Article | Published:

Genome-wide meta-analysis identifies 11 new loci for anthropometric traits and provides insights into genetic architecture

Nature Genetics volume 45, pages 501512 (2013) | Download Citation

Abstract

Approaches exploiting trait distribution extremes may be used to identify loci associated with common traits, but it is unknown whether these loci are generalizable to the broader population. In a genome-wide search for loci associated with the upper versus the lower 5th percentiles of body mass index, height and waist-to-hip ratio, as well as clinical classes of obesity, including up to 263,407 individuals of European ancestry, we identified 4 new loci (IGFBP4, H6PD, RSRC1 and PPP2R2A) influencing height detected in the distribution tails and 7 new loci (HNF4G, RPTOR, GNAT2, MRPS33P4, ADCY9, HS6ST3 and ZZZ3) for clinical classes of obesity. Further, we find a large overlap in genetic structure and the distribution of variants between traits based on extremes and the general population and little etiological heterogeneity between obesity subgroups.

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Acknowledgements

A full list of acknowledgments appears in the Supplementary Note. Funding was provided by the Aarno Koskelo Foundation; the Academy of Finland; the Agency for Science, Technology and Research of Singapore; the Australian National Health and Medical Research Council; the Australian Research Council; BDA Research; the BioSHaRE Consortium; the British Heart Foundation; the Cedars-Sinai Board of Governors' Chair in Medical Genetics; the Centre for Clinical Research at the University of Leipzig; the Centre of Excellence in Genomics and the University of Tartu; the Chief Scientist Office of the Scottish government; the City of Kuopio and the Social Insurance Institution of Finland; the Department of Educational Assistance, the University and Research of the Autonomous Province of Bolzano; the Donald W. Reynolds Foundation; the Dutch Ministry for Health, Welfare and Sports; the Dutch Ministry of Education, Culture and Science; Dutch BBRMI-NL; the Dutch Brain Foundation; the Dutch Centre for Medical Systems Biology; the Dutch Diabetes Research Foundation; the Dutch Government Economic Structure–Enhancing Fund; the Dutch Inter-University Cardiology Institute; the Dutch Kidney Foundation; the Dutch Ministry of Economic Affairs; the Dutch Ministry of Justice; the Dutch Research Institute for Diseases in the Elderly; Eleanor Nichols endowments; the Emil Aaltonen Foundation; Erasmus Medical Center and Erasmus University; the Estonian government; the European Commission; the European Regional Development Fund; the European Research Council; the European Science Foundation; the Faculty of Biology and Medicine of Lausanne; Finland's Slot Machine Association; the Finnish Cultural Foundation; the Finnish Diabetes Research Foundation; the Finnish Foundation for Cardiovascular Research; the Finnish Funding Agency for Technology and Innovation; the Finnish Heart Association; the Finnish Medical Society; the Finnish Ministry of Education and Culture; the Finnish Ministry of Health and Social Affairs; the Finnish National Institute for Health and Welfare; the Finnish Social Insurance Institution; Finska Läkaresällskapet; the Folkhälsan Research Foundation; the Foundation for Life and Health in Finland; the French Ministry of Research; the French National Research Agency; the Genetic Association Information Network; the German Diabetes Association; the German Federal Ministry of Education and Research; the German Ministry of Cultural Affairs; the German National Genome Research Network; the German Research Foundation; GlaxoSmithKline; the Göteborg Medical Society; the Greek General Secretary of Research and Technology; the Gyllenberg Foundation; Health Care Centers in Vasa, Närpes and Korsholm; the Heinz Nixdorf Foundation; Helmholtz Zentrum München–German Research Center for Environmental Health; the Icelandic Heart Association; the Icelandic Parliament; the Intramural Research Program of the Division of Cancer Epidemiology and Genetics, the National Cancer Institute, NIH; Italian Ministry of Education, Universities and Research; Italian Ministry of Health; Juho Vainio Foundation; Juvenile Diabetes Research Foundation International; the Knut and Alice Wallenberg Foundation; Kuopio, Tampere and Turku University Hospital Medical Funds; the Leducq Foundation; the Lundberg Foundation; the March of Dimes; the Munich Center of Health Sciences as part of LMUinnovativ; the Municipal Health Care Center and Hospital in Jakobstad; the Municipality of Rotterdam; the Närpes Health Care Foundation; National Alliance for Research on Schizophrenia and Depression Young Investigator Awards; the Netherlands Genomics Initiative; the Netherlands Organization for Health Research and Development; UK NHSBT; US National Institutes of Health; the Nordic Center of Cardiovascular Research; the Nordic Center of Excellence in Disease Genetics; the Nordic Centre of Excellence on Systems Biology in Controlled Dietary Interventions and Cohort Studies; the Northern Netherlands Collaboration of Provinces; the Novo Nordisk Foundation; the Ollqvist Foundation; the Orion-Farmos Research Foundation; the Paavo Nurmi Foundation; the Päivikki and Sakari Sohlberg Foundation; the Perklen Foundation; the Petrus and Augusta Hedlunds Foundation; the Province of Groningen; the Republic of Croatia Ministry of Science, Education and Sport; the Reynold's Foundation; the Royal Society; Samfundet Folkhälsan; the Signe and Ane Gyllenberg Foundation; the Sigrid Juselius Foundation; the Social Ministry of the Federal State of Mecklenburg–West Pomerania; the Sophia Foundation for Medical Research; the South Tyrolean Sparkasse Foundation; the Southern California Diabetes Endocrinology Research Center; the Stockholm County Council; the Strategic Cardiovascular Program of Karolinska Institutet; Strategic Support for Epidemiological Research at Karolinska Institutet; the Susan G. Komen Breast Cancer Foundation; the Swedish Ministry for Higher Education; the Swedish Cancer Society; the Swedish Cultural Foundation in Finland; the Swedish Diabetes Association; the Swedish Foundation for Strategic Research; the Swedish Heart-Lung Foundation; the Swedish Medical Research Council; the Swedish Ministry of Education; the Swedish Research Council; the Swedish Royal Academy of Science; the Swedish Society for Medical Research; the Swedish Society of Medicine; the Swiss National Science Foundation; the Tampere Tuberculosis Foundation; The Great Wine Estates of the Margaret River Region of Western Australia; The Paul Michael Donovan Charitable Foundation; the Torsten and Ragnar Söderberg Foundation; Cancer Research UK; the UK Diabetes Association; the UK Heart Foundation; the UK MRC; the UK NIHR, Biomedical Research Centre; UK West Anglia Primary and Community Care; the University Medical Center Groningen and the University of Groningen; the Västra Götaland Foundation; VU University: the Institute for Health and Care Research and the Neuroscience Campus Amsterdam; the Wellcome Trust; and the Yrjö Jahnsson Foundation.

Author information

Author notes

    • Sonja I Berndt
    • , Stefan Gustafsson
    • , Reedik Mägi
    •  & Andrea Ganna

    These authors contributed equally to this work.

    • Mark I McCarthy
    • , Elizabeth K Speliotes
    • , Kari E North
    •  & Ruth J F Loos

    These authors jointly directed this work.

Affiliations

  1. US Department of Health and Human Services, Division of Cancer Epidemiology and Genetics, National Cancer Institute, US National Institutes of Health, Bethesda, Maryland, USA.

    • Sonja I Berndt
    • , Jianxin Shi
    • , Kevin B Jacobs
    • , Zhaoming Wang
    •  & Stephen J Chanock
  2. Department of Medical Sciences, Molecular Epidemiology and Science for Life Laboratory, Uppsala University, Uppsala, Sweden.

    • Stefan Gustafsson
    •  & Erik Ingelsson
  3. Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden.

    • Stefan Gustafsson
    • , Andrea Ganna
    • , Tove Fall
    • , Henrik Grönberg
    • , Per Hall
    • , Patrik K Magnusson
    • , Nancy Pedersen
    • , Yudi Pawitan
    •  & Erik Ingelsson
  4. Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, UK.

    • Reedik Mägi
    • , Teresa Ferreira
    • , Joshua C Randall
    • , Josine L Min
    • , Inga Prokopenko
    • , Cecilia M Lindgren
    • , Andrew P Morris
    • , Mark I McCarthy
    •  & Erik Ingelsson
  5. Estonian Genome Center, University of Tartu, Tartu, Estonia.

    • Reedik Mägi
    • , Tõnu Esko
    • , Krista Fischer
    • , Markus Perola
    • , Evelin Mihailov
    • , Lili Milani
    •  & Andres Metspalu
  6. Wellcome Trust Sanger Institute, Hinxton, Cambridge, UK.

    • Eleanor Wheeler
    • , Joshua C Randall
    • , Stavroula Kanoni
    • , Sarah Edkins
    • , Sarah E Hunt
    • , Aarno Palotie
    • , Aparna Radhakrishnan
    • , So-Youn Shin
    • , Nicole Soranzo
    • , Kathleen Stirrups
    • , Willem H Ouwehand
    • , Samuli Ripatti
    • , Inês Barroso
    •  & Panos Deloukas
  7. Department of Genetics, Washington University School of Medicine, St. Louis, Missouri, USA.

    • Mary F Feitosa
    • , Aldi T Kraja
    • , Michael A Province
    •  & Ingrid B Borecki
  8. Department of Epidemiology, School of Public Health, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, USA.

    • Anne E Justice
    • , Keri L Monda
    •  & Kari E North
  9. Center for Observational Research, Amgen, Thousands Oaks, California, USA.

    • Keri L Monda
  10. Department of Genetics, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, USA.

    • Damien C Croteau-Chonka
    •  & Karen L Mohlke
  11. Medical Research Council (MRC) Epidemiology Unit, Institute of Metabolic Science, Addenbrooke's Hospital, Cambridge, UK.

    • Felix R Day
    • , Jian'an Luan
    • , Jing Hua Zhao
    • , Claudia Langenberg
    • , Ken K Ong
    • , Nicholas J Wareham
    •  & Ruth J F Loos
  12. Institute of Molecular and Cell Biology, University of Tartu, Tartu, Estonia.

    • Tõnu Esko
    • , Evelin Mihailov
    •  & Andres Metspalu
  13. Molecular Biology Department, Istituto Auxologico Italiano, Milan, Italy.

    • Davide Gentilini
    •  & Anna Maria Di Blasio
  14. Department of Biostatistics, Center for Statistical Genetics, University of Michigan, Ann Arbor, Michigan, USA.

    • Anne U Jackson
    • , Jennifer L Bragg-Gresham
    • , Goncalo R Abecasis
    •  & Michael Boehnke
  15. Divisions of Genetics and Endocrinology and Center for Basic and Translational Obesity Research, Children's Hospital, Boston, Massachusetts, USA.

    • Sailaja Vedantam
    •  & Joel N Hirschhorn
  16. Metabolism Initiative and Program in Medical and Population Genetics, Broad Institute, Cambridge, Massachusetts, USA.

    • Sailaja Vedantam
    • , Sekar Kathiresan
    •  & Joel N Hirschhorn
  17. Department of Genetics, Harvard Medical School, Boston, Massachusetts, USA.

    • Sailaja Vedantam
    •  & Joel N Hirschhorn
  18. Department of Internal Medicine (Cardiovascular), University of Michigan, Ann Arbor, Michigan, USA.

    • Cristen J Willer
  19. Department of Human Genetics, University of Michigan, Ann Arbor, Michigan, USA.

    • Cristen J Willer
  20. Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, Michigan, USA.

    • Cristen J Willer
    •  & Elizabeth K Speliotes
  21. Public Health and Gender Studies, Institute of Epidemiology and Preventive Medicine, Regensburg University Medical Center, Regensburg, Germany.

    • Thomas W Winkler
    •  & Iris M Heid
  22. Genetics of Complex Traits, Peninsula College of Medicine and Dentistry, University of Exeter, Exeter, UK.

    • Andrew R Wood
    • , Timothy Frayling
    •  & Erik Ingelsson
  23. Department of Nutrition, Harvard School of Public Health, Boston, Massachusetts, USA.

    • Tsegaselassie Workalemahu
    • , Frank B Hu
    • , Paul W Franks
    • , Lu Qi
    •  & David Hunter
  24. Channing Laboratory, Department of Medicine, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts, USA.

    • Tsegaselassie Workalemahu
    • , Lu Qi
    •  & David Hunter
  25. Department of Biostatistics and Bioinformatics, Emory University, Atlanta, Georgia, USA.

    • Yi-Juan Hu
  26. The Queensland Brain Institute, The University of Queensland, Brisbane, Queensland, Australia.

    • Sang Hong Lee
    •  & Peter M Visscher
  27. Department of Epidemiology, Harvard School of Public Health, Boston, Massachusetts, USA.

    • Liming Liang
  28. Department of Biostatistics, Harvard School of Public Health, Boston, Massachusetts, USA.

    • Liming Liang
  29. Department of Biostatistics, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, USA.

    • Dan-Yu Lin
  30. Analytic and Translational Genetics Unit, Massachusetts General Hospital, Boston, Massachusetts, USA.

    • Benjamin M Neale
  31. deCODE Genetics, Reykjavik, Iceland.

    • Gudmar Thorleifsson
    • , Kari Stefansson
    • , Valgerdur Steinthorsdottir
    •  & Unnur Thorsteinsdottir
  32. University of Queensland Diamantina Institute, The University of Queensland, Princess Alexandra Hospital, Brisbane, Queensland, Australia.

    • Jian Yang
    •  & Peter M Visscher
  33. Queensland Institute of Medical Research, Brisbane, Queensland, Australia.

    • Jian Yang
    • , Sarah E Medland
    • , Grant W Montgomery
    •  & Nicholas G Martin
  34. Institute of Genetic Epidemiology, Helmholtz Zentrum München–German Research Center for Environmental Health, Neuherberg, Germany.

    • Eva Albrecht
    • , Martina Müller-Nurasyid
    •  & Christian Gieger
  35. Department of Epidemiology, Erasmus Medical Center, Rotterdam, The Netherlands.

    • Najaf Amin
    • , Carolina Medina-Gomez
    • , Marjolein J Peters
    • , Karol Estrada
    • , Fernando Rivadeneira
    • , Joyce B J van Meurs
    • , Jacqueline C M Witteman
    • , Albert Hofman
    • , André G Uitterlinden
    •  & Cornelia M van Duijn
  36. Genetic Epidemiology and Biostatistics Platform, Ontario Institute for Cancer Research, Toronto, Ontario, Canada.

    • Gemma Cadby
    •  & Lyle J Palmer
  37. Prosserman Centre for Health Research, Samuel Lunenfeld Research Institute, Toronto, Ontario, Canada.

    • Gemma Cadby
    •  & Lyle J Palmer
  38. Centre for Genetic Epidemiology and Biostatistics, The University of Western Australia, Crawley, Western Australia, Australia.

    • Gemma Cadby
  39. Department of Internal Medicine, VU University Medical Centre, Amsterdam, The Netherlands.

    • Martin den Heijer
  40. Unit of Public Health Genomics, Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, Finland.

    • Niina Eklund
    • , Kati Kristiansson
    • , Markus Perola
    •  & Ida Surakka
  41. Cardiovascular Medicine, University of Oxford, Wellcome Trust Centre for Human Genetics, Oxford, UK.

    • Anuj Goel
    • , Martin Farrall
    • , Samuli Ripatti
    •  & Hugh Watkins
  42. Department of Biological Psychology, VU University Amsterdam, Amsterdam, The Netherlands.

    • Jouke-Jan Hottenga
    • , Gonneke Willemsen
    •  & Dorret I Boomsma
  43. MRC Human Genetics Unit, MRC Institute for Genetics and Molecular Medicine, Western General Hospital, Edinburgh, UK.

    • Jennifer E Huffman
    • , Caroline Hayward
    • , Veronique Vitart
    •  & Alan F Wright
  44. Institute of Medical Biometry and Epidemiology, University of Marburg, Marburg, Germany.

    • Ivonne Jarick
  45. Department of Immunology, Genetics and Pathology, Uppsala University, Uppsala, Sweden.

    • Åsa Johansson
    •  & Ulf Gyllensten
  46. Uppsala Clinical Research Center, Uppsala University Hospital, Uppsala, Sweden.

    • Åsa Johansson
  47. Genome Centre, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London, UK.

    • Toby Johnson
    • , Patricia B Munroe
    •  & Mark J Caulfield
  48. Clinical Pharmacology, William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London, UK.

    • Toby Johnson
    • , Patricia B Munroe
    •  & Mark J Caulfield
  49. Ludwigshafen Risk and Cardiovascular Health (LURIC) Study, Freiburg, Germany.

    • Marcus E Kleber
  50. Mannheim Institute of Public Health, Social and Preventive Medicine, Medical Faculty of Mannheim, University of Heidelberg, Mannheim, Germany.

    • Marcus E Kleber
    •  & Winfried März
  51. Institut für Medizinische Biometrie und Statistik, Universität zu Lübeck, Universitätsklinikum Schleswig-Holstein, Campus Lübeck, Lübeck, Germany.

    • Inke R König
    •  & Michael Preuss
  52. Department of Medical Genetics, University of Lausanne, Lausanne, Switzerland.

    • Zoltán Kutalik
    •  & Diana Marek
  53. Swiss Institute of Bioinformatics, Lausanne, Switzerland.

    • Zoltán Kutalik
    •  & Diana Marek
  54. Department of Medical Genetics, Division of Genetic Epidemiology, Molecular and Clinical Pharmacology, Innsbruck Medical University, Innsbruck, Austria.

    • Claudia Lamina
  55. University Lille Nord de France, Lille, France.

    • Cecile Lecoeur
    • , Boris Skrobek
    • , Philippe Froguel
    •  & David Meyre
  56. Centre National de la Recherche Scientifique (CNRS) Unité Mixte de Recherche (UMR) 8199, Institut de Biologie de Lille (IBL)–Institut Pasteur de Lille, Lille, France.

    • Cecile Lecoeur
    • , Boris Skrobek
    • , Philippe Froguel
    •  & David Meyre
  57. Cardiovascular Health Research Unit, University of Washington, Seattle, Washington, USA.

    • Guo Li
    •  & Bruce M Psaty
  58. Department of Twin Research and Genetic Epidemiology, King's College London, London, UK.

    • Massimo Mangino
    • , Nicole Soranzo
    •  & Timothy D Spector
  59. School of Social and Community Medicine, University of Bristol, Bristol, UK.

    • Wendy L McArdle
  60. Department of Internal Medicine, Erasmus Medical Center, Rotterdam, The Netherlands.

    • Carolina Medina-Gomez
    • , Marjolein J Peters
    • , Karol Estrada
    • , Fernando Rivadeneira
    • , Joyce B J van Meurs
    • , M Carola Zillikens
    •  & André G Uitterlinden
  61. Netherlands Genomics Initiative (NGI)-sponsored Netherlands Consortium for Healthy Aging (NCHA), The Netherlands.

    • Carolina Medina-Gomez
    • , Marjolein J Peters
    • , Karol Estrada
    • , Ben A Oostra
    • , Fernando Rivadeneira
    • , Joyce B J van Meurs
    • , Jacqueline C M Witteman
    • , M Carola Zillikens
    • , Albert Hofman
    • , André G Uitterlinden
    •  & Cornelia M van Duijn
  62. Department of Medicine I, University Hospital Grosshadern, Ludwig-Maximilians-Universität, Munich, Germany.

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

    • Martina Müller-Nurasyid
    •  & H-Erich Wichmann
  64. Chair of Genetic Epidemiology, Institute of Medical Informatics, Biometry and Epidemiology, Ludwig-Maximilians-Universität, Munich, Germany.

    • Martina Müller-Nurasyid
  65. Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts, USA.

    • Julius S Ngwa
    •  & L Adrienne Cupples
  66. Department of Epidemiology, University of Groningen, University Medical Center Groningen, Groningen, The Netherlands.

    • Ilja M Nolte
    • , Ronald P Stolk
    •  & Harold Snieder
  67. MRC Centre for Causal Analyses in Translational Epidemiology, School of Social and Community Medicine, University of Bristol, Bristol, UK.

    • Lavinia Paternoster
    •  & David M Evans
  68. Institute for Medical Informatics, Biometry and Epidemiology (IMIBE), University Hospital of Essen, University of Duisburg-Essen, Essen, Germany.

    • Sonali Pechlivanis
    • , Carolin Pütter
    • , Karl-Heinz Jöckel
    •  & André Scherag
  69. Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland.

    • Markus Perola
    • , Ida Surakka
    • , Aarno Palotie
    • , Elisabeth Widén
    •  & Samuli Ripatti
  70. Medizinische Klinik II, Universität zu Lübeck, Lübeck, Germany.

    • Michael Preuss
    •  & Jeanette Erdmann
  71. Division of Preventive Medicine, Brigham and Women's Hospital, Boston, Massachusetts, USA.

    • Lynda M Rose
    • , Daniel I Chasman
    •  & Paul M Ridker
  72. Genetic and Molecular Epidemiology Unit, Department of Clinical Sciences, Skåne University Hospital Malmö, Lund University, Malmö, Sweden.

    • Dmitry Shungin
    •  & Paul W Franks
  73. Department of Public Health & Clinical Medicine, Umeå University, Umeå, Sweden.

    • Dmitry Shungin
    •  & Paul W Franks
  74. Department of Odontology, Umeå University, Umeå, Sweden.

    • Dmitry Shungin
  75. Icelandic Heart Association, Kopavogur, Iceland.

    • Albert Vernon Smith
    •  & Vilmundur Gudnason
  76. Faculty of Medicine, University of Iceland, Reykjavik, Iceland.

    • Albert Vernon Smith
    • , Vilmundur Gudnason
    • , Kari Stefansson
    •  & Unnur Thorsteinsdottir
  77. Atherosclerosis Research Unit, Department of Medicine, Solna, Karolinska Institutet, Karolinska University Hospital, Stockholm, Sweden.

    • Rona J Strawbridge
    •  & Anders Hamsten
  78. Interfaculty Institute for Genetics and Functional Genomics, Ernst-Moritz-Arndt-University Greifswald, Greifswald, Germany.

    • Alexander Teumer
    •  & Georg Homuth
  79. Department of Vascular Medicine, Academic Medical Center, Amsterdam, The Netherlands.

    • Mieke D Trip
    • , Hanneke Basart
    •  & Kees G Hovingh
  80. Heart Failure Research Centre, Department of Clinical and Experimental Cardiology, Academic Medical Center, Amsterdam, The Netherlands.

    • Mieke D Trip
  81. Department of Oncology, University of Cambridge, Cambridge, UK.

    • Jonathan Tyrer
  82. Department of Endocrinology, University Medical Center Groningen, University of Groningen, Groningen, The Netherlands.

    • Jana V Van Vliet-Ostaptchouk
    • , Melanie M Van der Klauw
    •  & Bruce H R Wolffenbuttel
  83. LifeLines Cohort Study, University Medical Center Groningen, University of Groningen, Groningen, The Netherlands.

    • Jana V Van Vliet-Ostaptchouk
    • , Marcel Bruinenberg
    • , Lude Franke
    • , Ronald P Stolk
    • , Melanie M Van der Klauw
    • , Bruce H R Wolffenbuttel
    •  & Harold Snieder
  84. Department of Internal Medicine, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, Gothenburg, Sweden.

    • Liesbeth Vandenput
    •  & Mattias Lorentzon
  85. Hudson Alpha Institute for Biotechnology, Huntsville, Alabama, USA.

    • Lindsay L Waite
    • , Devin Absher
    •  & Claes Ohlsson
  86. Department of Cardiology, Division of Heart & Lungs, University Medical Center Utrecht, Utrecht, The Netherlands.

    • Folkert W Asselbergs
  87. Institute of Biomedicine/Physiology, University of Eastern Finland, Kuopio Campus, Kuopio, Finland.

    • Mustafa Atalay
    •  & Timo A Lakka
  88. National Institute for Health Research (NIHR) Cambridge Biomedical Research Centre, Cambridge, UK.

    • Antony P Attwood
    • , Augusto Rendon
    •  & Willem H Ouwehand
  89. Division of Epidemiology, Multidisciplinary Cardiovascular Research Centre (MCRC), Leeds Institute of Genetics, Health and Therapeutics (LIGHT), University of Leeds, Leeds, UK.

    • Anthony J Balmforth
  90. Department of Molecular Genetics, PathWest Laboratory of Western Australia, Queen Elizabeth II Medical Centre, Nedlands, Western Australia, Australia.

    • John Beilby
  91. Department of Surgery and Pathology, University of Western Australia, Nedlands, Western Australia, Australia.

    • John Beilby
  92. Genome Technology Branch, National Human Genome Research Institute, US National Institutes of Health, Bethesda, Maryland, USA.

    • Lori L Bonnycastle
    • , Peter S Chines
    • , Francis S Collins
    • , Narisu Narisu
    •  & Amy J Swift
  93. Dipartimento di Medicina Sperimentale. Università degli Studi Milano-Bicocca, Monza, Italy.

    • Paolo Brambilla
  94. Centre for Population Health Sciences, University of Edinburgh, Edinburgh, UK.

    • Harry Campbell
    • , Igor Rudan
    • , Sarah H Wild
    •  & James F Wilson
  95. Harvard Medical School, Boston, Massachusetts, USA.

    • Daniel I Chasman
    •  & Paul M Ridker
  96. British Heart Foundation Glasgow Cardiovascular Research Centre, University of Glasgow, Glasgow, UK.

    • John M Connell
  97. University of Dundee, Ninewells Hospital & Medical School, Dundee, UK.

    • John M Connell
  98. National Heart and Lung Institute, Imperial College London, London, UK.

    • William O Cookson
    •  & Miriam F Moffatt
  99. Division of Cardiovascular Epidemiology, Institute of Environmental Medicine, Karolinska Institutet, Stockholm, Sweden.

    • Ulf de Faire
  100. Department of Epidemiology, Biostatistics and HTA, Radboud University Nijmegen Medical Centre, Nijmegen, The Netherlands.

    • Femmie de Vegt
    • , Sita H Vermeulen
    •  & Lambertus A Kiemeney
  101. Istituto di Ricerca Genetica e Biomedica del Consiglio Nazionale delle Ricerche (CNR), Monserrato, Italy.

    • Mariano Dei
    •  & Serena Sanna
  102. Department of Dietetics-Nutrition, Harokopio University, Athens, Greece.

    • Maria Dimitriou
    • , Eirini V Theodoraki
    •  & George V Dedoussis
  103. Epidemiology and Preventive Medicine Research Center, Department of Clinical and Experimental Medicine, University of Insubria, Varese, Italy.

    • Marco M Ferrario
  104. Department of Cardiology, Toulouse University School of Medicine, Rangueil Hospital, Toulouse, France.

    • Jean Ferrières
  105. Department of Genetics, University Medical Center Groningen, University of Groningen, Groningen, The Netherlands.

    • Lude Franke
    •  & Pim van der Harst
  106. Department of Health Sciences, University of Milan, Ospedale San Paolo, Milan, Italy.

    • Francesca Frau
    •  & Daniele Cusi
  107. University of Chicago, Chicago, Illinois, USA.

    • Pablo V Gejman
    •  & Alan R Sanders
  108. Northshore University Health System, Evanston, Illinois, USA.

    • Pablo V Gejman
    •  & Alan R Sanders
  109. Research Unit for Molecular Epidemiology, Helmholtz Zentrum München–German Research Center for Environmental Health, Neuherberg, Germany.

    • Harald Grallert
    • , Annette Peters
    •  & Thomas Illig
  110. Division of Cardiovascular and Neuronal Remodelling, Multidisciplinary Cardiovascular Research Centre, LIGHT, University of Leeds, Leeds, UK.

    • Alistair S Hall
  111. Department of Clinical Sciences/Obstetrics and Gynecology, University of Oulu, Oulu, Finland.

    • Anna-Liisa Hartikainen
    •  & Anneli Pouta
  112. Department of Neurology, Boston University School of Medicine, Boston, Massachusetts, USA.

    • Nancy L Heard-Costa
  113. Department of Psychiatry, Washington University School of Medicine, St. Louis, Missouri, USA.

    • Andrew C Heath
    •  & Pamela A Madden
  114. Department of Child and Adolescent Psychiatry, University of Duisburg-Essen, Essen, Germany.

    • Johannes Hebebrand
    •  & Anke Hinney
  115. Centre For Paediatric Epidemiology and Biostatistics, MRC Centre of Epidemiology for Child Health, University College London Institute of Child Health, London, UK.

    • Elina Hyppönen
    •  & Chris Power
  116. Division of Research, Kaiser Permanente Northern California, Oakland, California, USA.

    • Carlos Iribarren
  117. Core Genotyping Facility, SAIC-Frederick, Inc., National Cancer Institute (NCI)-Frederick, Frederick, Maryland, USA.

    • Kevin B Jacobs
    •  & Zhaoming Wang
  118. Department of Physiology, Institute of Neuroscience and Physiology, Sahlgrenska Academy, University of Gothenburg, Gothenburg, Sweden.

    • John-Olov Jansson
  119. Population Studies Unit, Department of Chronic Disease Prevention, National Institute for Health and Welfare, Turku, Finland.

    • Antti Jula
  120. Department of Clinical Physiology, University of Tampere and Tampere University Hospital, Tampere, Finland.

    • Mika Kähönen
  121. Cardiovascular Research Center, Massachusetts General Hospital, Boston, Massachusetts, USA.

    • Sekar Kathiresan
  122. Cardiology Division, Massachusetts General Hospital, Boston, Massachusetts, USA.

    • Sekar Kathiresan
  123. Center for Human Genetic Research, Massachusetts General Hospital, Boston, Massachusetts, USA.

    • Sekar Kathiresan
  124. UK Clinical Research Collaboration (UKCRC) Centre of Excellence for Public Health (Northern Ireland), Queens University, Belfast, UK.

    • Frank Kee
  125. Department of Public Health and Primary Care, Institute of Public Health, University of Cambridge, Cambridge, UK.

    • Kay-Tee Khaw
  126. Department of Epidemiology and Public Health, University College London, London, UK.

    • Mika Kivimäki
    • , Meena Kumari
    • , Claudia Langenberg
    •  & Aroon Hingorani
  127. Department of Internal Medicine II–Cardiology, University of Ulm Medical Center, Ulm, Germany.

    • Wolfgang Koenig
  128. Chronic Disease Epidemiology and Prevention Unit, Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, Finland.

    • Kari Kuulasmaa
    • , Jarmo Virtamo
    •  & Veikko Salomaa
  129. Department of Medicine, University of Eastern Finland, Kuopio Campus and Kuopio University Hospital, Kuopio, Finland.

    • Johanna Kuusisto
    •  & Markku Laakso
  130. Finnish Institute of Occupational Health, Oulu, Finland.

    • Jaana H Laitinen
  131. Kuopio Research Institute of Exercise Medicine, Kuopio, Finland.

    • Timo A Lakka
    •  & Rainer Rauramaa
  132. Laboratory of Epidemiology, Demography, Biometry, National Institute on Aging, US National Institutes of Health, Bethesda, Maryland, USA.

    • Lenore J Launer
    •  & Tamara B Harris
  133. Department of Medical Sciences, Uppsala University, Akademiska Sjukhuset, Uppsala, Sweden.

    • Lars Lind
  134. Diabetes Prevention Unit, National Institute for Health and Welfare, Helsinki, Finland.

    • Jaana Lindström
    •  & Jaakko Tuomilehto
  135. Human Genetics, Genome Institute of Singapore, Singapore.

    • Jianjun Liu
  136. Department of Internal Medicine, Istituto Auxologico Italiano, Verbania, Italy.

    • Antonio Liuzzi
  137. Transplantation Laboratory, Haartman Institute, University of Helsinki, Helsinki, Finland.

    • Marja-Liisa Lokki
  138. Chair of Nephrology San Raffaele Scientific Institute, Università Vita-Salute San Raffaele, Unit Ospedaliera Nephrology and Dialysis, Milan, Italy.

    • Paolo Manunta
  139. Synlab Academy, Mannheim, Germany.

    • Winfried März
  140. Department of Cardiology, University Medical Center Groningen, University of Groningen, Groningen, The Netherlands.

    • Irene Mateo Leach
    •  & Pim van der Harst
  141. Department of Biostatistics, University of Washington, Seattle, Washington, USA.

    • Barbara McKnight
  142. Genetics Division, GlaxoSmithKline, King of Prussia, Pennsylvania, USA.

    • Vincent Mooser
  143. Institute of Human Genetics, University of Bonn, Bonn, Germany.

    • Thomas W Mühleisen
  144. Department of Genomics, Life & Brain Center, University of Bonn, Bonn, Germany.

    • Thomas W Mühleisen
  145. School of Population Health, The University of Western Australia, Nedlands, Western Australia, Australia.

    • Arthur W Musk
  146. Department of Respiratory Medicine, Sir Charles Gairdner Hospital, Nedlands, Western Australia, Australia.

    • Arthur W Musk
  147. Busselton Population Medical Research Foundation, Sir Charles Gairdner Hospital, Nedlands, Western Australia, Australia.

    • Arthur W Musk
  148. Department of Internal Medicine, University Medical Center Groningen, University of Groningen, Groningen, The Netherlands.

    • Gerjan Navis
  149. MRC Harwell, Harwell, UK.

    • George Nicholson
  150. Department of Statistics, University of Oxford, Oxford, UK.

    • George Nicholson
  151. Section of Epidemiology, Department of Public Health, Aarhus University, Aarhus, Denmark.

    • Ellen A Nohr
  152. MRC Unit for Lifelong Health & Ageing, London, UK.

    • Ken K Ong
    • , Andrew Wong
    •  & Diana Kuh
  153. Department of Clinical Genetics, Erasmus Medical Center, Rotterdam, The Netherlands.

    • Ben A Oostra
  154. Center of Medical Systems Biology, Leiden University Medical Center, Leiden, The Netherlands.

    • Ben A Oostra
    •  & Cornelia M van Duijn
  155. Medical Research Institute, University of Dundee, Ninewells Hospital and Medical School, Dundee, UK.

    • Colin N A Palmer
    •  & Andrew D Morris
  156. Illumina, Inc., Cambridge, UK.

    • John F Peden
  157. Institute of Epidemiology II, Helmholtz Zentrum München–German Research Center for Environmental Health, Neuherberg, Germany.

    • Annette Peters
    •  & Barbara Thorand
  158. Munich Heart Alliance, Munich, Germany.

    • Annette Peters
  159. Faculty of Medicine, University of Split, Split, Croatia.

    • Ozren Polasek
  160. National Institute for Health and Welfare, Oulu, Finland.

    • Anneli Pouta
    •  & Marjo-Riitta Jarvelin
  161. Center for Biomedicine, European Academy Bozen/Bolzano (EURAC), Bolzano/Bozen, Italy (affiliated institute of the University of Lübeck).

    • Peter P Pramstaller
    •  & Andrew A Hicks
  162. Department of Neurology, General Central Hospital, Bolzano, Italy.

    • Peter P Pramstaller
  163. Department of Neurology, University of Lübeck, Lübeck, Germany.

    • Peter P Pramstaller
  164. Oxford Centre for Diabetes, Endocrinology and Metabolism, University of Oxford, Oxford, UK.

    • Inga Prokopenko
    • , Fredrik Karpe
    •  & Mark I McCarthy
  165. Department of Haematology, University of Cambridge, Cambridge, UK.

    • Aparna Radhakrishnan
    • , Augusto Rendon
    • , Jennifer G Sambrook
    • , Jonathan C Stephens
    •  & Willem H Ouwehand
  166. National Health Service (NHS) Blood and Transplant, Cambridge Centre, Cambridge, UK.

    • Aparna Radhakrishnan
    • , Augusto Rendon
    • , Jennifer G Sambrook
    • , Jonathan C Stephens
    •  & Willem H Ouwehand
  167. Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku, Finland.

    • Olli Raitakari
  168. The Department of Clinical Physiology and Nuclear Medicine, Turku University Hospital, Turku, Finland.

    • Olli Raitakari
  169. MRC Biostatistics Unit, Institute of Public Health, Cambridge, UK.

    • Augusto Rendon
  170. Finnish Diabetes Association, Tampere, Finland.

    • Timo E Saaristo
  171. Pirkanmaa Hospital District, Tampere, Finland.

    • Timo E Saaristo
  172. South Karelia Central Hospital, Lappeenranta, Finland.

    • Jouko Saramies
  173. Institute for Community Medicine, University Medicine Greifswald, Greifswald, Germany.

    • Sabine Schipf
  174. Institute for Clinical Molecular Biology, Christian-Albrechts University, Kiel, Germany.

    • Stefan Schreiber
  175. Deutsches Herzzentrum München and DZHK (German Centre for Cardiovascular Research), partner site Munich Heart Alliance, Munich, Germany.

    • Heribert Schunkert
    •  & Jeanette Erdmann
  176. Azienda Ospedaliera di Desio e Vimercate, Milan, Italy.

    • Stefano Signorini
  177. Division of Cardiology, Cardiovascular Laboratory, Helsinki University Central Hospital, Helsinki, Finland.

    • Juha Sinisalo
    •  & Markku S Nieminen
  178. University of Eastern Finland and Kuopio University Hospital, Kuopio, Finland.

    • Alena Stančáková
  179. Klinik und Poliklinik für Innere Medizin II, Universitätklinikum Regensburg, Regensburg, Germany.

    • Klaus Stark
    •  & Christian Hengstenberg
  180. Department of Medicine, University of Leipzig, Leipzig, Germany.

    • Michael Stumvoll
    •  & Anke Tönjes
  181. Integrierte Forschungs- und Behandlungszentren (IFB) Adiposity Diseases, University of Leipzig, Leipzig, Germany.

    • Michael Stumvoll
    •  & Anke Tönjes
  182. Institut National de la Santé et de la Recherche Médicale (INSERM) Unité Mixte de Recherche Scientifique (UMRS) 937, Institute of Cardiometabolism And Nutrition (ICAN), Pierre et Marie Curie Medical School, Paris, France.

    • David-Alexandre Tregouet
  183. Dipartimento di Scienze Farmacologiche e Biomolecolari, Università di Milano, Centro Cardiologico Monzino, Istituto di Ricovero e Cura a Carattere Scientifico (IRCCS), Milan, Italy.

    • Elena Tremoli
  184. Department of Human Genetics, Radboud University Nijmegen Medical Centre, Nijmegen, The Netherlands.

    • Sita H Vermeulen
  185. Department of Medicine, University of Turku and Turku University Hospital, Turku, Finland.

    • Jorma Viikari
  186. Department of Internal Medicine, Centre Hospitalier Universitaire Vaudois (CHUV) University Hospital, Lausanne, Switzerland.

    • Gérard Waeber
    •  & Peter Vollenweider
  187. Cardiology Group, Frankfurt-Sachsenhausen, Germany.

    • Bernhard R Winkelmann
  188. Institut Pasteur de Lille, INSERM U744, Université Lille Nord de France, Lille, France.

    • Philippe Amouyel
  189. Department of Medicine, Division of Endocrinology and Diabetes, University Hospital, Ulm, Germany.

    • Bernhard O Boehm
  190. Human Genetics Center, University of Texas Health Science Center, Houston, Texas, USA.

    • Eric Boerwinkle
  191. Institute of Molecular Medicine, University of Texas Health Science Center, Houston, Texas, USA.

    • Eric Boerwinkle
  192. Fondazione Filarete, Milan, Italy.

    • Daniele Cusi
  193. Department of General Practice and Primary Health Care, University of Helsinki, Helsinki, Finland.

    • Johan G Eriksson
  194. National Institute for Health and Welfare, Helsinki, Finland.

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

    • Johan G Eriksson
  196. Department of Genomics of Common Disease, School of Public Health, Imperial College London, London, UK.

    • Philippe Froguel
  197. Helseundersøkelsen i Nord-Trøndelag (HUNT) Research Centre, Department of Public Health and General Practice, Norwegian University of Science and Technology, Levanger, Norway.

    • Kristian Hveem
  198. Hannover Unified Biobank, Hannover Medical School, Hannover, Germany.

    • Thomas Illig
  199. Department of Epidemiology and Biostatistics, School of Public Health, Faculty of Medicine, Imperial College London, London, UK.

    • Marjo-Riitta Jarvelin
  200. Institute of Health Sciences, University of Oulu, Oulu, Finland.

    • Marjo-Riitta Jarvelin
  201. Biocenter Oulu, University of Oulu, Oulu, Finland.

    • Marjo-Riitta Jarvelin
    •  & Sirkka M Keinanen-Kiukaanniemi
  202. Unit of General Practice, Oulu University Hospital, Oulu, Finland.

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

    • Lambertus A Kiemeney
  204. Comprehensive Cancer Center East, Nijmegen, The Netherlands.

    • Lambertus A Kiemeney
  205. Department of Clinical Chemistry, Fimlab Laboratories, University of Tampere and Tampere University Hospital, Tampere, Finland.

    • Terho Lehtimäki
  206. Stanford University School of Medicine, Stanford, California, USA.

    • Douglas F Levinson
  207. Department of Clinical Medicine, Faculty of Health Sciences, University of Tromsø, Tromsø, Norway.

    • Inger Njølstad
  208. Department of Community Medicine, Faculty of Health Sciences, University of Tromsø, Tromsø, Norway.

    • Inger Njølstad
  209. Interdisciplinary Center Psychopathology and Emotion Regulation, University of Groningen, University Medical Center Groningen, Groningen, The Netherlands.

    • Albertine J Oldehinkel
  210. Department of Psychiatry, University Medical Centre Groningen, Groningen, The Netherlands.

    • Brenda Penninx
  211. Department of Epidemiology, University of Washington, Seattle, Washington, USA.

    • Bruce M Psaty
  212. Department of Medicine, University of Washington, Seattle, Washington, USA.

    • Bruce M Psaty
  213. Department of Health Services, University of Washington, Seattle, Washington, USA.

    • Bruce M Psaty
  214. Group Health Research Institute, Group Health, Seattle, Washington, USA.

    • Bruce M Psaty
  215. Department of Clinical Physiology and Nuclear Medicine, Kuopio University Hospital, Kuopio, Finland.

    • Rainer Rauramaa
  216. Department of Cardiovascular Sciences, University of Leicester, Glenfield Hospital, Leicester, UK.

    • Nilesh J Samani
  217. Leicester NIHR Biomedical Research Unit in Cardiovascular Disease, Glenfield Hospital, Leicester, UK.

    • Nilesh J Samani
  218. Institute of Preventive Medicine, Bispebjerg University Hospital, Copenhagen, Denmark.

    • Thorkild I A Sørensen
  219. Novo Nordisk Foundation Center for Basic Metabolic Research, University of Copenhagen, Copenhagen, Denmark.

    • Thorkild I A Sørensen
  220. Centre for Vascular Prevention, Danube-University Krems, Krems, Austria.

    • Jaakko Tuomilehto
  221. King Abdulaziz University, Jeddah, Saudi Arabia.

    • Jaakko Tuomilehto
  222. Institute of Public Health and Clinical Nutrition, University of Eastern Finland, Kuopio, Finland.

    • Matti Uusitupa
  223. Research Unit, Kuopio University Hospital, Kuopio, Finland.

    • Matti Uusitupa
  224. Institute of Clinical Chemistry and Laboratory Medicine, University Medicine Greifswald, Greifswald, Germany.

    • Henri Wallaschofski
  225. Institute of Epidemiology I, Helmholtz Zentrum München–German Research Center for Environmental Health, Neuherberg, Germany.

    • H-Erich Wichmann
    •  & Iris M Heid
  226. Klinikum Grosshadern, Munich, Germany.

    • H-Erich Wichmann
  227. Department of Medicine, Stanford University School of Medicine, Stanford, California, USA.

    • Themistocles L Assimes
  228. University of Cambridge Metabolic Research Laboratories, Institute of Metabolic Science Addenbrooke's Hospital, Cambridge, UK.

    • Inês Barroso
  229. Division of Intramural Research, National Heart, Lung, and Blood Institute, Framingham Heart Study, Framingham, Massachusetts, USA.

    • Caroline S Fox
  230. Lund University Diabetes Centre, Department of Clinical Sciences, Lund University, Malmö, Sweden.

    • Leif C Groop
  231. Medical Genetics Institute, Cedars-Sinai Medical Center, Los Angeles, California, USA.

    • Talin Haritunian
  232. Department of Epidemiology and Population Health, Albert Einstein College of Medicine, Bronx, New York, USA.

    • Robert C Kaplan
  233. Oxford NIHR Biomedical Research Centre, Churchill Hospital, Oxford, UK.

    • Fredrik Karpe
    •  & Mark I McCarthy
  234. Department of Medicine, University of Maryland School of Medicine, Baltimore, Maryland, USA.

    • Jeffrey R O'Connell
  235. Department of Genetics and Genomic Sciences, Mount Sinai School of Medicine, New York, New York, USA.

    • Eric E Schadt
  236. Institute of Genomics and Multiscale Biology, Mount Sinai School of Medicine, New York, New York, USA.

    • Eric E Schadt
  237. Laboratory of Genetics, National Institute on Aging, Baltimore, Maryland, USA.

    • David Schlessinger
  238. Division of Population Health Sciences and Education, St George's, University of London, London, UK.

    • David P Strachan
  239. Department of Clinical Epidemiology and Biostatistics, McMaster University, Hamilton, Ontario, Canada.

    • David Meyre
  240. Department of Internal Medicine, Division of Gastroenterology, University of Michigan, Ann Arbor, Michigan, USA.

    • Elizabeth K Speliotes
  241. The Charles Bronfman Institute of Personalized Medicine, Icahn School of Medicine at Mount Sinai, New York, New York, USA.

    • Ruth J F Loos
  242. The Mindich Child Health and Development Institute, Icahn School of Medicine at Mount Sinai, New York, New York, USA.

    • Ruth J F Loos
  243. Department of Preventive Medicine, Icahn School of Medicine at Mount Sinai, New York, New York, USA.

    • Ruth J F Loos

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Contributions

Steering committee (oversaw the consortium): G.R.A., T.L.A., I.B., S.I.B., M. Boehnke, I.B.B., P.D., C.S.F., T. Frayling, L.C.G., T.H., I.M.H., D.H., E.I., R.C.K., R.J.F.L., M.I.M., K.L. Mohlke, K.E.N., J.R.O., D. Schlessinger, D.P.S., U.T. and C.M.v.D. Writing group (drafted and edited the manuscript): S.I.B., M.F.F., A. Ganna, S.G., E.I., A.E.J., C.M.L., R.J.F.L., R.M., M.I.M., D. Meyre, K.L. Monda, A.P.M., K.E.N., A. Scherag, E.K.S., E. Wheeler and C.J.W. Data cleaning and preparation: S.I.B., D.C.C.-C., F.R.D., T.E., T. Fall, T. Ferreira, S.G., I.M.H., E.I., A.U.J., C.M.L., J. Luan, R.M., J.C.R., A. Scherag, E.K.S., G.T., S.V., T.W.W. and A.R.W. Statistical advisors: S.H.L., B.M.N., Y.P., P.M.V., J.Y., D.-Y.L. and Y.-J.H. Gene expression (eQTL) analyses: L. Liang, W.O.C., M.F.M., G.R.A., V. Steinthorsdottir, G.T., J.L.M., G. Nicholson, F. Karpe., M.I.M. and E.E.S.

Project design, management and coordination of contributing studies

Stage 1—GWAS: (ADVANCE) T.L.A. and C.I.; (AGES) V.G., T.B.H. and L.J.L.; (ARIC) E.B. and K.E.N.; (B58C) D.P.S.; (BRIGHT Study) M.J.C. and P.B.M.; (CAPS) E.I.; (CHS) B.M. and B.M.P.; (CoLaus) V.M., P.V. and G. Waeber; (COROGENE) M.S.N. and J. Sinisalo; (deCODE) K. Stefansson and U.T.; (DGI) L.C.G. and J.N.H.; (EGCUT) A.M.; (EPIC) K.-T.K., R.J.F.L. and N.J.W.; (ERF) B.A.O. and C.M.v.D.; (FamHS) I.B.B. and M.A.P.; (Fenland) R.J.F.L. and N.J.W.; (FRAM) L.A.C. and C.S.F.; (FUSION GWAS) M. Boehnke and K.L. Mohlke; (Genmets) A.J., S.R. and V. Salomaa; (GerMIFS1) J.E. and H. Schunkert; (GerMIFS2) C. Hengstenberg and K. Stark; (GOOD) C.O.; (HBCS) J.G.E.; (KORA S3) H.-E.W.; (KORA S4) C.G., T.I., W.K. and A. Peters; (MGS) P.V.G. and D.F.L.; (MICROS (SOUTH TYROL)) P.P.P.; (MIGEN) J.N.H. and S. Kathiresan; (NESDA) B.P.; (NFBC 1966) M.-R.J.; (NHS) L.Q.; (Nijmegen Biomedical Study) L.A.K.; (NSPHS) U.G.; (NTR) D.I.B.; (ORCADES) J.F.W. and A.F.W.; (PLCO) S.J.C. and S.I.B.; (PROCARDIS) M.F. and H. Watkins; (RS-I) F.R. and A.G.U.; (RUNMC) L.A.K.; (SardiNIA) D. Schlessinger; (SASBAC) E.I.; (SHIP) H. Wallaschofski; (Sorbs) M.S. and A. Tönjes; (TwinsUK) T.D.S.; (VIS) I.R.; (WGHS) P.M.R.; (WTCC-T2D) M.I.M.; (WTCCC-CAD) A.J.B., A.S.H. and N.J.S.; and (YFS) M. Kähönen, T.L., O.R. and J. Viikari. Stage 2—Metabochip and in silico replication: (AMC-PAS) K.G.H.; (B58C) C. Power; (BHS) L.J.P.; (DILGOM) K. Kuulasmaa and V. Salomaa; (DPS) M.U.; (DR's EXTRA) T.A.L. and R.R.; (EPIC, Fenland and Ely) C. Langenberg, R.J.F.L. and N.J.W.; (FIN-D2D 2007) S.M.K.-K. and T.E.S.; (GLACIER) P.W.F.; (Go-DARTS (Dundee)) A.D.M. and C.N.A.P.; (HNR) K.-H.J.; (HUNT 2) K.H.; (Hypergenes) D.C.; (IMPROVE) U.d.F., A. Hamsten and E.T.; (KORA S3) I.M.H.; (LifeLines Cohort Study) H. Snieder, M.M.V.d.K. and B.H.R.W.; (LURIC) B.O.B., W.M. and B.R.W.; (METSIM) J.K. and M. Laakso; (MORGAM) P.A., P.B., M.M.F., J.F., F. Kee, D.-A.T. and J. Virtamo; (NSHD) D.K.; (PIVUS) E.I.; (PLCO2) S.I.B. and S.J.C.; (PREVEND) P.v.d.H.; (QIMR) N.G.M., G.W.M., A.C.H. and P.M.; (RS-II) A. Hofman and J.B.J.v.M.; (RS-III) C.M.v.D. and J.C.M.W.; (Swedish Twin Registry) E.I.; (THISEAS/AMCPAS/CARDIOGENICS) P.D.; (THISEAS) G.V.D.; (TRAILS) A.J.O.; (Tromsø 4) I.N.; (TWINGENE) E.I.; (UKBS2) W.H.O.; (ULSAM) E.I.; (Whitehall II) A. Hingorani and M. Kivimäki; and (WTCC-T2D) M.I.M. and C.M.L. Other contributing studies, clinical extremes: (French Extreme Obesity Study) D. Meyre and P.F.; (GEO-IT) A.M.D.B.; (Essen Obesity Study, Essen Case-Control & Essen Obesity Trio GWAS) J.H. and A. Hinney; and (GOYA) T.I.A.S. and E.A.N.

Genotyping of contributing studies

Stage 1—GWAS: (ADVANCE) D.A.; (ARIC) E.B.; (B58C) W.L.M.; (CAPS) H. Grönberg; (CHS) T.H.; (CoLaus) V.M.; (COROGENE) M. Perola; (EGCUT) T.E. and L.M.; (EPIC) I.B.; (ERF) B.A.O. and C.M.v.D.; (FamHS) I.B.B., M.A.P. and A.T.K.; (Fenland) J. Luan; (Genmets) S.R.; (GOOD) C.O., J.-O.J. and M. Lorentzon; (HBCS) A. Palotie and E. Widén; (MGS) P.V.G. and A.R.S.; (MICROS (SOUTH TYROL)) A.A.H.; (NHS) F.B.H. and D.H.; (NSPHS) Å.J.; (NTR and NESDA) J.-J.H.; (ORCADES) J.F.W.; (PLCO) S.J.C. and K.B.J.; (RS-I) F.R., A.G.U., K.E. and C.M.-G.; (SardiNIA) M. Dei; (SASBAC) P.H. and J. Liu; (SHIP) G.H.; (TwinsUK) M.M., S.-Y.S. and N.S.; (VIS) C. Hayward and V.V.; (WGHS) D.I.C.; (WTCC-T2D) M.I.M.; (WTCCC-CAD) A.J.B., A.S.H. and N.J.S.; and (YFS) T.L. Stage 2—Metabochip and in silico replication: (BHS) L.J.P. and J.B.; (CARDIOGENICS) S.E. and S.E.H.; (DPS) A.J.S.; (DR's EXTRA) M.A.; (EPIC, Fenland and Ely) J. Luan and K.K.O.; (FIN-D2D 2007) P.S.C.; (FUSION) F.S.C., J. Saramies and J. Tuomilehto; (GLACIER) I.B. and S.E.; (Go-DARTS (Dundee)) C.N.A.P.; (HNR) T.W.M.; (HUNT 2) N.N.; (Hypergenes) F.F.; (KORA S3) H. Grallert; (KORA S4) T.I.; (LifeLines Cohort Study) H. Snieder, B.H.R.W., M. Bruinenberg and L.F.; (NSHD) D.K., K.K.O. and A.W.; (PIVUS) E.I. and L. Lind; (PLCO2) S.J.C., K.B.J. and Z.W.; (PREVEND) P.v.d.H. and F.W.A.; (QIMR) N.G.M., G.W.M., A.C.H. and P.A.M.; (RS-II) M.J.P. and M. Dei; (Swedish Twin Registry) E.I., P.K.M. and N.P.; (THISEAS/AMCPAS/CARDIOGENICS) K. Stirrups; (TRAILS) A.J.O., I.M.N. and J.V.V.V.-O.; (Tromsø 4) L.L.B.; (TWINGENE) E.I., A. Hamsten and N.P.; (ULSAM) E.I.; (Whitehall II) C. Langenberg; and (WTCC-T2D) M.I.M. Other contributing studies, clinical extremes: (French Extreme Obesity Study) D. Meyre and P.F.; (GEO-IT) D.G.; and (GOYA) L.P. and D.M.E.

Phenotyping of contributing studies

Stage 1—GWAS: (ARIC) E.B.; (B58C) D.P.S.; (BRIGHT Study) J.M.C.; (CAPS) H. Grönberg; (CHS) B.M.P.; (CoLaus) P.V. and G. Waeber; (COROGENE) J. Sinisalo, M.-L.L.; (EGCUT) A.M. and K.F.; (EPIC) R.J.F.L.; (ERF) B.A.O. and C.M.v.D.; (FamHS) I.B.B., M.A.P. and M.F.F.; (Fenland) R.J.F.L.; (FRAM) C.S.F.; (Genmets) A.J. and V. Salomaa; (GerMIFS1) S. Schreiber; (GerMIFS2) A. Peters; (GOOD) C.O., J.-O.J., M. Lorentzon and L.V.; (MGS) P.V.G., A.R.S. and D.F.L.; (NFBC 1966) A.-L.H., J.H.L. and A. Pouta; (NHS) L.Q.; (Nijmegen Biomedical Study) F.d.V., M.d.H. and S.H.V.; (NSPHS) Å.J. and U.G.; (NTR and NESDA) G. Willemsen; (ORCADES) H.C. and S.H.W.; (PLCO) S.I.B.; (RS-I) F.R. and A.G.U.; (SASBAC) P.H.; (SHIP) S. Schipf; (Sorbs) A. Tönjes; (TwinsUK) M.M. and T.D.S.; (VIS) O.P.; (WTCC-T2D) M.I.M.; (WTCCC-CAD) A.J.B., A.S.H. and N.J.S.; and (YFS) M. Kähönen, O.R. and J. Viikari. Stage 2—Metabochip and in silico replication: (AMC-PAS) H.B. and M.D.T.; (B58C) C. Power and E.H.; (BHS) L.J.P., J.B. and A.W.M.; (DPS) J. Lindström; (EPIC, Fenland and Ely) R.J.F.L.; (GLACIER) P.W.F. and D. Shungin; (Go-DARTS (Dundee)) C.N.A.P. and A.D.M.; (Hypergenes) D.C. and P.M.; (KORA S3) B.T.; (KORA S4) A. Peters; (LifeLines Cohort Study) B.H.R.W. and M.M.V.d.K.; (METSIM) A. Stančáková and P.V.G.; (NSHD) D.K.; (PIVUS) E.I. and L. Lind; (PLCO2) S.I.B.; (PREVEND) G. Navis; (QIMR) N.G.M., A.C.H. and P.M.; (RS-II) M.C.Z.; (RS-III) J.C.M.W.; (Swedish Twin Registry) E.I., P.K.M. and N.P.; (THISEAS) M. Dimitriou and E.V.T.; (TRAILS) R.P.S.; (TWINGENE) E.I. and N.P.; (UKBS2) A. Rendon; (ULSAM) E.I.; (Whitehall II) M. Kumari; and (WTCC-T2D) M.I.M. Other contributing studies, clinical extremes: (Essen Obesity Study, Essen Case-Control GWAS & Essen Obesity Trio GWAS) J.H. and A. Hinney; (GEO-IT) A.L. and S. Signorini; and (GOYA) T.I.A.S. and E.A.N.

Analyses of contributing studies

Stage 1—GWAS: (ADVANCE) L.L.W.; (AGES) A.V.S.; (ARIC) K.E.N., A.E.J. and K.L. Monda; (B58C) D.P.S.; (BRIGHT Study) T.J.; (CAPS) E.I. and R.M.; (CHS) B.M. and G.L.; (CoLaus) D. Marek; (COROGENE) M. Perola; (deCODE) V. Steinthorsdottir and G.T.; (DGI) E.K.S. and S.V.; (EGCUT) K.F., T.E. and E.M.; (EPIC) J.H.Z.; (ERF) N.A.; (FamHS) M.F.F.; (Fenland) J. Luan; (FRAM) L.A.C., N.L.H.-C. and J.S.N.; (Genmets) I.S.; (GerMIFS1) M. Preuss; (GerMIFS2) I.R.K.; (GOOD) C.O., J.-O.J., M. Lorentzon and L.V.; (HBCS) N.E.; (KORA S3) C. Lamina; (KORA S4) E.A.; (MGS) D.F.L. and J. Shi; (MICROS (SOUTH TYROL)) J.E.H. and Å.J.; (MIGEN) E.K.S. and S.V.; (NFBC 1966) A. Pouta, R.M. and J.C.R.; (NHS) L.Q. and T.W.; (NSPHS) Å.J.; (NTR and NESDA) J.- J.H.; (ORCADES) Å.J.; (PLCO) K.B.J. and S.I.B.; (PROCARDIS) M.F., A. Goel and J.F.P.; (RS-I) F.R., K.E. and C.M.-G.; (SardiNIA) J.L.B.-G. and S. Sanna; (SASBAC) E.I. and R.M.; (SEARCH) J. Tyrer; (SHIP) A. Teumer; (Sorbs) R.M. and I.P.; (TwinsUK) M.M. and N.S.; (VIS) Å.J.; (WGHS) D.I.C.; (WTCC-T2D) C.M.L., R.M. and J.C.R.; (WTCCC-CAD) R.M. and J.C.R.; (WTCCC-NBS (UKBS-CC)) A.P.A., R.M., J.C.R., J.G.S. and J.C.S.; and (YFS) O.R. and T.L. Stage 2—Metabochip and in silico replication: (B58C) E.H. and T. Ferreira; (BHS) G.C.; (DILGOM) K. Kristiansson and K. Kuulasmaa; (DPS) A.U.J.; (DR's EXTRA) A.U.J.; (EPIC, Fenland and Ely) J. Luan and K.K.O.; (FIN-D2D 2007) A.U.J.; (FUSION) A.U.J.; (GLACIER) P.W.F. and D. Shungin; (HNR) S.P. and C. Pütter; (HUNT 2) A.U.J.; (Hypergenes) F.F. and Z.K.; (IMPROVE) R.J.S.; (KORA S3) I.M.H. and T.W.W.; (KORA S4) M.M.-N.; (LifeLines Cohort Study) M. Bruinenberg and L.F.; (LURIC) M.E.K.; (METSIM) A.U.J.; (NSHD) A.W. and J. Luan; (PIVUS) E.I., S.G.; (PLCO2) S.I.B., Z.W.; (PREVEND) P.v.d.H. and I.M.L.; (QIMR) S.E.M., J.Y.; (RS-II) M.J.P.; (Swedish Twin Registry) E.I. and S.G.; (THISEAS/AMCPAS/CARDIOGENICS) S. Kanoni; (TRAILS) I.M.N. and J.V.V.V.-O.; (Tromsø 4) A.U.J.; (TWINGENE) E.I. and S.G.; (UKBS2) A. Radhakrishnan; (ULSAM) E.I., A. Ganna and S.G.; (WGHS) L.M.R.; and (WTCC-T2D) R.M. and T. Ferreira. Other contributing studies, clinical extremes: (French Extreme Obesity Study) D. Meyre, C. Lecoeur and B.S.; (GEO-IT) A.M.D.B. and D.G.; (Essen Obesity Study, Essen Case-Control GWAS and Essen Obesity Trio GWAS) A. Scherag and I.J.; and (GOYA) L.P. and D.M.E.

Competing interests

K. Stefansson, V. Steinthorsdottir, G.T. and U.T. are employed by deCODE Genetics with stock and/or stock options in the company. I.B. and spouse own stock in Incyte, Ltd., and GlaxoSmithKline. E.E.S. is Chief Scientific Officer at Pacific Biosciences. T.I.A.S. has collaborated with Nestlé Research Centre and DSM, The Netherlands, on the metabolomics and genomics of obesity and related traits, respectively.

Corresponding author

Correspondence to Erik Ingelsson.

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