Article | Published:

Genetic studies of body mass index yield new insights for obesity biology

Nature volume 518, pages 197206 (12 February 2015) | Download Citation

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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Acknowledgements

A full list of acknowledgements can be found in the Supplementary Information.

Author information

Author notes

    • Jong-Young Lee

    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
    •  & Tune H. Pers

    These authors contributed equally to this work.

    • Kari E. North
    • , Erik Ingelsson
    • , Joel N. Hirschhorn
    • , Ruth J. F. Loos
    •  & Elizabeth K. Speliotes

    These authors jointly supervised this work.

Affiliations

  1. Center for Statistical Genetics, Department of Biostatistics, University of Michigan, Ann Arbor, Michigan 48109, USA.

    • Adam E. Locke
    • , Jennifer L. Bragg-Gresham
    • , Anne U. Jackson
    • , Heather M. Stringham
    • , Gonçalo R. Abecasis
    •  & Michael Boehnke
  2. Department of Internal Medicine, Division of Gastroenterology, and Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, Michigan 48109, USA.

    • Bratati Kahali
    • , Corey Powell
    •  & Elizabeth K. Speliotes
  3. Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, Maryland 20892, USA.

    • Sonja I. Berndt
    • , Zhaoming Wang
    • , Kevin B. Jacobs
    •  & Stephen J. Chanock
  4. 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
  5. Divisions of Endocrinology and Genetics and Center for Basic and Translational Obesity Research, Boston Children’s Hospital, Boston, Massachusetts 02115, USA.

    • Tune H. Pers
    • , Sailaja Vedantam
    • , Tonu Esko
    • , Cameron D. Palmer
    • , Rany M. Salem
    •  & Joel N. Hirschhorn
  6. Broad Institute of the Massachusetts Institute of Technology and Harvard University, Cambridge, Massachusetts 02142, 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
  7. Department of Genetics, Harvard Medical School, Boston, Massachusetts 02115, USA.

    • Tune H. Pers
    • , Tonu Esko
    • , Robert E. Handsaker
    • , Steven A. McCarroll
    • , Rany M. Salem
    •  & Joel N. Hirschhorn
  8. Center for Biological Sequence Analysis, Department of Systems Biology, Technical University of Denmark, Lyngby 2800, Denmark.

    • Tune H. Pers
  9. 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
  10. Department of Genetics, University of North Carolina, Chapel Hill, North Carolina 27599, USA.

    • Martin L. Buchkovich
    • , Damien C. Croteau-Chonka
    •  & Karen L. Mohlke
  11. Queensland Brain Institute, The University of Queensland, Brisbane 4072, Australia.

    • Jian Yang
    • , Anna A. E. Vinkhuyzen
    •  & Peter M. Visscher
  12. The University of Queensland Diamantina Institute, The Translation Research Institute, Brisbane 4012, Australia.

    • Jian Yang
    •  & Peter M. Visscher
  13. Channing Division of Network Medicine, Department of Medicine, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts 02115, USA.

    • Damien C. Croteau-Chonka
    • , Lu Qi
    •  & David J. Hunter
  14. 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
  15. Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm 17177, Sweden.

    • Tove Fall
    • , Henrik Grönberg
    • , Per Hall
    • , Patrik K. E. Magnusson
    •  & Nancy L. Pedersen
  16. Science for Life Laboratory, Uppsala University, Uppsala 75185, Sweden.

    • Tove Fall
    • , Stefan Gustafsson
    • , Åsa K. Hedman
    • , Johan Ärnlöv
    • , Ann-Christine Syvänen
    •  & Erik Ingelsson
  17. Department of Medical Sciences, Molecular Epidemiology, Uppsala University, Uppsala 75185, Sweden.

    • Tove Fall
    • , Stefan Gustafsson
    • , Åsa K. Hedman
    • , Johan Ärnlöv
    •  & Erik Ingelsson
  18. 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
  19. Institute of Social and Preventive Medicine (IUMSP), Centre Hospitalier Universitaire Vaudois (CHUV), Lausanne 1010, Switzerland.

    • Zoltán Kutalik
  20. Swiss Institute of Bioinformatics, Lausanne 1015, Switzerland.

    • Zoltán Kutalik
    •  & Jacques S. Beckmann
  21. Department of Medical Genetics, University of Lausanne, Lausanne 1005, Switzerland.

    • Zoltán Kutalik
    •  & Jacques S. Beckmann
  22. Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1SA, UK.

    • Joshua C. Randall
    • , Nigel W. Rayner
    • , Kathleen Stirrups
    • , Panos Deloukas
    •  & Inês Barroso
  23. 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
  24. 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
  25. Department of Nutrition, Harvard School of Public Health, Boston, Massachusetts 02115, USA.

    • Tsegaselassie Workalemahu
    • , Lu Qi
    • , Paul W. Franks
    •  & David J. Hunter
  26. Survey Research Center, Institute for Social Research, University of Michigan, Ann Arbor, Michigan 48104, USA.

    • Jessica D. Faul
    •  & David R. Weir
  27. Department of Epidemiology, University of Michigan, Ann Arbor, Michigan 48109, USA.

    • Jennifer A. Smith
    •  & Wei Zhao
  28. Department of Internal Medicine, Division of Cardiovascular Medicine, University of Michigan, Ann Arbor, Michigan 48109, USA.

    • Jin Chen
    •  & Cristen J. Willer
  29. 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
  30. Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, Michigan 48109, USA.

    • Ellen M. Schmidt
    •  & Cristen J. Willer
  31. HudsonAlpha Institute for Biotechnology, Huntsville, Alabama 35806, USA.

    • Devin Absher
    •  & Lindsay L. Waite
  32. Genetic Epidemiology Unit, Department of Epidemiology, Erasmus MC University Medical Center, 3015 GE Rotterdam, The Netherlands.

    • Najaf Amin
    • , Ayse Demirkan
    • , Aaron Isaacs
    • , Ben A. Oostra
    •  & Cornelia M. van Duijn
  33. Telethon Institute for Child Health Research, Centre for Child Health Research, The University of Western Australia, Perth, Western Australia 6008, Australia.

    • Denise Anderson
  34. 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
  35. Department of Molecular Epidemiology, Leiden University Medical Center, 2300 RC Leiden, The Netherlands.

    • Marian Beekman
    •  & P. Eline Slagboom
  36. 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
  37. Kidney Epidemiology and Cost Center, University of Michigan, Ann Arbor, Michigan 48109, USA.

    • Jennifer L. Bragg-Gresham
  38. Department of Statistics & Biostatistics, Rutgers University, Piscataway, New Jersey 08854, USA.

    • Steven Buyske
  39. Department of Genetics, Rutgers University, Piscataway, New Jersey 08854, USA.

    • Steven Buyske
    •  & Tara C. Matise
  40. Department of Human Genetics, Leiden University Medical Center, 2333 ZC Leiden, The Netherlands.

    • Ayse Demirkan
  41. Ealing Hospital NHS Trust, Middlesex UB1 3HW, UK.

    • Guohong Deng
    • , Weihua Zhang
    • , William R. Scott
    • , Sian-Tsung Tan
    • , Jaspal S. Kooner
    •  & John C. Chambers
  42. Department of Gastroenterology and Hepatology, Imperial College London, London W2 1PG, UK.

    • Guohong Deng
  43. Institute of infectious Diseases, Southwest Hospital, Third Military Medical University, Chongqing, China.

    • Guohong Deng
  44. Center for Complex Disease Genomics, McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University School of Medicine, Baltimore, Maryland 21205, USA.

    • Georg B. Ehret
    •  & Aravinda Chakravarti
  45. Cardiology, Department of Specialties of Internal Medicine, Geneva University Hospital, Geneva 1211, Switzerland.

    • Georg B. Ehret
    •  & François Mach
  46. Department of Epidemiology Research, Statens Serum Institut, Copenhagen DK-2300, Denmark.

    • Bjarke Feenstra
    • , Heather A. Boyd
    •  & Frank Geller
  47. Department of Genetics, Washington University School of Medicine, St Louis, Missouri 63110, USA.

    • Mary F. Feitosa
    • , E. Warwick Daw
    • , Mary K. Wojczynski
    • , Qunyuan Zhang
    • , Aldi T. Kraja
    • , D. C. Rao
    •  & Ingrid B. Borecki
  48. Division of Cardiovacular Medicine, Radcliffe Department of Medicine, University of Oxford, Oxford OX3 9DU, UK.

    • Anuj Goel
    • , Martin Farrall
    •  & Hugh Watkins
  49. Division of Public Health Sciences, Fred Hutchinson Cancer Research Center, Seattle, Washington 98109, USA.

    • Jian Gong
    • , Jeffrey Haessler
    • , Charles Kooperberg
    •  & Ulrike Peters
  50. 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
  51. 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
  52. Department of Internal Medicine II, Ulm University Medical Centre, D-89081 Ulm, Germany.

    • Marcus E. Kleber
    •  & Wolfgang Koenig
  53. 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
  54. Epidemiology Program, University of Hawaii Cancer Center, Honolulu, Hawaii 96813, USA.

    • Unhee Lim
    • , Lynne R. Wilkens
    •  & Loic Le Marchand
  55. The Charles Bronfman Institute for Personalized Medicine, Icahn School of Medicine at Mount Sinai, New York, New York 10029, USA.

    • Vaneet Lotay
    • , Janina M. Jeff
    • , Yingchang Lu
    • , Omri Gottesman
    • , Erwin P. Bottinger
    •  & Ruth J. F. Loos
  56. 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
  57. Department of Cardiology, University Medical Center Groningen, University of Groningen, 9700RB Groningen, The Netherlands.

    • Irene Mateo Leach
    • , Niek Verweij
    •  & Pim van der Harst
  58. 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
  59. 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
  60. 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
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  61. QIMR Berghofer Medical Research Institute, Brisbane, Queensland 4006, Australia.

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    • , Scott D. Gordon
    • , Grant W. Montgomery
    • , Dale R. Nyholt
    •  & Nicholas G. Martin
  62. Laboratory of Neurogenetics, National Institute on Aging, National Institutes of Health, Bethesda, Maryland 20892, USA.

    • Michael A. Nalls
  63. Institute for Medical Informatics, Biometry and Epidemiology (IMIBE), University Hospital Essen, 45147 Essen, Germany.

    • Sonali Pechlivanis
    • , Karl-Heinz Jöckel
    • , Susanne Moebus
    •  & André Scherag
  64. 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
  65. Department of Genomics of Common Disease, School of Public Health, Imperial College London, Hammersmith Hospital, London W12 0NN, UK.

    • Inga Prokopenko
    •  & Philippe Froguel
  66. 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
  67. Department of Public Health and Clinical Medicine, Unit of Medicine, Umeå University, Umeå 901 87, Sweden.

    • Dmitry Shungin
    •  & Paul W. Franks
  68. Department of Odontology, Umeå University, Umeå 901 85, Sweden.

    • Dmitry Shungin
  69. University of Eastern Finland, FI-70210 Kuopio, Finland.

    • Alena Stančáková
  70. Atherosclerosis Research Unit, Center for Molecular Medicine, Department of Medicine, Karolinska Institutet, Stockholm 17176, Sweden.

    • Rona J. Strawbridge
    • , Lasse Folkersen
    • , Per Eriksson
    • , Jesper R. Gådin
    • , Ferdinand M. van’t Hooft
    •  & Anders Hamsten
  71. Division of Biostatistics, Washington University School of Medicine, St Louis, Missouri 63110, USA.

    • Yun Ju Sung
    • , D. C. Rao
    •  & Treva K. Rice
  72. Translational Gerontology Branch, National Institute on Aging, Baltimore, Maryland 21225, USA.

    • Toshiko Tanaka
    •  & Luigi Ferrucci
  73. Interfaculty Institute for Genetics and Functional Genomics, University Medicine Greifswald, D-17475 Greifswald, Germany.

    • Alexander Teumer
    • , Georg Homuth
    •  & Uwe Völker
  74. Department of Cardiology, Leiden University Medical Center, 2300 RC Leiden, The Netherlands.

    • Stella Trompet
    •  & J. Wouter Jukema
  75. Department of Gerontology and Geriatrics, Leiden University Medical Center, 2300 RC Leiden, The Netherlands.

    • Stella Trompet
    •  & Anton J. M. de Craen
  76. Experimental Cardiology Laboratory, Division Heart and Lungs, University Medical Center Utrecht, 3584 CX Utrecht, The Netherlands.

    • Sander W. van der Laan
    • , Hester M. den Ruijter
    •  & Gerard Pasterkamp
  77. Department of Medical Genetics, University Medical Center Utrecht, 3584 CX Utrecht, The Netherlands.

    • Jessica van Setten
    •  & Paul I. W. de Bakker
  78. Department of Endocrinology, University of Groningen, University Medical Center Groningen, 9700 RB Groningen, The Netherlands.

    • Jana V. Van Vliet-Ostaptchouk
  79. Core Genotyping Facility, SAIC-Frederick, Inc., NCI-Frederick, Frederick, Maryland 21702, USA.

    • Zhaoming Wang
    •  & Kevin B. Jacobs
  80. CNRS UMR 8199, F-59019 Lille, France.

    • Loïc Yengo
    • , Elodie Eury
    • , Stéphane Lobbens
    • , Stéphane Cauchi
    •  & Philippe Froguel
  81. European Genomic Institute for Diabetes, F-59000 Lille, France.

    • Loïc Yengo
    • , Elodie Eury
    • , Stéphane Lobbens
    • , Stéphane Cauchi
    •  & Philippe Froguel
  82. Université de Lille 2, F-59000 Lille, France.

    • Loïc Yengo
    • , Elodie Eury
    • , Stéphane Lobbens
    • , Stéphane Cauchi
    •  & Philippe Froguel
  83. Department of Epidemiology and Biostatistics, Imperial College London, London W2 1PG, UK.

    • Weihua Zhang
    • , William R. Scott
    • , Marjo-Riitta Jarvelin
    •  & John C. Chambers
  84. Center for Medical Sytems Biology, 2300 RC Leiden, The Netherlands.

    • Aaron Isaacs
    • , Ben A. Oostra
    •  & Cornelia M. van Duijn
  85. 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
  86. School of Health and Social Studies, Dalarna University, SE-791 88 Falun, Sweden.

    • Johan Ärnlöv
  87. PathWest Laboratory Medicine of Western Australia, Nedlands, Western Australia 6009, Australia.

    • Gillian M. Arscott
    • , John Beilby
    •  & Jennie Hui
  88. Department of Haematology, University of Cambridge, Cambridge CB2 0PT, UK.

    • Antony P. Attwood
    •  & Jennifer Jolley
  89. NHS Blood and Transplant, Cambridge CB2 0PT, UK.

    • Antony P. Attwood
    •  & Jennifer Jolley
  90. Geriatric Unit, Azienda Sanitaria Firenze (ASF), 50125 Florence, Italy.

    • Stefania Bandinelli
  91. USC-Office of Population Studies Foundation, Inc., University of San Carlos, Cebu City 6000, Philippines.

    • Isabelita N. Bas
    •  & Nanette R. Lee
  92. Department of Genetics, Texas Biomedical Research Institute, San Antonio, Texas 78227, USA.

    • Claire Bellis
    •  & John Blangero
  93. Genomics Research Centre, Institute of Health and Biomedical Innovation, Queensland University of Technology, Brisbane, Queensland 4001, Australia.

    • Claire Bellis
  94. Department of Medical Sciences, Endocrinology, Diabetes and Metabolism, Uppsala University, Uppsala 75185, Sweden.

    • Christian Berne
  95. Division of Endocrinology, Diabetes and Metabolism, Ulm University Medical Centre, D-89081 Ulm, Germany.

    • Roza Blagieva
    •  & Sigrun Merger
  96. 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
  97. Department of Medicine, University of Leipzig, D-04103 Leipzig, Germany.

    • Matthias Blüher
    • , Peter Kovacs
    • , Anke Tönjes
    •  & Michael Stumvoll
  98. Department of Medical Statistics and Bioinformatics, Leiden University Medical Center, 2300 RC Leiden, The Netherlands.

    • Stefan Böhringer
    • , Quinta Helmer
    •  & Hae-Won Uh
  99. 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
  100. LifeLines Cohort Study, University Medical Center Groningen, University of Groningen, 9700 RB Groningen, The Netherlands.

    • Marcel Bruinenberg
    •  & Salome Scholtens
  101. Department of Biology, Norwegian University of Science and Technology, 7491 Trondheim, Norway.

    • Ida H. Caspersen
    •  & Berit Johansen
  102. Department of Pediatrics, University of California Los Angeles, Torrance, California 90502, USA.

    • Yii-Der Ida Chen
  103. Transgenomics Institute, Los Angeles Biomedical Research Institute, Torrance, California 90502, USA.

    • Yii-Der Ida Chen
  104. Clinical Trial Service Unit and Epidemiological Studies Unit, Nuffield Department of Population Health, University of Oxford, Oxford OX3 7LF, UK.

    • Robert Clarke
  105. Department of Dietetics-Nutrition, Harokopio University, 17671 Athens, Greece.

    • Maria Dimitriou
    •  & George Dedoussis
  106. 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
  107. Institute for Molecular Medicine, University of Helsinki, FI-00014 Helsinki, Finland.

    • Niina Eklund
    • , Jaakko Kaprio
    • , Leif C. Groop
    •  & Markus Perola
  108. Analytic and Translational Genetics Unit, Massachusetts General Hospital and Harvard Medical School, Boston, Massachusetts 02114, USA.

    • Karol Estrada
    •  & Stephan Ripke
  109. Laboratory of Epidemiology and Population Sciences, National Institute on Aging, NIH, Bethesda, Maryland 20892, USA.

    • Melissa E. Garcia
    •  & Tamara B. Harris
  110. Department of Public Health and Caring Sciences, Geriatrics, Uppsala University, Uppsala 75185, Sweden.

    • Vilmantas Giedraitis
  111. Division of Cardiovascular Epidemiology, Institute of Environmental Medicine, Karolinska Institutet, Stockholm, Sweden, Stockholm 17177, Sweden.

    • Bruna Gigante
    • , Karin Leander
    •  & Ulf de Faire
  112. Kaiser Permanente, Division of Research, Oakland, California 94612, USA.

    • Alan S. Go
  113. Service of Therapeutic Education for Diabetes, Obesity and Chronic Diseases, Geneva University Hospital, Geneva CH-1211, Switzerland.

    • Alain Golay
  114. Department of Cardiovascular Sciences, University of Leicester, Glenfield Hospital, Leicester LE3 9QP, UK.

    • Alison H. Goodall
    •  & Nilesh J. Samani
  115. National Institute for Health Research (NIHR) Leicester Cardiovascular Biomedical Research Unit, Glenfield Hospital, Leicester LE3 9QP, UK.

    • Alison H. Goodall
    •  & Nilesh J. Samani
  116. Department of Nephrology, University Hospital Regensburg, D-93053 Regensburg, Germany.

    • Mathias Gorski
  117. Department of Psychiatry and Psychotherapy, University Medicine Greifswald, HELIOS-Hospital Stralsund, D-17475 Greifswald, Germany.

    • Hans-Jörgen Grabe
  118. German Center for Neurodegenerative Diseases (DZNE), Rostock, Greifswald, D-17475 Greifswald, Germany.

    • Hans-Jörgen Grabe
  119. Research Unit of Molecular Epidemiology, Helmholtz Zentrum München - German Research Center for Environmental Health, D-85764 Neuherberg, Germany.

    • Harald Grallert
    • , Thomas Illig
    •  & Annette Peters
  120. German Center for Diabetes Research (DZD), 85764 Neuherberg, Germany.

    • Harald Grallert
    •  & Barbara Thorand
  121. 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
  122. Institut inter Régional pour la Santé, Synergies, F-37520 La Riche, France.

    • Gaëlle Gusto
  123. Department of Public Health and Clinical Medicine, Unit of Nutritional Research, Umeå University, Umeå 90187, Sweden.

    • Goran Hallmans
  124. Department of Psychiatry, University of Groningen, University Medical Center Groningen, 9700RB Groningen, The Netherlands.

    • Catharina A. Hartman
    •  & Albertine J. Oldehinkel
  125. Kuopio Research Institute of Exercise Medicine, 70100 Kuopio, Finland.

    • Maija Hassinen
    • , Timo A. Lakka
    •  & Rainer Rauramaa
  126. 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
  127. National Heart, Lung, and Blood Institute, the Framingham Heart Study, Framingham, Massachusetts 01702, USA.

    • Nancy L. Heard-Costa
    • , Joanne M. Murabito
    • , L. Adrienne Cupples
    •  & Caroline S. Fox
  128. Department of Neurology, Boston University School of Medicine, Boston, Massachusetts 02118, USA.

    • Nancy L. Heard-Costa
  129. Faculty of Psychology and Education, VU University Amsterdam, 1081BT Amsterdam, The Netherlands.

    • Quinta Helmer
  130. 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
  131. Deutsches Herzzentrum München, Technische Universität München, D-80636 Munich, Germany.

    • Christian Hengstenberg
    •  & Heribert Schunkert
  132. Department of Public Health and General Practice, Norwegian University of Science and Technology, Trondheim 7489, Norway.

    • Oddgeir Holmen
    •  & Kristian Hveem
  133. Biological Psychology, VU University Amsterdam, 1081BT Amsterdam, The Netherlands.

    • Jouke-Jan Hottenga
    •  & Gonneke Willemsen
  134. Department of Pulmonary Physiology and Sleep Medicine, Nedlands, Western Australia 6009, Australia.

    • Alan L. James
  135. School of Medicine and Pharmacology, University of Western Australia, Crawley 6009, Australia.

    • Alan L. James
  136. Uppsala University, Department of Immunology, Genetics, Pathology, SciLifeLab, Rudbeck Laboratory, SE-751 85 Uppsala, Sweden.

    • Åsa Johansson
    •  & Ulf Gyllensten
  137. Hjelt Institute Department of Public Health, University of Helsinki, FI-00014 Helsinki, Finland.

    • Markku Koskenvuo
    •  & Jaakko Kaprio
  138. Department of Internal Medicine I, Ulm University Medical Centre, D-89081 Ulm, Germany.

    • Wolfgang Kratzer
    •  & Thomas Seufferlein
  139. Finnish Institute of Occupational Health, FI-90100 Oulu, Finland.

    • Jaana Laitinen
  140. Division of Genetic Epidemiology, Department of Medical Genetics, Molecular and Clinical Pharmacology, Innsbruck Medical University, 6020 Innsbruck, Austria.

    • Claudia Lamina
  141. Institute of Human Genetics, Helmholtz Zentrum München - German Research Center for Environmental Health, D-85764 Neuherberg, Germany.

    • Peter Lichtner
  142. Department of Medical Sciences, Cardiovascular Epidemiology, Uppsala University, Uppsala 75185, Sweden.

    • Lars Lind
    •  & Johan Sundström
  143. Montreal Heart Institute, Montreal, Quebec H1T 1C8, Canada.

    • Ken Sin Lo
    •  & John D. Rioux
  144. Institute for Community Medicine, University Medicine Greifswald, D-17475 Greifswald, Germany.

    • Roberto Lorbeer
  145. The Genetics of Obesity and Related Metabolic Traits Program, The Icahn School of Medicine at Mount Sinai, New York, New York 10029, USA.

    • Yingchang Lu
    •  & Ruth J. F. Loos
  146. School of Social and Community Medicine, University of Bristol, Bristol BS8 2BN, UK.

    • Wendy L. McArdle
    •  & Josine L. Min
  147. Institute of Molecular and Cell Biology, University of Tartu, Tartu 51010, Estonia.

    • Evelin Mihailov
    •  & Andres Metspalu
  148. Farr Institute of Health Informatics Research, University College London, London NW1 2DA, UK.

    • Alireza Moayyeri
  149. The Center for Observational Research, Amgen, Inc., Thousand Oaks, California 91320, USA.

    • Keri L. Monda
  150. Istituto di Ricerca Genetica e Biomedica (IRGB), Consiglio Nazionale delle Ricerche, Cagliari, Sardinia 09042, Italy.

    • Antonella Mulas
    •  & Serena Sanna
  151. Center for Evidence-based Healthcare, University Hospital Carl Gustav Carus, Technische Universität Dresden, D-01307 Dresden, Germany.

    • Gabriele Müller
  152. Department of Medicine I, University Hospital Grosshadern, Ludwig-Maximilians-Universität, D-81377 Munich, Germany.

    • Martina Müller-Nurasyid
  153. Institute of Medical Informatics, Biometry and Epidemiology, Chair of Genetic Epidemiology, Ludwig-Maximilians-Universität, D-81377 Munich, Germany.

    • Martina Müller-Nurasyid
    •  & Konstantin Strauch
  154. Department of Respiratory Medicine, Sir Charles Gairdner Hospital, Nedlands, Western Australia 6009, Australia.

    • Arthur W. Musk
  155. Laboratory of Genetics, National Institute on Aging, Baltimore, Maryland 21224, USA.

    • Ramaiah Nagaraja
    •  & David Schlessinger
  156. Department of Genomics, Life & Brain Center, University of Bonn, 53127 Bonn, Germany.

    • Markus M. Nöthen
  157. Institute of Human Genetics, University of Bonn, 53127 Bonn, Germany.

    • Markus M. Nöthen
  158. Department of Epidemiology, University Medical Center Groningen, University of Groningen, 9700 RB Groningen, The Netherlands.

    • Ilja M. Nolte
  159. 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
  160. Department of Internal Medicine, Division of Endocrinology and Metabolism, Medical University of Graz, 8036 Graz, Austria.

    • Stefan Pilz
  161. Institute of Physiology, University Medicine Greifswald, D-17495 Karlsburg, Germany.

    • Rainer Rettig
  162. Stanley Center for Psychiatric Research, Broad Institute of MIT and Harvard, Cambridge, Massachusetts 02142, USA.

    • Stephan Ripke
    •  & Steven A. McCarroll
  163. Division of Preventive Medicine, Brigham and Women's Hospital, Boston, Massachusetts 02215, USA.

    • Lynda M. Rose
    • , Daniel I. Chasman
    •  & Paul M. Ridker
  164. Clinical Institute of Medical and Chemical Laboratory Diagnostics, Medical University of Graz, Graz 8036, Austria.

    • Hubert Scharnagl
    •  & Winfried März
  165. Department of Preventive Medicine, Keck School of Medicine, University of Southern California, Los Angeles, California 90089, USA.

    • Fredrick R. Schumacher
    •  & Christopher A. Haiman
  166. National Cancer Institute, Bethesda, Maryland 20892, USA.

    • Jianxin Shi
  167. Icelandic Heart Association, Kopavogur 201, Iceland.

    • Albert Vernon Smith
    •  & Vilmundur Gudnason
  168. University of Iceland, Reykjavik 101, Iceland.

    • Albert Vernon Smith
    •  & Vilmundur Gudnason
  169. 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
  170. Molecular & Cellular Therapeutics, Royal College of Surgeons in Ireland, 123 St Stephen’s Green, Dublin 2, Ireland.

    • Alice V. Stanton
  171. deCODE Genetics, Amgen Inc., Reykjavik 101, Iceland.

    • Valgerdur Steinthorsdottir
    • , Gudmar Thorleifsson
    • , Unnur Thorsteinsdottir
    •  & Kari Stefansson
  172. Department of Medical Sciences, Molecular Medicine, Uppsala University, Uppsala 75144, Sweden.

    • Ann-Christine Syvänen
  173. National Heart and Lung Institute, Imperial College London, London SW3 6LY, UK.

    • Sian-Tsung Tan
    • , Miriam F. Moffatt
    •  & Jaspal S. Kooner
  174. Department of Public Health Sciences, Stritch School of Medicine, Loyola University of Chicago, Maywood, Illinois 61053, USA.

    • Bamidele O. Tayo
    •  & Richard S. Cooper
  175. Institute of Epidemiology II, Helmholtz Zentrum München - German Research Center for Environmental Health, Neuherberg, Germany, D-85764 Neuherberg, Germany.

    • Barbara Thorand
    •  & Annette Peters
  176. Department of Oncology, University of Cambridge, Cambridge CB2 0QQ, UK.

    • Jonathan P. Tyrer
  177. Centre for Bone and Arthritis Research, Department of Internal Medicine and Clinical Nutrition, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, Gothenburg 413 45, Sweden.

    • Liesbeth Vandenput
    •  & Claes Ohlsson
  178. Department of Child and Adolescent Psychiatry/Psychology, Erasmus MC University Medical Centre, 3000 CB Rotterdam, The Netherlands.

    • Frank C. Verhulst
  179. Department for Health Evidence, Radboud University Medical Centre, 6500 HB Nijmegen, The Netherlands.

    • Sita H. Vermeulen
    •  & Lambertus A. Kiemeney
  180. Department of Genetics, Radboud University Medical Centre, 6500 HB Nijmegen, The Netherlands.

    • Sita H. Vermeulen
  181. 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
  182. Genetics, GlaxoSmithKline, King of Prussia, Pennsylvania 19406, USA.

    • Dawn Waterworth
  183. German Center for Cardiovascular Research, partner site Hamburg/Lubeck/Kiel, 23562 Lubeck, Germany.

    • Christina Willenborg
    •  & Jeanette Erdmann
  184. Institut für Integrative und Experimentelle Genomik, Universität zu Lübeck, D-23562 Lübeck, Germany.

    • Christina Willenborg
    •  & Jeanette Erdmann
  185. Department of Community Medicine, Faculty of Health Sciences, UiT The Arctic University of Norway, 9037 Tromsø, Norway.

    • Tom Wilsgaard
    •  & Inger Njølstad
  186. MRC Unit for Lifelong Health and Ageing at University College London, London WC1B 5JU, UK.

    • Andrew Wong
    • , Ken K. Ong
    •  & Diana Kuh
  187. Diabetes Complications Research Centre, Conway Institute, School of Medicine and Medical Sciences, University College Dublin, Dublin 4, Ireland.

    • Eoin P. Brennan
  188. Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul, Korea.

    • Murim Choi
  189. Lady Davis Institute, Departments of Human Genetics, Epidemiology and Biostatistics, McGill University, Montréal, Québec H3T1E2, Canada.

    • Zari Dastani
  190. Cardiothoracic Surgery Unit, Department of Molecular Medicine and Surgery, Karolinska Institutet, Stockholm 17176, Sweden.

    • Anders Franco-Cereceda
  191. Department of Medicine, Columbia University College of Physicians and Surgeons, New York 10032, USA.

    • Ali G. Gharavi
    •  & Krzysztof Kiryluk
  192. Biosciences Research Division, Department of Primary Industries, Victoria 3083, Australia.

    • Michael E. Goddard
  193. Department of Food and Agricultural Systems, University of Melbourne, Victoria 3010, Australia.

    • Michael E. Goddard
  194. Department of Epidemiology, Harvard School of Public Health, Boston, Massachusetts 02115, USA.

    • Jinyan Huang
    • , Liming Liang
    • , Baoshan Ma
    •  & David J. Hunter
  195. 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
  196. NIHR Oxford Biomedical Research Centre, OUH Trust, Oxford OX3 7LE, UK.

    • Fredrik Karpe
  197. Cardiovascular Research Center, Massachusetts General Hospital, Harvard Medical School, Boston, Massachusetts, USA.

    • Sekar Kathiresan
  198. Laboratory for Genotyping Development, RIKEN Center for Integrative Medical Sciences, Yokohama 230-0045, Japan.

    • Michiaki Kubo
  199. Center for Genome Science, National Institute of Health, Chungcheongbuk-do, Chungbuk 363–951, Republic of Korea.

    • Jong-Young Lee
  200. Harvard School of Public Health, Department of Biostatistics, Harvard University, Boston, Massachusetts 2115, USA.

    • Liming Liang
  201. Department of Genetics, Howard Hughes Medical Institute, Yale University School of Medicine, New Haven, New Haven, Connecticut 06520, USA.

    • Richard P. Lifton
  202. College of Information Science and Technology, Dalian Maritime University, Dalian, Liaoning 116026, China.

    • Baoshan Ma
  203. Nephrology Research, Centre for Public Health, Queen’s University of Belfast, Belfast, County Down BT9 7AB, UK.

    • Amy J. McKnight
  204. Section of General Internal Medicine, Boston University School of Medicine, Boston, Massachusetts 02118, USA.

    • Joanne M. Murabito
  205. Department of Statistics, University of Oxford, 1 South Parks Road, Oxford OX1 3TG, UK.

    • George Nicholson
  206. MRC Harwell, Harwell Science and Innovation Campus, Harwell OX11 0QG, UK.

    • George Nicholson
  207. Institute of Health and Biomedical Innovation, Queensland University of Technology, Brisbane, Queensland 4059, Australia.

    • Dale R. Nyholt
  208. Laboratory for Statistical Analysis, RIKEN Center for Integrative Medical Sciences, Yokohama 230-0045, Japan.

    • Yukinori Okada
    •  & Atsushi Takahashi
  209. 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
  210. Genome Institute of Singapore, Agency for Science, Technology and Research, 138672 Singapore.

    • Rajkumar Dorajoo
  211. Department of Biomedical Engineering and Computational Science, Aalto University School of Science, Helsinki FI-00076, Finland.

    • Niina Sandholm
  212. Department of Medicine, Division of Nephrology, Helsinki University Central Hospital, FI-00290 Helsinki, Finland.

    • Niina Sandholm
  213. Folkhälsan Institute of Genetics, Folkhälsan Research Center, FI-00290 Helsinki, Finland.

    • Niina Sandholm
  214. Laboratory for Cardiovascular Diseases, RIKEN Center for Integrative Medical Sciences, Yokohama 230-0045, Japan.

    • Toshihiro Tanaka
  215. Division of Disease Diversity, Bioresource Research Center, Tokyo Medical and Dental University, 113-8510 Tokyo, Japan.

    • Toshihiro Tanaka
  216. Division of Epidemiology, Department of Medicine; Vanderbilt Epidemiology Center; and Vanderbilt-Ingram Cancer Center, Vanderbilt University Medical Center, Nashville, Tennessee 37075, USA.

    • Wei Zheng
  217. Nuffield Department of Obstetrics & Gynaecology, University of Oxford, Oxford OX3 7BN, UK.

    • Krina T. Zondervan
  218. Department of Psychiatry, Washington University School of Medicine, St Louis, Missouri 63110, USA.

    • Andrew C. Heath
    • , Pamela A. F. Madden
    • , D. C. Rao
    •  & Treva K. Rice
  219. Department of Epidemiology and Public Health, EA3430, University of Strasbourg, Faculty of Medicine, Strasbourg, France.

    • Dominique Arveiler
  220. Department of Internal Medicine, University Medical Center Groningen, University of Groningen, 9700RB Groningen, The Netherlands.

    • Stephan J. L. Bakker
    •  & Ron T. Gansevoort
  221. Pathology and Laboratory Medicine, The University of Western Australia, Perth, Western Australia 6009, Australia.

    • John Beilby
    •  & Jennie Hui
  222. Cedars-Sinai Diabetes and Obesity Research Institute, Los Angeles, California 90048, USA.

    • Richard N. Bergman
  223. Institute of Social and Preventive Medicine (IUMSP), Centre Hospitalier Universitaire Vaudois and University of Lausanne, 1010 Lausanne, Switzerland.

    • Pascal Bovet
    •  & Murielle Bochud
  224. Ministry of Health, Victoria, Republic of Seychelles.

    • Pascal Bovet
    •  & Murielle Bochud
  225. University of Milano, Bicocca, 20126, Italy.

    • Giancarlo Cesana
  226. Harvard Medical School, Boston, Massachusetts 02115, USA.

    • Daniel I. Chasman
    •  & Paul M. Ridker
  227. Center for Human Genetics Research, Vanderbilt University Medical Center, Nashville, Tennessee 37203, USA.

    • Dana C. Crawford
  228. Department of Molecular Physiology and Biophysics, Vanderbilt University, Nashville, Tennessee 37232, USA.

    • Dana C. Crawford
  229. Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts 02118, USA.

    • L. Adrienne Cupples
  230. Department of Health Sciences, University of Milano, I 20142, Italy.

    • Daniele Cusi
  231. Fondazione Filarete, Milano I 20139, Italy.

    • Daniele Cusi
  232. Department of Public Health and Primary Care, University of Cambridge, Cambridge CB1 8RN, UK.

    • John Danesh
    •  & Danish Saleheen
  233. Julius Center for Health Sciences and Primary Care, University Medical Center Utrecht, 3584 CX Utrecht, The Netherlands.

    • Hester M. den Ruijter
  234. Institute of Cardiovascular and Medical Sciences, College of Medical, Veterinary and Life Sciences, University of Glasgow, Glasgow G12 8TA, UK.

    • Anna F. Dominiczak
  235. Clinic of Cardiology, West-German Heart Centre, University Hospital Essen, 45147 Essen, Germany.

    • Raimund Erbel
  236. Department of General Practice and Primary Health Care, University of Helsinki, FI-00290 Helsinki, Finland.

    • Johan G. Eriksson
  237. Unit of General Practice, Helsinki University Central Hospital, Helsinki 00290, Finland.

    • Johan G. Eriksson
  238. Department of Internal Medicine B, University Medicine Greifswald, D-17475 Greifswald, Germany.

    • Stephan B. Felix
  239. 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
  240. Department of Internal Medicine, University of Pisa, 56100 Pisa, Italy.

    • Ele Ferrannini
  241. National Research Council Institute of Clinical Physiology, University of Pisa, 56124 Pisa, Italy.

    • Ele Ferrannini
  242. Department of Cardiology, Toulouse University School of Medicine, Rangueil Hospital, 31400 Toulouse, France.

    • Jean Ferrières
  243. Robertson Center for Biostatistics, University of Glasgow, Glasgow G12 8QQ, UK.

    • Ian Ford
  244. UWI Solutions for Developing Countries, The University of the West Indies, Mona, Kingston 7, Jamaica.

    • Terrence Forrester
    •  & Colin A. McKenzie
  245. NorthShore University HealthSystem, Evanston, IL 60201, University of Chicago, Chicago, Illinois, USA.

    • Pablo V. Gejman
  246. Leeds MRC Medical Bioinformatics Centre, University of Leeds, Leeds LS2 9LU, UK.

    • Alistair S. Hall
  247. Institute of Biomedical & Clinical Science, University of Exeter, Barrack Road, Exeter EX2 5DW, UK.

    • Andrew T. Hattersley
  248. 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
  249. Division of Genomic Medicine, National Human Genome Research Institute, National Institutes of Health, Bethesda, Maryland 20892, USA.

    • Lucia A. Hindorff
  250. Institute of Cardiovascular Science, University College London, London WC1E 6BT, UK.

    • Aroon D. Hingorani
    •  & Folkert W. Asselbergs
  251. Department of Vascular Medicine, Academic Medical Center, 1105 AZ Amsterdam, The Netherlands.

    • G. Kees Hovingh
    •  & John J. P. Kastelein
  252. Centre for Cardiovascular Genetics, Institute Cardiovascular Sciences, University College London, London WC1E 6JJ, UK.

    • Steve E. Humphries
  253. Cardiovascular Genetics Division, Department of Internal Medicine, University of Utah, Salt Lake City, Utah 84108, USA.

    • Steven C. Hunt
  254. Sansom Institute for Health Research, University of South Australia, Adelaide 5000, South Australia, Australia.

    • Elina Hyppönen
  255. School of Population Health, University of South Australia, Adelaide 5000, South Australia, Australia.

    • Elina Hyppönen
  256. South Australian Health and Medical Research Institute, Adelaide, South Australia 5000, Australia.

    • Elina Hyppönen
  257. Population, Policy, and Practice, University College London Institute of Child Health, London WC1N 1EH, UK.

    • Elina Hyppönen
    •  & Chris Power
  258. Hannover Unified Biobank, Hannover Medical School, Hannover, D-30625 Hannover, Germany.

    • Thomas Illig
  259. National Institute for Health and Welfare, FI-90101 Oulu, Finland.

    • Marjo-Riitta Jarvelin
  260. MRC Health Protection Agency (HPA) Centre for Environment and Health, School of Public Health, Imperial College London, London W2 1PG, UK.

    • Marjo-Riitta Jarvelin
  261. Unit of Primary Care, Oulu University Hospital, FI-90220 Oulu, Finland.

    • Marjo-Riitta Jarvelin
  262. Biocenter Oulu, University of Oulu, FI-90014 Oulu, Finland.

  263. Institute of Health Sciences, University of Oulu, FI-90014 Oulu, Finland.

    • Marjo-Riitta Jarvelin
    •  & Sirkka M. Keinanen-Kiukaanniemi
  264. 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
  265. Interuniversity Cardiology Institute of the Netherlands (ICIN), 3501 DG Utrecht, The Netherlands.

    • J. Wouter Jukema
  266. Unit of Primary Health Care/General Practice, Oulu University Hospital, FI-90220 Oulu, Finland.

    • Sirkka M. Keinanen-Kiukaanniemi
  267. Department of Urology, Radboud University Medical Centre, 6500 HB Nijmegen, The Netherlands.

    • Lambertus A. Kiemeney
  268. Imperial College Healthcare NHS Trust, London W12 0HS, UK.

    • Jaspal S. Kooner
    •  & John C. Chambers
  269. Department of Epidemiology and Public Health, University College London, London WC1E 6BT, UK.

    • Meena Kumari
    • , Claudia Langenberg
    •  & Mika Kivimaki
  270. Department of Biological and Social Epidemiology, University of Essex, Wivenhoe Park, Colchester, Essex CO4 3SQ, UK.

    • Meena Kumari
  271. Department of Medicine, Kuopio University Hospital and University of Eastern Finland, FI-70210 Kuopio, Finland.

    • Johanna Kuusisto
    •  & Markku Laakso
  272. Department of Physiology, Institute of Biomedicine, University of Eastern Finland, Kuopio Campus, FI-70211 Kuopio, Finland.

    • Timo A. Lakka
  273. Department of Clinical Physiology and Nuclear Medicine, Kuopio University Hospital and University of Eastern Finland, FI-70210 Kuopio, Finland.

    • Timo A. Lakka
    •  & Rainer Rauramaa
  274. Department of Clinical Chemistry, Fimlab Laboratories and School of Medicine University of Tampere, FI-33520 Tampere, Finland.

    • Terho Lehtimäki
  275. Steno Diabetes Center A/S, Gentofte DK-2820, Denmark.

    • Valeriya Lyssenko
  276. Lund University Diabetes Centre and Department of Clinical Science, Diabetes & Endocrinology Unit, Lund University, Malmö 221 00, Sweden.

    • Valeriya Lyssenko
    •  & Leif C. Groop
  277. Institut Universitaire de Cardiologie et de Pneumologie de Québec, Faculty of Medicine, Laval University, Quebec, QC G1V 0A6, Canada.

    • André Marette
  278. Institute of Nutrition and Functional Foods, Laval University, Quebec, QC G1V 0A6, Canada.

    • André Marette
    • , Marie-Claude Vohl
    •  & Louis Pérusse
  279. Department of Biostatistics, University of Washington, Seattle, Washington 98195, USA.

    • Barbara McKnight
  280. Department of Surgery, University Medical Center Utrecht, 3584 CX Utrecht, The Netherlands.

    • Frans L. Moll
  281. Department of Biostatistics, University of Liverpool, Liverpool L69 3GA, UK.

    • Andrew P. Morris
  282. Department of Pediatrics, University of Iowa, Iowa City, Iowa 52242, USA.

    • Jeffrey C. Murray
  283. Illumina, Inc, Little Chesterford, Cambridge CB10 1XL, UK.

    • John F. Peden
  284. University of Groningen, University Medical Center Groningen, Department of Pulmonary Medicine and Tuberculosis, Groningen, The Netherlands.

    • Dirkje S. Postma
  285. Department of Neurology, General Central Hospital, Bolzano 39100, Italy.

    • Peter P. Pramstaller
  286. Department of Clinical Physiology and Nuclear Medicine, Turku University Hospital, FI-20521 Turku, Finland.

    • Olli T. Raitakari
  287. Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, FI-20521 Turku, Finland.

    • Olli T. Raitakari
  288. Human Genomics Laboratory, Pennington Biomedical Research Center, Baton Rouge, Louisiana 70808, USA.

    • Tuomo Rankinen
    • , Mark A. Sarzynski
    •  & Claude Bouchard
  289. Université de Montréal, Montreal, Quebec H1T 1C8, Canada.

    • John D. Rioux
  290. Center for Systems Genomics, The Pennsylvania State University, University Park, Pennsylvania 16802, USA.

    • Marylyn D. Ritchie
  291. Croatian Centre for Global Health, Faculty of Medicine, University of Split, 21000 Split, Croatia.

    • Igor Rudan
  292. South Carelia Central Hospital, 53130 Lappeenranta, Finland.

    • Jouko Saramies
  293. Paul Langerhans Institute Dresden, German Center for Diabetes Research (DZD), 01307 Dresden, Germany.

    • Peter E. H. Schwarz
  294. International Centre for Circulatory Health, Imperial College London, London W2 1PG, UK.

    • Peter Sever
  295. Division of Endocrinology, Diabetes and Nutrition, University of Maryland School of Medicine, Baltimore, Maryland 21201, USA.

    • Alan R. Shuldiner
    •  & Jeffrey R. O’Connell
  296. Program for Personalized and Genomic Medicine, University of Maryland School of Medicine, Baltimore, Maryland 21201, USA.

    • Alan R. Shuldiner
    •  & Jeffrey R. O’Connell
  297. Geriatric Research and Education Clinical Center, Vetrans Administration Medical Center, Baltimore, Maryland 21201, USA.

    • Alan R. Shuldiner
  298. Helsinki University Central Hospital Heart and Lung Center, Department of Medicine, Helsinki University Central Hospital, FI-00290 Helsinki, Finland.

    • Juha Sinisalo
  299. Sorbonne Universités, UPMC Univ Paris 06, UMR S 1166, F-75013 Paris, France.

    • David-Alexandre Trégouët
  300. INSERM, UMR S 1166, Team Genomics and Physiopathology of Cardiovascular Diseases, F-75013 Paris, France.

    • David-Alexandre Trégouët
  301. Institute for Cardiometabolism And Nutrition (ICAN), F-75013 Paris, France.

    • David-Alexandre Trégouët
  302. Department of Kinesiology, Laval University, Quebec QC G1V 0A6, Canada.

    • Angelo Tremblay
    •  & Louis Pérusse
  303. 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
  304. Department of Food Science and Nutrition, Laval University, Quebec QC G1V 0A6, Canada.

    • Marie-Claude Vohl
  305. Department of Internal Medicine, University Hospital (CHUV) and University of Lausanne, Lausanne 1011, Switzerland.

    • Gérard Waeber
  306. Department of Nutrition, University of North Carolina, Chapel Hill, North Carolina 27599, USA.

    • Linda S. Adair
  307. Institut Pasteur de Lille; INSERM, U744; Université de Lille 2; F-59000 Lille, France.

    • Philippe Amouyel
  308. Department of Cardiology, Division Heart and Lungs, University Medical Center Utrecht, 3584 CX Utrecht, The Netherlands.

    • Folkert W. Asselbergs
  309. Department of Medicine, Stanford University School of Medicine, Palo Alto, California 94304, USA.

    • Themistocles L. Assimes
    • , Mads Melbye
    •  & Thomas Quertermous
  310. Lee Kong Chian School of Medicine, Imperial College London and Nanyang Technological University, Singapore, 637553 Singapore, Singapore.

    • Bernhard O. Boehm
  311. Department of Internal Medicine I, Ulm University Medical Centre, D-89081 Ulm, Germany.

    • Bernhard O. Boehm
  312. Health Science Center at Houston, University of Texas, Houston, Texas 77030, USA.

    • Eric Boerwinkle
  313. Department of Medicine, Division of Genetics, Brigham and Women’s Hospital, Harvard Medical School, Boston, Massachusetts 02115, USA.

    • Paul I. W. de Bakker
  314. Department of Epidemiology, University Medical Center Utrecht, 3584 CX Utrecht, The Netherlands.

    • Paul I. W. de Bakker
  315. School of Population Health, The University of Western Australia, Nedlands, Western Australia 6009, Australia.

    • Jennie Hui
  316. Albert Einstein College of Medicine, Department of Epidemiology and Population Health, Belfer 1306, New York 10461, USA.

    • Robert C. Kaplan
  317. Center for Human Genetics, Division of Public Health Sciences, Wake Forest School of Medicine, Winston-Salem, North Carolina 27157, USA.

    • Yongmei Liu
  318. Synlab Academy, Synlab Services GmbH, 68163 Mannheim, Germany.

    • Winfried März
  319. Department of Clinical Medicine, Copenhagen University, 2200 Copenhagen, Denmark.

    • Mads Melbye
  320. Department of Clinical Genetics, Erasmus MC University Medical Center, 3000 CA Rotterdam, The Netherlands.

    • Ben A. Oostra
  321. Finnish Diabetes Association, Kirjoniementie 15, FI-33680 Tampere, Finland.

    • Timo E. Saaristo
  322. Pirkanmaa Hospital District, FI-33521 Tampere, Finland.

    • Timo E. Saaristo
  323. Center for Non-Communicable Diseases, Karatchi, Pakistan.

    • Danish Saleheen
  324. Department of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania 19104, USA.

    • Danish Saleheen
  325. BHF Glasgow Cardiovascular Research Centre, Division of Cardiovascular and Medical Sciences, University of Glasgow, Glasgow G12 8TA, UK.

    • Naveed Sattar
  326. Icahn Institute for Genomics and Multiscale Biology, Icahn School of Medicine at Mount Sinai, New York, New York 10580, USA.

    • Eric E. Schadt
  327. Faculty of Medicine, University of Iceland, Reykjavik 101, Iceland.

    • Unnur Thorsteinsdottir
    •  & Kari Stefansson
  328. Institute for Health Research, University Hospital of La Paz (IdiPaz), 28046 Madrid, Spain.

    • Jaakko Tuomilehto
  329. Diabetes Research Group, King Abdulaziz University, 21589 Jeddah, Saudi Arabia.

    • Jaakko Tuomilehto
  330. Centre for Vascular Prevention, Danube-University Krems, 3500 Krems, Austria.

    • Jaakko Tuomilehto
  331. Department of Public Health and Clinical Nutrition, University of Eastern Finland, Finland.

    • Matti Uusitupa
  332. Research Unit, Kuopio University Hospital, FI-70210 Kuopio, Finland.

    • Matti Uusitupa
  333. Institute of Cellular Medicine, Newcastle University, Newcastle NE1 7RU, UK.

    • Mark Walker
  334. Institute of Clinical Chemistry and Laboratory Medicine, University Medicine Greifswald, D-17475 Greifswald, Germany.

    • Henri Wallaschofski
  335. Institute of Medical Informatics, Biometry and Epidemiology, Chair of Epidemiology, Ludwig-Maximilians-Universität, D-85764 Munich, Germany.

    • H-Erich Wichmann
  336. Klinikum Grosshadern, D-81377 Munich, Germany.

    • H-Erich Wichmann
  337. Institute of Epidemiology I, Helmholtz Zentrum München - German Research Center for Environmental Health, Neuherberg, Germany, D-85764 Neuherberg, Germany.

    • H-Erich Wichmann
  338. Department of Pulmonology, University Medical Center Utrecht, 3584 CX Utrecht, The Netherlands.

    • Pieter Zanen
  339. Princess Al-Jawhara Al-Brahim Centre of Excellence in Research of Hereditary Disorders (PACER-HD), King Abdulaziz University, 21589 Jeddah, Saudi Arabia.

    • Panos Deloukas
  340. Division of Population Health Sciences & Education, St George’s, University of London, London SW17 0RE, UK.

    • David P. Strachan
  341. Oxford NIHR Biomedical Research Centre, Oxford University Hospitals NHS Trust, Oxford OX3 7LJ, UK.

    • Mark I. McCarthy
  342. Clinical Epidemiology, Integrated Research and Treatment Center, Center for Sepsis Control and Care (CSCC), Jena University Hospital, 07743 Jena, Germany.

    • André Scherag
  343. Department of Human Genetics, University of Michigan, Ann Arbor, Michigan 48109, USA.

    • Cristen J. Willer
  344. Service of Medical Genetics, CHUV University Hospital, 1011 Lausanne, Switzerland.

    • Jacques S. Beckmann
  345. University of Cambridge Metabolic Research Laboratories, Institute of Metabolic Science, Addenbrooke’s Hospital, Cambridge CB2 OQQ, UK.

    • Inês Barroso
  346. NIHR Cambridge Biomedical Research Centre, Institute of Metabolic Science, Addenbrooke’s Hospital, Cambridge CB2 OQQ, UK.

    • Inês Barroso
  347. Carolina Center for Genome Sciences, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina 27599, USA.

    • Kari E. North
  348. The Mindich Child Health and Development Institute, Icahn School of Medicine at Mount Sinai, New York, New York 10029, USA.

    • Ruth J. F. Loos

Consortia

  1. The LifeLines Cohort Study

    A list of authors and affiliations appears in the Supplementary Information.

  2. The ADIPOGen Consortium

    A list of authors and affiliations appears in the Supplementary Information.

  3. The AGEN-BMI Working Group

    A list of authors and affiliations appears in the Supplementary Information.

  4. The CARDIOGRAMplusC4D Consortium

    A list of authors and affiliations appears in the Supplementary Information.

  5. The CKDGen Consortium

    A list of authors and affiliations appears in the Supplementary Information.

  6. The GLGC

    A list of authors and affiliations appears in the Supplementary Information.

  7. The ICBP

    A list of authors and affiliations appears in the Supplementary Information.

  8. The MAGIC Investigators

    A list of authors and affiliations appears in the Supplementary Information.

  9. The MuTHER Consortium

    A list of authors and affiliations appears in the Supplementary Information.

  10. The MIGen Consortium

    A list of authors and affiliations appears in the Supplementary Information.

  11. The PAGE Consortium

    A list of authors and affiliations appears in the Supplementary Information.

  12. The ReproGen Consortium

    A list of authors and affiliations appears in the Supplementary Information.

  13. The GENIE Consortium

    A list of authors and affiliations appears in the Supplementary Information.

  14. The International Endogene Consortium

    A list of authors and affiliations appears in the Supplementary Information.

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