomedical Campus, Cambridge CB2 0QQ, UK.,
Felix R. Day, Jian’an Luan, Jing Hua Zhao, Robert A. Scott, Nita G. Forouhi, Claudia Langenberg, Ken K. Ong, Nicholas J. Wareham & Ruth J. F. Loos
Department of Genetics, University of North Carolina, Chapel Hill, 27599, North Carolina, USA
Martin L. Buchkovich, Damien C. Croteau-Chonka & Karen L. Mohlke
Queensland Brain Institute, The University of Queensland, Brisbane 4072, Australia.,
Jian Yang, Anna A. E. Vinkhuyzen & Peter M. Visscher
The University of Queensland Diamantina Institute, The Translation Research Institute, Brisbane 4012, Australia.,
Jian Yang & Peter M. Visscher
Channing Division of Network Medicine, Department of Medicine, Brigham and Women's Hospital and Harvard Medical School, Boston, 02115, Massachusetts, USA
Damien C. Croteau-Chonka, Lu Qi & David J. Hunter
Estonian Genome Center, University of Tartu, Tartu 51010, Estonia.,
Tonu Esko, Reedik Mägi, Krista Fischer, Toomas Haller, Evelin Mihailov, Lili Milani, Eva Reinmaa, Andrew P. Morris, Mari Nelis, Andres Metspalu & Markus Perola
Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm 17177, Sweden.,
Tove Fall, Henrik Grönberg, Per Hall, Patrik K. E. Magnusson & Nancy L. Pedersen
Science for Life Laboratory, Uppsala University, Uppsala 75185, Sweden.,
Tove Fall, Stefan Gustafsson, Åsa K. Hedman, Johan Ärnlöv, Ann-Christine Syvänen & Erik Ingelsson
Department of Medical Sciences, Molecular Epidemiology, Uppsala University, Uppsala 75185, Sweden.,
Tove Fall, Stefan Gustafsson, Åsa K. Hedman, Johan Ärnlöv & Erik Ingelsson
Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford OX3 7BN, UK.,
Teresa Ferreira, Reedik Mägi, Joshua C. Randall, Åsa K. Hedman, Anuj Goel, Inga Prokopenko, Thorhildur Juliusdottir, Anubha Mahajan, Nigel W. Rayner, Neil R. Robertson, Alexander W. Drong, Sarah Keildson, John R. B. Perry, Krina T. Zondervan, Martin Farrall, Andrew P. Morris, Hugh Watkins, Mark I. McCarthy, Cecilia M. Lindgren & Erik Ingelsson
Institute of Social and Preventive Medicine (IUMSP), Centre Hospitalier Universitaire Vaudois (CHUV), Lausanne 1010, Switzerland.,
Zoltán Kutalik
Swiss Institute of Bioinformatics, Lausanne 1015, Switzerland.,
Zoltán Kutalik & Jacques S. Beckmann
Department of Medical Genetics, University of Lausanne, Lausanne 1005, Switzerland.,
Zoltán Kutalik & Jacques S. Beckmann
Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1SA, UK.,
Joshua C. Randall, Nigel W. Rayner, Kathleen Stirrups, Panos Deloukas & Inês Barroso
Department of Genetic Epidemiology, Institute of Epidemiology and Preventive Medicine, University of Regensburg, D-93053 Regensburg, Germany.,
Thomas W. Winkler, Mathias Gorski & Iris M. Heid
Genetics of Complex Traits, University of Exeter Medical School, University of Exeter, Exeter EX1 2LU, UK.,
Andrew R. Wood, Dorota Pasko, Michael N. Weedon, John R. B. Perry & Timothy M. Frayling
Department of Nutrition, Harvard School of Public Health, Boston, 02115, Massachusetts, USA
Tsegaselassie Workalemahu, Lu Qi, Paul W. Franks & David J. Hunter
Survey Research Center, Institute for Social Research, University of Michigan, Ann Arbor, 48104, Michigan, USA
Jessica D. Faul & David R. Weir
Department of Epidemiology, University of Michigan, Ann Arbor, 48109, Michigan, USA
Jennifer A. Smith & Wei Zhao
Department of Internal Medicine, Division of Cardiovascular Medicine, University of Michigan, Ann Arbor, 48109, Michigan, USA
Jin Chen & Cristen J. Willer
Department of Genetics, University Medical Center Groningen, University of Groningen, 9700 RB Groningen, The Netherlands.,
Rudolf Fehrmann, Juha Karjalainen, Joanna Smolonska, Morris A. Swertz, Harm-Jan Westra, Pim van der Harst & Lude Franke
Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, 48109, Michigan, USA
Ellen M. Schmidt & Cristen J. Willer
HudsonAlpha Institute for Biotechnology, Huntsville, 35806, Alabama, USA
Devin Absher & Lindsay L. Waite
Department of Epidemiology, Genetic Epidemiology Unit, Erasmus MC University Medical Center, 3015 GE Rotterdam, The Netherlands.,
Najaf Amin, Ayse Demirkan, Aaron Isaacs, Ben A. Oostra & Cornelia M. van Duijn
Telethon Institute for Child Health Research, Centre for Child Health Research, The University of Western Australia, Perth, 6008, Western Australia, Australia
Denise Anderson
Netherlands Consortium for Healthy Aging (NCHA), Leiden University Medical Center, Leiden 2300 RC, The Netherlands.,
Marian Beekman, Stefan Böhringer, Quinta Helmer, Hae-Won Uh, Lisette Stolk & P. Eline Slagboom
Department of Molecular Epidemiology, Leiden University Medical Center, 2300 RC Leiden, The Netherlands.,
Marian Beekman & P. Eline Slagboom
Centre for Population Health Sciences, University of Edinburgh, Teviot Place, Edinburgh EH8 9AG, UK.,
Jennifer L. Bolton, Ross M. Fraser, Stela McLachlan, Harry Campbell, Jackie F. Price, Igor Rudan & James F. Wilson
Kidney Epidemiology and Cost Center, University of Michigan, Ann Arbor, 48109, Michigan, USA
Jennifer L. Bragg-Gresham
Department of Statistics & Biostatistics, Rutgers University, Piscataway, 08854, New Jersey, USA
Steven Buyske
Department of Genetics, Rutgers University, Piscataway, 08854, New Jersey, USA
Steven Buyske & Tara C. Matise
Department of Human Genetics, Leiden University Medical Center, 2333 ZC Leiden, The Netherlands.,
Ayse Demirkan
Ealing Hospital NHS Trust, Middlesex UB1 3HW, UK.,
Guohong Deng, Weihua Zhang, William R. Scott, Sian-Tsung Tan, Jaspal S. Kooner & John C. Chambers
Department of Gastroenterology and Hepatology, Imperial College London, London W2 1PG, UK.,
Guohong Deng
Institute of infectious Diseases, Southwest Hospital, Third Military Medical University, Chongqing, China
Guohong Deng
Center for Complex Disease Genomics, McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University School of Medicine, Baltimore, 21205, Maryland, USA
Georg B. Ehret & Aravinda Chakravarti
Department of Specialties of Internal Medicine, Cardiology, Geneva University Hospital, Geneva 1211, Switzerland.,
Georg B. Ehret & François Mach
Department of Epidemiology Research, Statens Serum Institut, Copenhagen DK-2300, Denmark.,
Bjarke Feenstra, Heather A. Boyd & Frank Geller
Department of Genetics, Washington University School of Medicine, St Louis, 63110, Missouri, USA
Mary F. Feitosa, E. Warwick Daw, Mary K. Wojczynski, Qunyuan Zhang, Aldi T. Kraja, D. C. Rao & Ingrid B. Borecki
Division of Cardiovacular Medicine, Radcliffe Department of Medicine, University of Oxford, Oxford OX3 9DU, UK.,
Anuj Goel, Martin Farrall & Hugh Watkins
Division of Public Health Sciences, Fred Hutchinson Cancer Research Center, Seattle, 98109, Washington, USA
Jian Gong, Jeffrey Haessler, Charles Kooperberg & Ulrike Peters
William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London EC1M 6BQ, UK.,
Stavroula Kanoni, Kathleen Stirrups & Panos Deloukas
Vth Department of Medicine (Nephrology, Hypertensiology, Endocrinology, Diabetology, Rheumatology), Medical Faculty of Mannheim, University of Heidelberg, D-68187 Mannheim, Germany.,
Marcus E. Kleber, Graciela Delgado, Tanja B. Grammer & Winfried März
Department of Internal Medicine II, Ulm University Medical Centre, D-89081 Ulm, Germany.,
Marcus E. Kleber & Wolfgang Koenig
National Institute for Health and Welfare, FI-00271 Helsinki, Finland.,
Kati Kristiansson, Niina Eklund, Leena Kinnunen, Jaana Lindström, Johan G. Eriksson, Pekka Jousilahti, Antti M. Jula, Jaakko Kaprio, Paul Knekt, Satu Männistö, Veikko Salomaa, Jarmo Virtamo, Markus Perola & Jaakko Tuomilehto
Epidemiology Program, University of Hawaii Cancer Center, Honolulu, 96813, Hawaii, USA
Unhee Lim, Lynne R. Wilkens & Loic Le Marchand
The Charles Bronfman Institute for Personalized Medicine, Icahn School of Medicine at Mount Sinai, New York, 10029, New York, USA
Vaneet Lotay, Janina M. Jeff, Yingchang Lu, Omri Gottesman, Erwin P. Bottinger & Ruth J. F. Loos
Department of Twin Research and Genetic Epidemiology, King’s College London, London SE1 7EH, UK.,
Massimo Mangino, Cristina Menni, Alireza Moayyeri, John R. B. Perry & Tim D. Spector
Department of Cardiology, University Medical Center Groningen, University of Groningen, 9700RB Groningen, The Netherlands.,
Irene Mateo Leach, Niek Verweij & Pim van der Harst
Netherlands Consortium for Healthy Aging (NCHA), 3015GE Rotterdam, The Netherlands.,
Carolina Medina-Gomez, Marjolein J. Peters, Oscar H. Franco, Albert Hofman, M. Carola Zillikens, Fernando Rivadeneira, André G. Uitterlinden & Cornelia M. van Duijn
Department of Epidemiology, Erasmus MC University Medical Center, 3015GE Rotterdam, The Netherlands.,
Carolina Medina-Gomez, Oscar H. Franco, Albert Hofman, Jacqueline C. Witteman, Fernando Rivadeneira, André G. Uitterlinden & Cornelia M. van Duijn
Department of Internal Medicine, Erasmus MC University Medical Center, 3015GE Rotterdam, The Netherlands.,
Carolina Medina-Gomez, Marjolein J. Peters, Karol Estrada, Lisette Stolk, M. Carola Zillikens, Fernando Rivadeneira & André G. Uitterlinden
QIMR Berghofer Medical Research Institute, Brisbane, Queensland 4006, Australia.,
Sarah E. Medland, Scott D. Gordon, Grant W. Montgomery, Dale R. Nyholt & Nicholas G. Martin
Laboratory of Neurogenetics, National Institute on Aging, National Institutes of Health, Bethesda, 20892, Maryland, USA
Michael A. Nalls
Institute for Medical Informatics, Biometry and Epidemiology (IMIBE), University Hospital Essen, 45147 Essen, Germany.,
Sonali Pechlivanis, Karl-Heinz Jöckel, Susanne Moebus & André Scherag
Oxford Centre for Diabetes, Endocrinology and Metabolism, University of Oxford, Oxford OX3 7LJ, UK.,
Inga Prokopenko, Amy Barrett, Amanda J. Bennett, Christopher J. Groves, Nigel W. Rayner, Neil R. Robertson, Fredrik Karpe & Mark I. McCarthy
Department of Genomics of Common Disease, School of Public Health, Imperial College London, Hammersmith Hospital, London W12 0NN, UK.,
Inga Prokopenko & Philippe Froguel
Department of Clinical Sciences, Genetic & Molecular Epidemiology Unit, Lund University Diabetes Center, Skåne University Hosptial, Malmö 205 02, Sweden.,
Dmitry Shungin, Frida Renstrom & Paul W. Franks
Department of Public Health and Clinical Medicine, Unit of Medicine, Umeå University, Umeå 901 87, Sweden.,
Dmitry Shungin & Paul W. Franks
Department of Odontology, Umeå University, Umeå 901 85, Sweden.,
Dmitry Shungin
University of Eastern Finland, FI-70210 Kuopio, Finland.,
Alena Stančáková
Department of Medicine, Atherosclerosis Research Unit, Center for Molecular Medicine, Karolinska Institutet, Stockholm 17176, Sweden.,
Rona J. Strawbridge, Lasse Folkersen, Per Eriksson, Jesper R. Gådin, Ferdinand M. van’t Hooft & Anders Hamsten
Division of Biostatistics, Washington University School of Medicine, St Louis, 63110, Missouri, USA
Yun Ju Sung, D. C. Rao & Treva K. Rice
Translational Gerontology Branch, National Institute on Aging, Baltimore, 21225, Maryland, USA
Toshiko Tanaka & Luigi Ferrucci
Interfaculty Institute for Genetics and Functional Genomics, University Medicine Greifswald, D-17475 Greifswald, Germany.,
Alexander Teumer, Georg Homuth & Uwe Völker
Department of Cardiology, Leiden University Medical Center, 2300 RC Leiden, The Netherlands.,
Stella Trompet & J. Wouter Jukema
Department of Gerontology and Geriatrics, Leiden University Medical Center, 2300 RC Leiden, The Netherlands.,
Stella Trompet & Anton J. M. de Craen
Division Heart and Lungs, Experimental Cardiology Laboratory, University Medical Center Utrecht, 3584 CX Utrecht, The Netherlands.,
Sander W. van der Laan, Hester M. den Ruijter & Gerard Pasterkamp
Department of Medical Genetics, University Medical Center Utrecht, 3584 CX Utrecht, The Netherlands.,
Jessica van Setten & Paul I. W. de Bakker
Department of Endocrinology, University of Groningen, University Medical Center Groningen, 9700 RB Groningen, The Netherlands.,
Jana V. Van Vliet-Ostaptchouk
Core Genotyping Facility, SAIC-Frederick, Inc., NCI-Frederick, Frederick, Maryland 21702, USA.,
Zhaoming Wang & Kevin B. Jacobs
CNRS UMR 8199, F-59019 Lille, France.,
Loïc Yengo, Elodie Eury, Stéphane Lobbens, Stéphane Cauchi & Philippe Froguel
European Genomic Institute for Diabetes, F-59000 Lille, France.,
Loïc Yengo, Elodie Eury, Stéphane Lobbens, Stéphane Cauchi & Philippe Froguel
Université de Lille 2, F-59000 Lille, France.,
Loïc Yengo, Elodie Eury, Stéphane Lobbens, Stéphane Cauchi & Philippe Froguel
Department of Epidemiology and Biostatistics, Imperial College London, London W2 1PG, UK.,
Weihua Zhang, William R. Scott, Marjo-Riitta Jarvelin & John C. Chambers
Center for Medical Sytems Biology, 2300 RC Leiden, The Netherlands.,
Aaron Isaacs, Ben A. Oostra & Cornelia M. van Duijn
Institute of Genetic Epidemiology, Helmholtz Zentrum München - German Research Center for Environmental Health, D-85764 Neuherberg, Germany.,
Eva Albrecht, Harald Grallert, Martina Müller-Nurasyid, Janina S. Ried, Christian Gieger, Konstantin Strauch & Iris M. Heid
School of Health and Social Studies, Dalarna University, SE-791 88 Falun, Sweden.,
Johan Ärnlöv
PathWest Laboratory Medicine of Western Australia, Nedlands, 6009, Western Australia, Australia
Gillian M. Arscott, John Beilby & Jennie Hui
Department of Haematology, University of Cambridge, Cambridge CB2 0PT, UK.,
Antony P. Attwood & Jennifer Jolley
NHS Blood and Transplant, Cambridge CB2 0PT, UK.,
Antony P. Attwood & Jennifer Jolley
Geriatric Unit, Azienda Sanitaria Firenze (ASF), 50125 Florence, Italy.,
Stefania Bandinelli
USC-Office of Population Studies Foundation, Inc., University of San Carlos, Cebu City 6000, Philippines.,
Isabelita N. Bas & Nanette R. Lee
Department of Genetics, Texas Biomedical Research Institute, San Antonio, 78227, Texas, USA
Claire Bellis & John Blangero
Genomics Research Centre, Institute of Health and Biomedical Innovation, Queensland University of Technology, Brisbane, 4001, Queensland, Australia
Claire Bellis
Department of Medical Sciences, Endocrinology, Diabetes and Metabolism, Uppsala University, Uppsala 75185, Sweden.,
Christian Berne
Division of Endocrinology, Diabetes and Metabolism, Ulm University Medical Centre, D-89081 Ulm, Germany.,
Roza Blagieva & Sigrun Merger
Integrated Research and Treatment Center (IFB) Adiposity Diseases, University of Leipzig, D-04103 Leipzig, Germany.,
Matthias Blüher, Yvonne Böttcher, Peter Kovacs, Anke Tönjes & Michael Stumvoll
Department of Medicine, University of Leipzig, D-04103 Leipzig, Germany.,
Matthias Blüher, Peter Kovacs, Anke Tönjes & Michael Stumvoll
Department of Medical Statistics and Bioinformatics, Leiden University Medical Center, 2300 RC Leiden, The Netherlands.,
Stefan Böhringer, Quinta Helmer & Hae-Won Uh
Medical Genomics and Metabolic Genetics Branch, National Human Genome Research Institute, NIH, Bethesda, Maryland 20892, USA.,
Lori L. Bonnycastle, Mario A. Morken, Amy J. Swift, Peter S. Chines & Francis S. Collins
LifeLines Cohort Study, University Medical Center Groningen, University of Groningen, 9700 RB Groningen, The Netherlands.,
Marcel Bruinenberg & Salome Scholtens
Department of Biology, Norwegian University of Science and Technology, 7491 Trondheim, Norway.,
Ida H. Caspersen & Berit Johansen
Department of Pediatrics, University of California Los Angeles, Torrance, 90502, California, USA
Yii-Der Ida Chen
Transgenomics Institute, Los Angeles Biomedical Research Institute, Torrance, 90502, California, USA
Yii-Der Ida Chen
Nuffield Department of Population Health, Clinical Trial Service Unit and Epidemiological Studies Unit, University of Oxford, Oxford OX3 7LF, UK.,
Robert Clarke
Department of Dietetics-Nutrition, Harokopio University, 17671 Athens, Greece.,
Maria Dimitriou & George Dedoussis
Medical Research Institute, University of Dundee, Ninewells Hospital and Medical School, Dundee DD1 9SY, UK.,
Alex S. F. Doney, Andrew D. Morris & Colin N. A. Palmer
Institute for Molecular Medicine, University of Helsinki, FI-00014 Helsinki, Finland.,
Niina Eklund, Jaakko Kaprio, Leif C. Groop & Markus Perola
Analytic and Translational Genetics Unit, Massachusetts General Hospital and Harvard Medical School, Boston, 02114, Massachusetts, USA
Karol Estrada & Stephan Ripke
Laboratory of Epidemiology and Population Sciences, National Institute on Aging, NIH, Bethesda, 20892, Maryland, USA
Melissa E. Garcia & Tamara B. Harris
Department of Public Health and Caring Sciences, Geriatrics, Uppsala University, Uppsala 75185, Sweden.,
Vilmantas Giedraitis
Division of Cardiovascular Epidemiology, Institute of Environmental Medicine, Karolinska Institutet, Stockholm, Sweden, Stockholm 17177, Sweden.,
Bruna Gigante, Karin Leander & Ulf de Faire
Division of Research, Kaiser Permanente, Oakland, 94612, California, USA
Alan S. Go
Service of Therapeutic Education for Diabetes, Obesity and Chronic Diseases, Geneva University Hospital, Geneva CH-1211, Switzerland.,
Alain Golay
Department of Cardiovascular Sciences, University of Leicester, Glenfield Hospital, Leicester LE3 9QP, UK.,
Alison H. Goodall & Nilesh J. Samani
National Institute for Health Research (NIHR) Leicester Cardiovascular Biomedical Research Unit, Glenfield Hospital, Leicester LE3 9QP, UK.,
Alison H. Goodall & Nilesh J. Samani
Department of Nephrology, University Hospital Regensburg, D-93053 Regensburg, Germany.,
Mathias Gorski
Department of Psychiatry and Psychotherapy, University Medicine Greifswald, HELIOS-Hospital Stralsund, D-17475 Greifswald, Germany.,
Hans-Jörgen Grabe
German Center for Neurodegenerative Diseases (DZNE), Rostock, Greifswald, D-17475 Greifswald, Germany.,
Hans-Jörgen Grabe
Research Unit of Molecular Epidemiology, Helmholtz Zentrum München - German Research Center for Environmental Health, D-85764 Neuherberg, Germany.,
Harald Grallert, Thomas Illig & Annette Peters
German Center for Diabetes Research (DZD), 85764 Neuherberg, Germany.,
Harald Grallert & Barbara Thorand
Department of Medicine III, University Hospital Carl Gustav Carus, Technische Universität Dresden, D-01307 Dresden, Germany.,
Jürgen Gräßler, Peter E. H. Schwarz & Stefan R. Bornstein
Institut inter Régional pour la Santé, Synergies, F-37520 La Riche, France.,
Gaëlle Gusto
Department of Public Health and Clinical Medicine, Unit of Nutritional Research, Umeå University, Umeå 90187, Sweden.,
Goran Hallmans
Department of Psychiatry, University of Groningen, University Medical Center Groningen, 9700RB Groningen, The Netherlands.,
Catharina A. Hartman & Albertine J. Oldehinkel
Kuopio Research Institute of Exercise Medicine, 70100 Kuopio, Finland.,
Maija Hassinen, Timo A. Lakka & Rainer Rauramaa
MRC Human Genetics Unit, Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh EH4 2XU, UK.,
Caroline Hayward & Alan F. Wright
National Heart, Lung, and Blood Institute, the Framingham Heart Study, Framingham, 01702, Massachusetts, USA
Nancy L. Heard-Costa, Joanne M. Murabito, L. Adrienne Cupples & Caroline S. Fox
Department of Neurology, Boston University School of Medicine, Boston, 02118, Massachusetts, USA
Nancy L. Heard-Costa
Faculty of Psychology and Education, VU University Amsterdam, 1081BT Amsterdam, The Netherlands.,
Quinta Helmer
Deutsches Forschungszentrum für Herz-Kreislauferkrankungen (DZHK) (German Research Centre for Cardiovascular Research), Munich Heart Alliance, D-80636 Munich, Germany.,
Christian Hengstenberg, Martina Müller-Nurasyid, Annette Peters & Heribert Schunkert
Deutsches Herzzentrum München, Technische Universität München, D-80636 Munich, Germany.,
Christian Hengstenberg & Heribert Schunkert
Department of Public Health and General Practice, Norwegian University of Science and Technology, Trondheim 7489, Norway.,
Oddgeir Holmen & Kristian Hveem
Biological Psychology, VU University Amsterdam, 1081BT Amsterdam, The Netherlands.,
Jouke-Jan Hottenga & Gonneke Willemsen
Department of Pulmonary Physiology and Sleep Medicine, Nedlands, 6009, Western Australia, Australia
Alan L. James
School of Medicine and Pharmacology, University of Western Australia, Crawley 6009, Australia.,
Alan L. James
Department of Immunology, Uppsala University, Genetics, Pathology, SciLifeLab, Rudbeck Laboratory, SE-751 85 Uppsala, Sweden.,
Åsa Johansson & Ulf Gyllensten
Hjelt Institute Department of Public Health, University of Helsinki, FI-00014 Helsinki, Finland.,
Markku Koskenvuo & Jaakko Kaprio
Department of Internal Medicine I, Ulm University Medical Centre, D-89081 Ulm, Germany.,
Wolfgang Kratzer & Thomas Seufferlein
Finnish Institute of Occupational Health, FI-90100 Oulu, Finland.,
Jaana Laitinen
Division of Genetic Epidemiology, Department of Medical Genetics, Molecular and Clinical Pharmacology, Innsbruck Medical University, 6020 Innsbruck, Austria.,
Claudia Lamina
Institute of Human Genetics, Helmholtz Zentrum München - German Research Center for Environmental Health, D-85764 Neuherberg, Germany.,
Peter Lichtner
Department of Medical Sciences, Cardiovascular Epidemiology, Uppsala University, Uppsala 75185, Sweden.,
Lars Lind & Johan Sundström
Montreal Heart Institute, Montreal, Quebec H1T 1C8, Canada.,
Ken Sin Lo & John D. Rioux
Institute for Community Medicine, University Medicine Greifswald, D-17475 Greifswald, Germany.,
Roberto Lorbeer
The Genetics of Obesity and Related Metabolic Traits Program, The Icahn School of Medicine at Mount Sinai, New York, 10029, New York, USA
Yingchang Lu & Ruth J. F. Loos
School of Social and Community Medicine, University of Bristol, Bristol BS8 2BN, UK.,
Wendy L. McArdle & Josine L. Min
Institute of Molecular and Cell Biology, University of Tartu, Tartu 51010, Estonia.,
Evelin Mihailov & Andres Metspalu
Farr Institute of Health Informatics Research, University College London, London NW1 2DA, UK.,
Alireza Moayyeri
The Center for Observational Research, Amgen, Inc., Thousand Oaks, 91320, California, USA
Keri L. Monda
Istituto di Ricerca Genetica e Biomedica (IRGB), Consiglio Nazionale delle Ricerche, Cagliari, Sardinia 09042, Italy.,
Antonella Mulas & Serena Sanna
Center for Evidence-based Healthcare, University Hospital Carl Gustav Carus, Technische Universität Dresden, D-01307 Dresden, Germany.,
Gabriele Müller
Department of Medicine I, University Hospital Grosshadern, Ludwig-Maximilians-Universität, D-81377 Munich, Germany.,
Martina Müller-Nurasyid
Institute of Medical Informatics, Biometry and Epidemiology, Chair of Genetic Epidemiology, Ludwig-Maximilians-Universität, D-81377 Munich, Germany.,
Martina Müller-Nurasyid & Konstantin Strauch
Department of Respiratory Medicine, Sir Charles Gairdner Hospital, Nedlands, 6009, Western Australia, Australia
Arthur W. Musk
Laboratory of Genetics, National Institute on Aging, Baltimore, 21224, Maryland, USA
Ramaiah Nagaraja & David Schlessinger
Department of Genomics, Life & Brain Center, University of Bonn, 53127 Bonn, Germany.,
Markus M. Nöthen
Institute of Human Genetics, University of Bonn, 53127 Bonn, Germany.,
Markus M. Nöthen
Department of Epidemiology, University Medical Center Groningen, University of Groningen, 9700 RB Groningen, The Netherlands.,
Ilja M. Nolte
Department of Epidemiology and Biostatistics, Institute for Research in Extramural Medicine, Institute for Health and Care Research, VU University Medical Center, 1081BT Amsterdam, The Netherlands.,
Stefan Pilz
Department of Internal Medicine, Division of Endocrinology and Metabolism, Medical University of Graz, 8036 Graz, Austria.,
Stefan Pilz
Institute of Physiology, University Medicine Greifswald, D-17495 Karlsburg, Germany.,
Rainer Rettig
Stanley Center for Psychiatric Research, Broad Institute of MIT and Harvard, Cambridge, 02142, Massachusetts, USA
Stephan Ripke & Steven A. McCarroll
Division of Preventive Medicine, Brigham and Women's Hospital, Boston, 02215, Massachusetts, USA
Lynda M. Rose, Daniel I. Chasman & Paul M. Ridker
Clinical Institute of Medical and Chemical Laboratory Diagnostics, Medical University of Graz, Graz 8036, Austria.,
Hubert Scharnagl & Winfried März
Department of Preventive Medicine, Keck School of Medicine, University of Southern California, Los Angeles, 90089, California, USA
Fredrick R. Schumacher & Christopher A. Haiman
National Cancer Institute, Bethesda, 20892, Maryland, USA
Jianxin Shi
Icelandic Heart Association, Kopavogur 201, Iceland.,
Albert Vernon Smith & Vilmundur Gudnason
University of Iceland, Reykjavik 101, Iceland.,
Albert Vernon Smith & Vilmundur Gudnason
Department of Epidemiology, University Medical Center Groningen, University of Groningen, 9700 RB Groningen, The Netherlands.,
Joanna Smolonska, Judith M. Vonk, Ronald P. Stolk & Harold Snieder
Molecular & Cellular Therapeutics, Royal College of Surgeons in Ireland, 123 St Stephen’s Green, Dublin 2, Ireland.,
Alice V. Stanton
deCODE Genetics, Amgen Inc., Reykjavik 101, Iceland.,
Valgerdur Steinthorsdottir, Gudmar Thorleifsson, Unnur Thorsteinsdottir & Kari Stefansson
Department of Medical Sciences, Molecular Medicine, Uppsala University, Uppsala 75144, Sweden.,
Ann-Christine Syvänen
National Heart and Lung Institute, Imperial College London, London SW3 6LY, UK.,
Sian-Tsung Tan, Miriam F. Moffatt & Jaspal S. Kooner
Department of Public Health Sciences, Stritch School of Medicine, Loyola University of Chicago, Maywood, 61053, Illinois, USA
Bamidele O. Tayo & Richard S. Cooper
Institute of Epidemiology II, Helmholtz Zentrum München - German Research Center for Environmental Health, Neuherberg, Germany, D-85764 Neuherberg, Germany.,
Barbara Thorand & Annette Peters
Department of Oncology, University of Cambridge, Cambridge CB2 0QQ, UK.,
Jonathan P. Tyrer
Department of Internal Medicine and Clinical Nutrition, Centre for Bone and Arthritis Research, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, Gothenburg 413 45, Sweden.,
Liesbeth Vandenput & Claes Ohlsson
Department of Child and Adolescent Psychiatry/Psychology, Erasmus MC University Medical Centre, 3000 CB Rotterdam, The Netherlands.,
Frank C. Verhulst
Department for Health Evidence, Radboud University Medical Centre, 6500 HB Nijmegen, The Netherlands.,
Sita H. Vermeulen & Lambertus A. Kiemeney
Department of Genetics, Radboud University Medical Centre, 6500 HB Nijmegen, The Netherlands.,
Sita H. Vermeulen
Department of Clinical Pharmacology, William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London EC1M 6BQ, UK.,
Helen R. Warren, Mark J. Caulfield & Patricia B. Munroe
Genetics, GlaxoSmithKline, King of Prussia, 19406, Pennsylvania, USA
Dawn Waterworth
German Center for Cardiovascular Research, partner site Hamburg/Lubeck/Kiel, 23562 Lubeck, Germany.,
Christina Willenborg & Jeanette Erdmann
Institut für Integrative und Experimentelle Genomik, Universität zu Lübeck, D-23562 Lübeck, Germany.,
Christina Willenborg & Jeanette Erdmann
Department of Community Medicine, Faculty of Health Sciences, UiT The Arctic University of Norway, 9037 Tromsø, Norway.,
Tom Wilsgaard & Inger Njølstad
MRC Unit for Lifelong Health and Ageing at University College London, London WC1B 5JU, UK.,
Andrew Wong, Ken K. Ong & Diana Kuh
Diabetes Complications Research Centre, Conway Institute, School of Medicine and Medical Sciences, University College Dublin, Dublin 4, Ireland.,
Eoin P. Brennan
Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea
Murim Choi
Departments of Human Genetics, Lady Davis Institute, Epidemiology and Biostatistics, McGill University, Montréal, Québec H3T1E2, Canada.,
Zari Dastani
Department of Molecular Medicine and Surgery, Cardiothoracic Surgery Unit, Karolinska Institutet, Stockholm 17176, Sweden.,
Anders Franco-Cereceda
Department of Medicine, Columbia University College of Physicians and Surgeons, 10032, New York, USA
Ali G. Gharavi & Krzysztof Kiryluk
Biosciences Research Division, Department of Primary Industries, Victoria 3083, Australia.,
Michael E. Goddard
Department of Food and Agricultural Systems, University of Melbourne, Victoria 3010, Australia.,
Michael E. Goddard
Department of Epidemiology, Harvard School of Public Health, Boston, 02115, Massachusetts, USA
Jinyan Huang, Liming Liang, Baoshan Ma & David J. Hunter
State Key Laboratory of Medical Genomics, Shanghai Institute of Hematology, Rui Jin Hospital Affiliated with Shanghai Jiao Tong University School of Medicine, Shanghai, China
Jinyan Huang
NIHR Oxford Biomedical Research Centre, OUH Trust, Oxford OX3 7LE, UK.,
Fredrik Karpe
Cardiovascular Research Center, Massachusetts General Hospital, Harvard Medical School, Boston, Massachusetts, USA
Sekar Kathiresan
Laboratory for Genotyping Development, RIKEN Center for Integrative Medical Sciences, Yokohama 230-0045, Japan.,
Michiaki Kubo
Center for Genome Science, National Institute of Health, Chungcheongbuk-do, Chungbuk 363–951, Republic of Korea.,
Jong-Young Lee
Department of Biostatistics, Harvard School of Public Health, Harvard University, Boston, 2115, Massachusetts, USA
Liming Liang
Department of Genetics, Howard Hughes Medical Institute, Yale University School of Medicine, New Haven, New Haven, 06520, Connecticut, USA
Richard P. Lifton
College of Information Science and Technology, Dalian Maritime University, Dalian, 116026, Liaoning, China
Baoshan Ma
Nephrology Research, Centre for Public Health, Queen’s University of Belfast, Belfast, County Down BT9 7AB, UK.,
Amy J. McKnight
Section of General Internal Medicine, Boston University School of Medicine, Boston, 02118, Massachusetts, USA
Joanne M. Murabito
Department of Statistics, University of Oxford, 1 South Parks Road, Oxford OX1 3TG, UK.,
George Nicholson
MRC Harwell, Harwell Science and Innovation Campus, Harwell OX11 0QG, UK.,
George Nicholson
Institute of Health and Biomedical Innovation, Queensland University of Technology, Brisbane, 4059, Queensland, Australia
Dale R. Nyholt
Laboratory for Statistical Analysis, RIKEN Center for Integrative Medical Sciences, Yokohama 230-0045, Japan.,
Yukinori Okada & Atsushi Takahashi
Department of Human Genetics and Disease Diversity, Graduate School of Medical and Dental Sciences, Tokyo Medical and Dental University, 113-8510 Tokyo, Japan.,
Yukinori Okada & Toshihiro Tanaka
Genome Institute of Singapore, Agency for Science, Technology and Research, 138672 Singapore.,
Rajkumar Dorajoo
Department of Biomedical Engineering and Computational Science, Aalto University School of Science, Helsinki FI-00076, Finland.,
Niina Sandholm
Department of Medicine, Division of Nephrology, Helsinki University Central Hospital, FI-00290 Helsinki, Finland.,
Niina Sandholm
Folkhälsan Institute of Genetics, Folkhälsan Research Center, FI-00290 Helsinki, Finland.,
Niina Sandholm
Laboratory for Cardiovascular Diseases, RIKEN Center for Integrative Medical Sciences, Yokohama 230-0045, Japan.,
Toshihiro Tanaka
Division of Disease Diversity, Bioresource Research Center, Tokyo Medical and Dental University, 113-8510 Tokyo, Japan.,
Toshihiro Tanaka
Division of Epidemiology, Department of Medicine; Vanderbilt Epidemiology Center,
Wei Zheng
Vanderbilt Epidemiology Center, Nashville, 37075, Tennessee, USA
Wei Zheng
Nuffield Department of Obstetrics & Gynaecology, University of Oxford, Oxford OX3 7BN, UK.,
Krina T. Zondervan
Department of Psychiatry, Washington University School of Medicine, St Louis, 63110, Missouri, USA
Andrew C. Heath, Pamela A. F. Madden, D. C. Rao & Treva K. Rice
Department of Epidemiology and Public Health, EA3430, University of Strasbourg, Faculty of Medicine, Strasbourg, France
Dominique Arveiler
Department of Internal Medicine, University Medical Center Groningen, University of Groningen, 9700RB Groningen, The Netherlands.,
Stephan J. L. Bakker & Ron T. Gansevoort
Pathology and Laboratory Medicine, The University of Western Australia, Perth, 6009, Western Australia, Australia
John Beilby & Jennie Hui
Cedars-Sinai Diabetes and Obesity Research Institute, Los Angeles, 90048, California, USA
Richard N. Bergman
Institute of Social and Preventive Medicine (IUMSP), Centre Hospitalier Universitaire Vaudois and University of Lausanne, 1010 Lausanne, Switzerland.,
Pascal Bovet & Murielle Bochud
Ministry of Health, Victoria, Republic of Seychelles
Pascal Bovet & Murielle Bochud
University of Milano, Bicocca, 20126, Italy.,
Giancarlo Cesana
Harvard Medical School, Boston, 02115, Massachusetts, USA
Daniel I. Chasman & Paul M. Ridker
Center for Human Genetics Research, Vanderbilt University Medical Center, Nashville, 37203, Tennessee, USA
Dana C. Crawford
Department of Molecular Physiology and Biophysics, Vanderbilt University, Nashville, 37232, Tennessee, USA
Dana C. Crawford
Department of Biostatistics, Boston University School of Public Health, Boston, 02118, Massachusetts, USA
L. Adrienne Cupples
Department of Health Sciences, University of Milano, I 20142, Italy.,
Daniele Cusi
Fondazione Filarete, Milano I 20139, Italy.,
Daniele Cusi
Department of Public Health and Primary Care, University of Cambridge, Cambridge CB1 8RN, UK.,
John Danesh & Danish Saleheen
Julius Center for Health Sciences and Primary Care, University Medical Center Utrecht, 3584 CX Utrecht, The Netherlands.,
Hester M. den Ruijter
Institute of Cardiovascular and Medical Sciences, College of Medical, Veterinary and Life Sciences, University of Glasgow, Glasgow G12 8TA, UK.,
Anna F. Dominiczak
Clinic of Cardiology, West-German Heart Centre, University Hospital Essen, 45147 Essen, Germany.,
Raimund Erbel
Department of General Practice and Primary Health Care, University of Helsinki, FI-00290 Helsinki, Finland.,
Johan G. Eriksson
Unit of General Practice, Helsinki University Central Hospital, Helsinki 00290, Finland.,
Johan G. Eriksson
Department of Internal Medicine B, University Medicine Greifswald, D-17475 Greifswald, Germany.,
Stephan B. Felix
DZHK (Deutsches Zentrum für Herz-Kreislaufforschung – German Centre for Cardiovascular Research), partner site Greifswald, D-17475 Greifswald, Germany.,
Stephan B. Felix, Uwe Völker & Henri Wallaschofski
Department of Internal Medicine, University of Pisa, 56100 Pisa, Italy.,
Ele Ferrannini
National Research Council Institute of Clinical Physiology, University of Pisa, 56124 Pisa, Italy.,
Ele Ferrannini
Department of Cardiology, Toulouse University School of Medicine, Rangueil Hospital, 31400 Toulouse, France.,
Jean Ferrières
Robertson Center for Biostatistics, University of Glasgow, Glasgow G12 8QQ, UK.,
Ian Ford
UWI Solutions for Developing Countries, The University of the West Indies, Mona, Kingston 7, Jamaica.,
Terrence Forrester & Colin A. McKenzie
NorthShore University HealthSystem, Evanston, IL 60201, University of Chicago, Chicago, Illinois, USA
Pablo V. Gejman
Leeds MRC Medical Bioinformatics Centre, University of Leeds, Leeds LS2 9LU, UK.,
Alistair S. Hall
Institute of Biomedical & Clinical Science, University of Exeter, Barrack Road, Exeter EX2 5DW, UK.,
Andrew T. Hattersley
Center for Biomedicine, European Academy Bozen, Bolzano (EURAC), Bolzano 39100, Italy (affiliated institute of the University of Lübeck, D-23562 Lübeck, Germany).,
Andrew A. Hicks & Peter P. Pramstaller
Division of Genomic Medicine, National Human Genome Research Institute, National Institutes of Health, Bethesda, 20892, Maryland, USA
Lucia A. Hindorff
Institute of Cardiovascular Science, University College London, London WC1E 6BT, UK.,
Aroon D. Hingorani & Folkert W. Asselbergs
Department of Vascular Medicine, Academic Medical Center, 1105 AZ Amsterdam, The Netherlands.,
G. Kees Hovingh & John J. P. Kastelein
Centre for Cardiovascular Genetics, Institute Cardiovascular Sciences, University College London, London WC1E 6JJ, UK.,
Steve E. Humphries
Cardiovascular Genetics Division, Department of Internal Medicine, University of Utah, Salt Lake City, 84108, Utah, USA
Steven C. Hunt
Sansom Institute for Health Research, University of South Australia, Adelaide, 5000, South Australia, Australia
Elina Hyppönen
School of Population Health, University of South Australia, Adelaide, 5000, South Australia, Australia
Elina Hyppönen
South Australian Health and Medical Research Institute, Adelaide, 5000, South Australia, Australia
Elina Hyppönen
Population, Policy, and Practice, University College London Institute of Child Health, London WC1N 1EH, UK.,
Elina Hyppönen & Chris Power
Hannover Unified Biobank, Hannover Medical School, Hannover, D-30625 Hannover, Germany.,
Thomas Illig
National Institute for Health and Welfare, FI-90101 Oulu, Finland.,
Marjo-Riitta Jarvelin
MRC Health Protection Agency (HPA) Centre for Environment and Health, School of Public Health, Imperial College London, London W2 1PG, UK.,
Marjo-Riitta Jarvelin
Unit of Primary Care, Oulu University Hospital, FI-90220 Oulu, Finland.,
Marjo-Riitta Jarvelin
Institute of Health Sciences, University of Oulu, FI-90014 Oulu, Finland.,
Marjo-Riitta Jarvelin, Marjo-Riitta Jarvelin & Sirkka M. Keinanen-Kiukaanniemi
Durrer Center for Cardiogenetic Research, Interuniversity Cardiology Institute Netherlands (ICIN), 3501 DG Utrecht, The Netherlands.,
J. Wouter Jukema, Folkert W. Asselbergs & Pim van der Harst
Interuniversity Cardiology Institute of the Netherlands (ICIN), 3501 DG Utrecht, The Netherlands.,
J. Wouter Jukema
Unit of Primary Health Care/General Practice, Oulu University Hospital, FI-90220 Oulu, Finland.,
Sirkka M. Keinanen-Kiukaanniemi
Department of Urology, Radboud University Medical Centre, 6500 HB Nijmegen, The Netherlands.,
Lambertus A. Kiemeney
Imperial College Healthcare NHS Trust, London W12 0HS, UK.,
Jaspal S. Kooner & John C. Chambers
Department of Epidemiology and Public Health, University College London, London WC1E 6BT, UK.,
Meena Kumari, Claudia Langenberg & Mika Kivimaki
Department of Biological and Social Epidemiology, University of Essex, Wivenhoe Park, Colchester, Essex CO4 3SQ, UK.,
Meena Kumari
Department of Medicine, Kuopio University Hospital and University of Eastern Finland, FI-70210 Kuopio, Finland.,
Johanna Kuusisto & Markku Laakso
Department of Physiology, Institute of Biomedicine, University of Eastern Finland, Kuopio Campus, FI-70211 Kuopio, Finland.,
Timo A. Lakka
Department of Clinical Physiology and Nuclear Medicine, Kuopio University Hospital and University of Eastern Finland, FI-70210 Kuopio, Finland.,
Timo A. Lakka & Rainer Rauramaa
Department of Clinical Chemistry, Fimlab Laboratories and School of Medicine University of Tampere, FI-33520 Tampere, Finland.,
Terho Lehtimäki
Steno Diabetes Center A/S, Gentofte DK-2820, Denmark.,
Valeriya Lyssenko
Lund University Diabetes Centre and Department of Clinical Science, Diabetes & Endocrinology Unit, Lund University, Malmö 221 00, Sweden.,
Valeriya Lyssenko & Leif C. Groop
Institut Universitaire de Cardiologie et de Pneumologie de Québec, Faculty of Medicine, Laval University, Quebec, QC G1V 0A6, Canada.,
André Marette
Institute of Nutrition and Functional Foods, Laval University, Quebec, QC G1V 0A6, Canada.,
André Marette, Marie-Claude Vohl & Louis Pérusse
Department of Biostatistics, University of Washington, Seattle, 98195, Washington, USA
Barbara McKnight
Department of Surgery, University Medical Center Utrecht, 3584 CX Utrecht, The Netherlands.,
Frans L. Moll
Department of Biostatistics, University of Liverpool, Liverpool L69 3GA, UK.,
Andrew P. Morris
Department of Pediatrics, University of Iowa, Iowa City, 52242, Iowa, USA
Jeffrey C. Murray
Illumina, Inc, Little Chesterford, Cambridge CB10 1XL, UK.,
John F. Peden
Department of Pulmonary Medicine and Tuberculosis, University of Groningen, University Medical Center Groningen, Groningen, The Netherlands
Dirkje S. Postma
Department of Neurology, General Central Hospital, Bolzano 39100, Italy.,
Peter P. Pramstaller
Department of Clinical Physiology and Nuclear Medicine, Turku University Hospital, FI-20521 Turku, Finland.,
Olli T. Raitakari
Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, FI-20521 Turku, Finland.,
Olli T. Raitakari
Human Genomics Laboratory, Pennington Biomedical Research Center, Baton Rouge, 70808, Louisiana, USA
Tuomo Rankinen, Mark A. Sarzynski & Claude Bouchard
Université de Montréal, Montreal, Quebec H1T 1C8, Canada.,
John D. Rioux
Center for Systems Genomics, The Pennsylvania State University, University Park, 16802, Pennsylvania, USA
Marylyn D. Ritchie
Croatian Centre for Global Health, Faculty of Medicine, University of Split, 21000 Split, Croatia.,
Igor Rudan
South Carelia Central Hospital, 53130 Lappeenranta, Finland.,
Jouko Saramies
Paul Langerhans Institute Dresden, German Center for Diabetes Research (DZD), 01307 Dresden, Germany.,
Peter E. H. Schwarz
International Centre for Circulatory Health, Imperial College London, London W2 1PG, UK.,
Peter Sever
Division of Endocrinology, Diabetes and Nutrition, University of Maryland School of Medicine, Baltimore, 21201, Maryland, USA
Alan R. Shuldiner & Jeffrey R. O’Connell
Program for Personalized and Genomic Medicine, University of Maryland School of Medicine, Baltimore, 21201, Maryland, USA
Alan R. Shuldiner & Jeffrey R. O’Connell
Geriatric Research and Education Clinical Center, Vetrans Administration Medical Center, Baltimore, 21201, Maryland, USA
Alan R. Shuldiner
Department of Medicine, Helsinki University Central Hospital Heart and Lung Center, Helsinki University Central Hospital, FI-00290 Helsinki, Finland.,
Juha Sinisalo
Sorbonne Universités, UPMC Univ Paris 06, UMR S 1166, F-75013 Paris, France.,
David-Alexandre Trégouët
INSERM, UMR S 1166, Team Genomics and Physiopathology of Cardiovascular Diseases, F-75013 Paris, France.,
David-Alexandre Trégouët
Institute for Cardiometabolism And Nutrition (ICAN), F-75013 Paris, France.,
David-Alexandre Trégouët
Department of Kinesiology, Laval University, Quebec QC G1V 0A6, Canada.,
Angelo Tremblay & Louis Pérusse
Dipartimento di Scienze Farmacologiche e Biomolecolari, Università di Milano & Centro Cardiologico Monzino, Instituto di Ricovero e Cura a Carattere Scientifico, Milan 20133, Italy.,
Elena Tremoli
Department of Food Science and Nutrition, Laval University, Quebec QC G1V 0A6, Canada.,
Marie-Claude Vohl
Department of Internal Medicine, University Hospital (CHUV) and University of Lausanne, Lausanne 1011, Switzerland.,
Gérard Waeber
Department of Nutrition, University of North Carolina, Chapel Hill, 27599, North Carolina, USA
Linda S. Adair
Institut Pasteur de Lille; INSERM, U744; Université de Lille 2; F-59000 Lille, France.,
Philippe Amouyel
INSERM, U744,
Philippe Amouyel
Department of Cardiology, Division Heart and Lungs, University Medical Center Utrecht, 3584 CX Utrecht, The Netherlands.,
Folkert W. Asselbergs
Department of Medicine, Stanford University School of Medicine, Palo Alto, 94304, California, USA
Themistocles L. Assimes, Mads Melbye & Thomas Quertermous
Lee Kong Chian School of Medicine, Imperial College London and Nanyang Technological University, Singapore, 637553 Singapore, Singapore.,
Bernhard O. Boehm
Department of Internal Medicine I, Ulm University Medical Centre, D-89081 Ulm, Germany.,
Bernhard O. Boehm
Health Science Center at Houston, University of Texas, Houston, 77030, Texas, USA
Eric Boerwinkle
Department of Medicine, Division of Genetics, Brigham and Women’s Hospital, Harvard Medical School, Boston, 02115, Massachusetts, USA
Paul I. W. de Bakker
Department of Epidemiology, University Medical Center Utrecht, 3584 CX Utrecht, The Netherlands.,
Paul I. W. de Bakker
School of Population Health, The University of Western Australia, Nedlands, 6009, Western Australia, Australia
Jennie Hui
Department of Epidemiology and Population Health, Albert Einstein College of Medicine, Belfer 1306, New York 10461, USA.,
Robert C. Kaplan
Division of Public Health Sciences, Center for Human Genetics, Wake Forest School of Medicine, Winston-Salem, 27157, North Carolina, USA
Yongmei Liu
Synlab Academy, Synlab Services GmbH, 68163 Mannheim, Germany.,
Winfried März
Department of Clinical Medicine, Copenhagen University, 2200 Copenhagen, Denmark.,
Mads Melbye
Department of Clinical Genetics, Erasmus MC University Medical Center, 3000 CA Rotterdam, The Netherlands.,
Ben A. Oostra
Finnish Diabetes Association, Kirjoniementie 15, FI-33680 Tampere, Finland.,
Timo E. Saaristo
Pirkanmaa Hospital District, FI-33521 Tampere, Finland.,
Timo E. Saaristo
Center for Non-Communicable Diseases, Karatchi, Pakistan
Danish Saleheen
Department of Medicine, University of Pennsylvania, Philadelphia, 19104, Pennsylvania, USA
Danish Saleheen
Division of Cardiovascular and Medical Sciences, BHF Glasgow Cardiovascular Research Centre, University of Glasgow, Glasgow G12 8TA, UK.,
Naveed Sattar
Icahn Institute for Genomics and Multiscale Biology, Icahn School of Medicine at Mount Sinai, New York, 10580, New York, USA
Eric E. Schadt
Faculty of Medicine, University of Iceland, Reykjavik 101, Iceland.,
Unnur Thorsteinsdottir & Kari Stefansson
Institute for Health Research, University Hospital of La Paz (IdiPaz), 28046 Madrid, Spain.,
Jaakko Tuomilehto
Diabetes Research Group, King Abdulaziz University, 21589 Jeddah, Saudi Arabia.,
Jaakko Tuomilehto
Centre for Vascular Prevention, Danube-University Krems, 3500 Krems, Austria.,
Jaakko Tuomilehto
Department of Public Health and Clinical Nutrition, University of Eastern Finland, Finland
Matti Uusitupa
Research Unit, Kuopio University Hospital, FI-70210 Kuopio, Finland.,
Matti Uusitupa
Institute of Cellular Medicine, Newcastle University, Newcastle NE1 7RU, UK.,
Mark Walker
Institute of Clinical Chemistry and Laboratory Medicine, University Medicine Greifswald, D-17475 Greifswald, Germany.,
Henri Wallaschofski
Institute of Medical Informatics, Biometry and Epidemiology, Chair of Epidemiology, Ludwig-Maximilians-Universität, D-85764 Munich, Germany.,
H-Erich Wichmann
Klinikum Grosshadern, D-81377 Munich, Germany.,
H-Erich Wichmann
Institute of Epidemiology I, Helmholtz Zentrum München - German Research Center for Environmental Health, Neuherberg, Germany, D-85764 Neuherberg, Germany.,
H-Erich Wichmann
Department of Pulmonology, University Medical Center Utrecht, 3584 CX Utrecht, The Netherlands.,
Pieter Zanen
Princess Al-Jawhara Al-Brahim Centre of Excellence in Research of Hereditary Disorders (PACER-HD), King Abdulaziz University, 21589 Jeddah, Saudi Arabia.,
Panos Deloukas
Division of Population Health Sciences & Education, St George’s, University of London, London SW17 0RE, UK.,
David P. Strachan
Oxford NIHR Biomedical Research Centre, Oxford University Hospitals NHS Trust, Oxford OX3 7LJ, UK.,
Mark I. McCarthy
Clinical Epidemiology, Integrated Research and Treatment Center, Center for Sepsis Control and Care (CSCC), Jena University Hospital, 07743 Jena, Germany.,
André Scherag
Department of Human Genetics, University of Michigan, Ann Arbor, 48109, Michigan, USA
Cristen J. Willer
Service of Medical Genetics, CHUV University Hospital, 1011 Lausanne, Switzerland.,
Jacques S. Beckmann
University of Cambridge Metabolic Research Laboratories, Institute of Metabolic Science, Addenbrooke’s Hospital, Cambridge CB2 OQQ, UK.,
Inês Barroso
NIHR Cambridge Biomedical Research Centre, Institute of Metabolic Science, Addenbrooke’s Hospital, Cambridge CB2 OQQ, UK.,
Inês Barroso
Carolina Center for Genome Sciences, University of North Carolina at Chapel Hill, Chapel Hill, 27599, North Carolina, USA
Kari E. North
The Mindich Child Health and Development Institute, Icahn School of Medicine at Mount Sinai, New York, 10029, New York, USA
Ruth J. F. Loos
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Consortia
The LifeLines Cohort Study
The ADIPOGen Consortium
The AGEN-BMI Working Group
The CARDIOGRAMplusC4D Consortium
The CKDGen Consortium
The GLGC
The ICBP
The MAGIC Investigators
The MuTHER Consortium
The MIGen Consortium
The PAGE Consortium
The ReproGen Consortium
The GENIE Consortium
The International Endogene Consortium
Contributions
A full list of author contributions can be found in the Supplementary Information.
Corresponding authors
Correspondence to Joel N. Hirschhorn, Ruth J. F. Loos or Elizabeth K. Speliotes.
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Competing interests
Competing interests statement: G.T., V.S., U.T. and K.S. are employed by deCODE Genetics/Amgen, Inc. I.B. and spouse own stock in GlaxoSmithKline and Incyte, Ltd. C.B. is a consultant for Weight Watchers, Pathway Genomics, NIKE and Gatorade PepsiCo.
Additional information
A list of authors and affiliations appears in the Supplementary Information.
A list of authors and affiliations appears in the Supplementary Information.
A list of authors and affiliations appears in the Supplementary Information.
A list of authors and affiliations appears in the Supplementary Information.
A list of authors and affiliations appears in the Supplementary Information.
A list of authors and affiliations appears in the Supplementary Information.
A list of authors and affiliations appears in the Supplementary Information.
A list of authors and affiliations appears in the Supplementary Information.
A list of authors and affiliations appears in the Supplementary Information.
A list of authors and affiliations appears in the Supplementary Information.
A list of authors and affiliations appears in the Supplementary Information.
A list of authors and affiliations appears in the Supplementary Information.
A list of authors and affiliations appears in the Supplementary Information.
A list of authors and affiliations appears in the Supplementary Information.
Extended data figures and tables
Extended Data Figure 1 Study design.
*The SNP counts reflect sample size filter of n ≥ 50,000. §Counts represent the primary European sex-combined analysis. Please see Extended Data Table 1 for counts for secondary analyses.
Extended Data Figure 2 Genetic characterization of BMI-associated variants.
a, Plot of the cumulative phenotypic variance explained by each locus ordered by decreasing effect size. b, The relationship between effect size and allele frequency. Previously identified loci are blue circles and novel loci are red triangles. c, Quantile–quantile (Q–Q) plot of meta-analysis P values for all 1,909 BMI-replication SNPs (blue) and after removing SNPs near the 97 associated loci (green). d, Histogram of cumulative effect of BMI risk alleles. Mean BMI for each bin is shown by the black dots (with standard deviation) and corresponds to the right-hand y axis.
Extended Data Figure 3 Partitioning the variance in and risk prediction from SNP-derived predictor.
a, b, The analyses were performed using 2,758 full sibling pairs from the TwinGene cohort (a) and 1,622 pairs from the QIMR cohort (b). The SNP-based predictor was adjusted for the first 20 principal components. The variance of the SNP-based predictor can be partitioned into four components (Vg, Ve, Cg and Ce) using the within-family prediction analysis, in which Vg is the variance explained by real SNP effects, Cg is the covariance between predictors attributed to the real effects of SNPs that are not in LD but correlated due to population stratification, Ve is the accumulated variance due to the errors in estimating SNP effects, and Ce is the covariance between predictors attributed to errors in estimating the effects of SNPs that are correlated due to population stratification. Error bars reflect s.e.m. of estimates. c, The prediction R2 shown on the y axis is the squared correlation between phenotype and SNP-based genetic predictor in unrelated individuals from the TwinGene (n = 5,668) and QIMR (n = 3,953) studies. The number shown in each column is the number of SNPs selected from the GCTA joint and conditional analysis at a range of P-value thresholds. In each case, the predictor was adjusted by the first 20 principal components. The column in orange is the average prediction R2 weighted by sample size over the two cohorts. The dashed grey line is the value inferred from the within-family prediction analyses using this equation R2 = (Vg + Cg)2/(Vg + Ve + Cg + Ce).
Extended Data Figure 4 Comparison of BMI-associated index SNPs across ethnicities.
a, b, BMI effects observed in European ancestry individuals (x axes) compared to African ancestry (a) or Asian ancestry (b) individuals (y axes). c, d, Allele frequencies between ancestry groups, as in a and b. e, f, Comparison of the estimates of explained variance. In all plots, novel loci are in red and previously identified loci are in blue.
Extended Data Figure 5 Effects of BMI-associated loci on related metabolic traits.
Unsupervised hierarchical clustering of the 97 BMI-associated loci (y axis) on 23 related metabolic traits (x axis). The top row shows the a priori expected relationship with BMI (green is concordant effect direction, purple is opposite). Loci with statistically significant concordant direction of effect are highlighted in green, and significant but opposing effects are in purple. Grey indicates a non-significant relationship and those with no information are in white. The key in the top left corner also shows the count of gene–phenotype pairs in each category (cyan bars).
Extended Data Figure 6 Bubble chart representing the genetic overlap across traits at BMI susceptibility loci.
Each bubble represents a trait for which association results were requested for the 97 GWS BMI loci. The size of the bubble is proportional to the number of BMI-increasing loci with a significant association. A line connects each pair of bubbles with thickness proportional to the number of significant loci shared between the traits. Traits tested include the current study BMI SNPs, African-American BMI (AA BMI), hip circumference (HIP), HIP adjusted for BMI (HIPadjBMI), waist circumference (WC), waist circumference adjusted for BMI (WCadjBMI), waist-to-hip ratio (WHR), waist-to-hip ratio adjusted for BMI (WHRadjBMI), height, adiponectin, coronary artery disease (CAD), diastolic blood pressure (DBP), systolic blood pressure (SBP), high-density lipoprotein (HDL), low-density lipoprotein (LDL), total cholesterol (TC), triglycerides (TG), type 2 diabetes (T2D), fasting glucose (FG), fasting insulin (FI), fasting insulin adjusted for BMI (FIadjBMI), two-hour glucose (Glu2hr), diabetic nephropathy (Diab_Neph), age at menopause (AgeMenopause), and age at menarche (AgeMenarche).
Supplementary information
Supplementary Information
This file contains Supplementary Figures 1-10, Author Contributions and detailed Acknowledgements. (PDF 9393 kb)
Supplementary Data
This file contains Supplementary Tables 1-25. (XLSX 1588 kb)
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Locke, A., Kahali, B., Berndt, S. et al. Genetic studies of body mass index yield new insights for obesity biology. Nature 518, 197–206 (2015). https://doi.org/10.1038/nature14177
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DOI: https://doi.org/10.1038/nature14177
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