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Defining the role of common variation in the genomic and biological architecture of adult human height

Nature Genetics 46, 1173–1186 (2014) | Download Citation

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Abstract

Using genome-wide data from 253,288 individuals, we identified 697 variants at genome-wide significance that together explained one-fifth of the heritability for adult height. By testing different numbers of variants in independent studies, we show that the most strongly associated ∼2,000, ∼3,700 and ∼9,500 SNPs explained ∼21%, ∼24% and ∼29% of phenotypic variance. Furthermore, all common variants together captured 60% of heritability. The 697 variants clustered in 423 loci were enriched for genes, pathways and tissue types known to be involved in growth and together implicated genes and pathways not highlighted in earlier efforts, such as signaling by fibroblast growth factors, WNT/β-catenin and chondroitin sulfate–related genes. We identified several genes and pathways not previously connected with human skeletal growth, including mTOR, osteoglycin and binding of hyaluronic acid. Our results indicate a genetic architecture for human height that is characterized by a very large but finite number (thousands) of causal variants.

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Acknowledgements

A full list of acknowledgments appears in the Supplementary Note.

Author information

    • Andrew R Wood
    • , Tonu Esko
    • , Jian Yang
    • , Sailaja Vedantam
    • , Tune H Pers
    • , Stefan Gustafsson
    •  & Michael E Goddard

    These authors contributed equally to this work.

    • Goncalo R Abecasis
    • , Daniel I Chasman
    • , Michael E Goddard
    • , Peter M Visscher
    • , Joel N Hirschhorn
    •  & Timothy M Frayling

    These authors jointly directed this work.

Affiliations

  1. Genetics of Complex Traits, University of Exeter Medical School, University of Exeter, Exeter, UK.

    • Andrew R Wood
    • , Dorota Pasko
    • , Michael N Weedon
    •  & Timothy M Frayling

  2. Estonian Genome Center, University of Tartu, Tartu, Estonia.

    • Tonu Esko
    • , Reedik Mägi
    • , Evelin Mihailov
    • , Krista Fischer
    • , Toomas Haller
    • , Lili Milani
    • , Andrew P Morris
    • , Mari Nelis
    • , Markus Perola
    • , Eva Reinmaa
    •  & Andres Metspalu

  3. Division of Endocrinology, Genetics and Basic and Translational Obesity Research, Boston Children's Hospital, Boston, Massachusetts, USA.

    • Tonu Esko
    • , Sailaja Vedantam
    • , Tune H Pers
    • , Cameron D Palmer
    •  & Joel N Hirschhorn

  4. Broad Institute of MIT and Harvard University, Cambridge, Massachusetts, USA.

    • Tonu Esko
    • , Sailaja Vedantam
    • , Tune H Pers
    • , Karol Estrada
    • , Cameron D Palmer
    • , Cecilia M Lindgren
    • , Alkes L Price
    •  & Joel N Hirschhorn

  5. Department of Genetics, Harvard Medical School, Boston, Massachusetts, USA.

    • Tonu Esko
    • , Tune H Pers
    •  & Joel N Hirschhorn

  6. Queensland Brain Institute, University of Queensland, Brisbane, Queensland, Australia.

    • Jian Yang
    • , Anna A E Vinkhuyzen
    • , Gibran Hemani
    • , Joseph E Powell
    •  & Peter M Visscher

  7. University of Queensland Diamantina Institute, Translation Research Institute, Brisbane, Queensland, Australia.

    • Jian Yang
    • , Gibran Hemani
    • , Joseph E Powell
    •  & Peter M Visscher

  8. Center for Biological Sequence Analysis, Department of Systems Biology, Technical University of Denmark, Lyngby, Denmark.

    • Tune H Pers

  9. Science for Life Laboratory, Uppsala University, Uppsala, Sweden.

    • Stefan Gustafsson
    • , Tove Fall
    • , Johan Ärnlöv
    • , Ann-Christine Syvänen
    •  & Erik Ingelsson

  10. Department of Medical Sciences, Molecular Epidemiology, Uppsala University, Uppsala, Sweden.

    • Stefan Gustafsson
    • , Tove Fall
    • , Johan Ärnlöv
    •  & Erik Ingelsson

  11. Division of Preventive Medicine, Brigham and Women's Hospital, Boston, Massachusetts, USA.

    • Audrey Y Chu
    • , Lynda M Rose
    • , Paul M Ridker
    •  & Daniel I Chasman

  12. Department of Internal Medicine, Erasmus Medical Center, Rotterdam, the Netherlands.

    • Karol Estrada
    • , Carolina Medina-Gomez
    • , Suzanne van Dijk
    • , Nathalie van der Velde
    • , Fernando Rivadeneira
    •  & André G Uitterlinden

  13. Analytic and Translational Genetics Unit, Massachusetts General Hospital and Harvard Medical School, Boston, Massachusetts, USA.

    • Karol Estrada
    •  & Stephan Ripke

  14. Medical Research Council (MRC) Epidemiology Unit, University of Cambridge, Institute of Metabolic Science, Addenbrooke's Hospital, Cambridge, UK.

    • Jian'an Luan
    • , Felix R Day
    • , Jing Hua Zhao
    • , Robert A Scott
    • , Nita G Forouhi
    • , Claudia Langenberg
    • , Ken K Ong
    • , Nicholas J Wareham
    •  & Ruth J F Loos

  15. Institute of Social and Preventive Medicine (IUMSP), Centre Hospitalier Universitaire Vaudois (CHUV), Lausanne, Switzerland.

    • Zoltán Kutalik

  16. Swiss Institute of Bioinformatics, Lausanne, Switzerland.

    • Zoltán Kutalik
    • , Sven Bergmann
    •  & Jacques S Beckmann

  17. Department of Medical Genetics, University of Lausanne, Lausanne, Switzerland.

    • Zoltán Kutalik
    • , Sven Bergmann
    •  & Jacques S Beckmann

  18. Genetic Epidemiology Unit, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, the Netherlands.

    • Najaf Amin
    • , Ayse Demirkan
    • , Aaron Isaacs
    • , Ben A Oostra
    •  & Cornelia M van Duijn

  19. Department of Genetics, University of North Carolina, Chapel Hill, North Carolina, USA.

    • Martin L Buchkovich
    • , Damien C Croteau-Chonka
    •  & Karen L Mohlke

  20. Channing Division of Network Medicine, Department of Medicine, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts, USA.

    • Damien C Croteau-Chonka
    • , Lu Qi
    •  & David J Hunter

  21. Department of Genetics, Division of Statistical Genomics, Washington University School of Medicine, St. Louis, Missouri, USA.

    • Yanan Duan

  22. Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden.

    • Tove Fall
    • , Henrik Grönberg
    • , Per Hall
    • , Patrik K E Magnusson
    •  & Nancy L Pedersen

  23. Department of Genetics, University Medical Center Groningen, University of Groningen, Groningen, the Netherlands.

    • Rudolf Fehrmann
    • , Juha Karjalainen
    • , Harm-Jan Westra
    • , Joanna Smolonska
    • , Morris A Swertz
    • , Pim van der Harst
    •  & Lude Franke

  24. Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, UK.

    • Teresa Ferreira
    • , Reedik Mägi
    • , Joshua C Randall
    • , Anuj Goel
    • , Inga Prokopenko
    • , Thorhildur Juliusdottir
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    • , Andrew P Morris
    • , Hugh Watkins
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    • , Mark I McCarthy
    •  & Erik Ingelsson

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

    • Anne U Jackson
    • , Adam E Locke
    • , Heather M Stringham
    • , Michael Boehnke
    •  & Goncalo R Abecasis

  26. Montreal Heart Institute, Montreal, Quebec, Canada.

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  27. Institute of Molecular and Cell Biology, University of Tartu, Tartu, Estonia.

    • Evelin Mihailov
    •  & Andres Metspalu

  28. Istituto di Ricerca Genetica e Biomedica (IRGB), Consiglio Nazionale delle Ricerche, Cagliari, Italy.

    • Eleonora Porcu
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  29. Wellcome Trust Sanger Institute, Hinxton, UK.

    • Joshua C Randall
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    • , Kathleen Stirrups
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  30. Institute for Medical Informatics, Biometry and Epidemiology (IMIBE), University Hospital Essen, Essen, Germany.

    • André Scherag
    • , Sonali Pechlivanis
    •  & Susanne Moebus

  31. Clinical Epidemiology, Integrated Research and Treatment Center, Center for Sepsis Control and Care (CSCC), Jena University Hospital, Jena, Germany.

    • André Scherag

  32. Department of Genetic Epidemiology, Institute of Epidemiology and Preventive Medicine, University of Regensburg, Regensburg, Germany.

    • Thomas W Winkler
    •  & Iris M Heid

  33. Department of Nutrition, Harvard School of Public Health, Harvard University, Boston, Massachusetts, USA.

    • Tsegaselassie Workalemahu
    • , Lu Qi
    • , Paul W Franks
    •  & David J Hunter

  34. HudsonAlpha Institute for Biotechnology, Huntsville, Alabama, USA.

    • Devin Absher
    •  & Lindsay L Waite

  35. Institute of Genetic Epidemiology, Helmholtz Zentrum München–German Research Center for Environmental Health, Neuherberg, Germany.

    • Eva Albrecht
    • , Janina S Ried
    • , Harald Grallert
    • , Martina Müller-Nurasyid
    • , Christian Gieger
    • , Konstantin Strauch
    •  & Iris M Heid

  36. Telethon Institute for Child Health Research, Centre for Child Health Research, University of Western Australia, Perth, Western Australia, Australia.

    • Denise Anderson
    •  & Matthew Cooper

  37. Section on Growth and Development, Program in Developmental Endocrinology and Genetics, Eunice Kennedy Shriver National Institute of Child Health and Human Development, US National Institutes of Health, Bethesda, Maryland, USA.

    • Jeffrey Baron
    •  & Julian C Lui

  38. Netherlands Consortium for Healthy Aging (NCHA), Leiden University Medical Center, Leiden, the Netherlands.

    • Marian Beekman
    • , Joris Deelen
    • , Quinta Helmer
    • , Jeanine J Houwing-Duistermaat
    • , Diana van Heemst
    •  & P Eline Slagboom

  39. Department of Molecular Epidemiology, Leiden University Medical Center, Leiden, the Netherlands.

    • Marian Beekman
    • , Joris Deelen
    •  & P Eline Slagboom

  40. Department of Human Genetics, Leiden University Medical Center, Leiden, the Netherlands.

    • Ayse Demirkan

  41. Center for Complex Disease Genomics, McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University School of Medicine, Baltimore, Maryland, USA.

    • Georg B Ehret
    •  & Aravinda Chakravarti

  42. Department of Specialties of Internal Medicine, Division of Cardiology, Geneva University Hospital, Geneva, Switzerland.

    • Georg B Ehret

  43. Department of Epidemiology Research, Statens Serum Institut, Copenhagen, Denmark.

    • Bjarke Feenstra
    • , Heather A Boyd
    • , Frank Geller
    •  & Mads Melbye

  44. Department of Genetics, Washington University School of Medicine, St. Louis, Missouri, USA.

    • Mary F Feitosa
    • , E Warwick Daw
    • , Mary K Wojczynski
    • , Qunyuan Zhang
    • , Aldi T Kraja
    • , D C Rao
    •  & Ingrid B Borecki

  45. Centre for Population Health Sciences, University of Edinburgh, Edinburgh, UK.

    • Ross M Fraser
    • , Jennifer L Bolton
    • , Stela McLachlan
    • , Harry Campbell
    • , Jackie F Price
    • , Igor Rudan
    •  & James F Wilson

  46. Radcliffe Department of Medicine, Division of Cardiovascular Medicine, University of Oxford, Oxford, UK.

    • Anuj Goel
    • , Martin Farrall
    •  & Hugh Watkins

  47. Division of Public Health Sciences, Fred Hutchinson Cancer Research Center, Seattle, Washington, USA.

    • Jian Gong
    • , Jeffrey Haessler
    • , Charles Kooperberg
    •  & Ulrike Peters

  48. Department of Epidemiology, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, USA.

    • Anne E Justice
    • , Keri L Monda
    •  & Kari E North

  49. William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London, UK.

    • Stavroula Kanoni
    • , Kathleen Stirrups
    •  & Panos Deloukas

  50. Vth Department of Medicine (Nephrology, Hypertensiology, Endocrinology, Diabetology, Rheumatology), Medical Faculty of Mannheim, University of Heidelberg, Heidelberg, Germany.

    • Marcus E Kleber
    • , Graciela Delgado
    • , Tanja B Grammer
    •  & Winfried März

  51. Department of Internal Medicine II, Ulm University Medical Centre, Ulm, Germany.

    • Marcus E Kleber

  52. National Institute for Health and Welfare, Helsinki, Finland.

    • Kati Kristiansson
    • , Niina Eklund
    • , Leena Kinnunen
    • , Jaana Lindström
    • , Johan G Eriksson
    • , Pekka Jousilahti
    • , Antti M Jula
    • , Jaakko Kaprio
    • , Seppo Koskinen
    • , Satu Männistö
    • , Veikko Salomaa
    • , Jarmo Virtamo
    • , Markus Perola
    •  & Jaakko Tuomilehto

  53. Epidemiology Program, University of Hawaii Cancer Center, Honolulu, Hawaii, USA.

    • Unhee Lim
    • , Lynne R Wilkens
    •  & Loic Le Marchand

  54. Charles Bronfman Institute for Personalized Medicine, Icahn School of Medicine at Mount Sinai, New York, New York, USA.

    • Vaneet Lotay
    • , Janina Jeff
    • , Yingchang Lu
    • , Omri Gottesman
    • , Erwin P Bottinger
    •  & Ruth J F Loos

  55. Department of Twin Research and Genetic Epidemiology, King's College London, London, UK.

    • Massimo Mangino
    • , Cristina Menni
    • , Alireza Moayyeri
    •  & Tim D Spector

  56. Department of Cardiology, University Medical Center Groningen, University of Groningen, Groningen, the Netherlands.

    • Irene Mateo Leach
    • , Hans L Hillege
    • , Niek Verweij
    •  & Pim van der Harst

  57. Netherlands Consortium for Healthy Aging (NCHA), Rotterdam, the Netherlands.

    • Carolina Medina-Gomez
    • , Albert Hofman
    • , Manfred Kayser
    • , Fernando Rivadeneira
    • , André G Uitterlinden
    •  & Cornelia M van Duijn

  58. Department of Epidemiology, Erasmus Medical Center, Rotterdam, the Netherlands.

    • Carolina Medina-Gomez
    • , Albert Hofman
    • , Fernando Rivadeneira
    • , André G Uitterlinden
    •  & Cornelia M van Duijn

  59. Laboratory of Neurogenetics, National Institute on Aging, US National Institutes of Health, Bethesda, Maryland, USA.

    • Michael A Nalls

  60. QIMR Berghofer Medical Research Institute, Brisbane, Queensland, Australia.

    • Dale R Nyholt
    • , Anjali K Henders
    • , Grant W Montgomery
    •  & Nicholas G Martin

  61. Oxford Centre for Diabetes, Endocrinology and Metabolism, University of Oxford, Oxford, UK.

    • Inga Prokopenko
    • , Amy Barrett
    • , Amanda J Bennett
    • , Christopher J Groves
    • , Nigel W Rayner
    • , Neil R Robertson
    •  & Mark I McCarthy

  62. Department of Genomics of Common Disease, School of Public Health, Imperial College London, Hammersmith Hospital, London, UK.

    • Inga Prokopenko
    •  & Philippe Froguel

  63. Stanley Center for Psychiatric Research, Broad Institute of MIT and Harvard University, Cambridge, Massachusetts, USA.

    • Stephan Ripke

  64. Department of Clinical Sciences, Genetic and Molecular Epidemiology Unit, Lund University Diabetes Center, Skåne University Hospital, Malmö, Sweden.

    • Dmitry Shungin
    • , Frida Renstrom
    •  & Paul W Franks

  65. Department of Public Health and Clinical Medicine, Unit of Medicine, Umeå University, Umeå, Sweden.

    • Dmitry Shungin
    •  & Paul W Franks

  66. Department of Odontology, Umeå University, Umeå, Sweden.

    • Dmitry Shungin

  67. Department of Medicine, University of Eastern Finland, Kuopio, Finland.

    • Alena Stancáková

  68. Atherosclerosis Research Unit, Center for Molecular Medicine, Department of Medicine, Karolinska Institutet, Stockholm, Sweden.

    • Rona J Strawbridge
    • , Lasse Folkersen
    •  & Anders Hamsten

  69. Division of Biostatistics, Washington University School of Medicine, St. Louis, Missouri, USA.

    • Yun Ju Sung
    • , D C Rao
    •  & Treva K Rice

  70. Translational Gerontology Branch, National Institute on Aging, Baltimore, Maryland, USA.

    • Toshiko Tanaka
    •  & Luigi Ferrucci

  71. Interfaculty Institute for Genetics and Functional Genomics, University Medicine Greifswald, Greifswald, Germany.

    • Alexander Teumer

  72. Department of Cardiology, Leiden University Medical Center, Leiden, the Netherlands.

    • Stella Trompet
    •  & J Wouter Jukema

  73. Department of Gerontology and Geriatrics, Leiden University Medical Center, Leiden, the Netherlands.

    • Stella Trompet
    •  & Diana van Heemst

  74. Experimental Cardiology Laboratory, Division of Heart and Lungs, University Medical Center Utrecht, Utrecht, the Netherlands.

    • Sander W van der Laan
    • , Hester M den Ruijter
    •  & Gerard Pasterkamp

  75. Department of Medical Genetics, University Medical Center Utrecht, Utrecht, the Netherlands.

    • Jessica van Setten
    •  & Paul I W de Bakker

  76. Department of Endocrinology, University of Groningen, University Medical Center Groningen, Groningen, the Netherlands.

    • Jana V Van Vliet-Ostaptchouk

  77. DZHK (Deutsches Zentrum für Herz-Kreislaufforschung–German Centre for Cardiovascular Research), partner site Greifswald, Greifswald, Germany.

    • Zhaoming Wang
    • , Marcus Dörr
    • , Wolfgang Hoffmann
    • , Matthias Nauck
    •  & Henry Völzke

  78. Institute of Clinical Chemistry and Laboratory Medicine, University Medicine Greifswald, Greifswald, Germany.

    • Zhaoming Wang
    • , Anke Hannemann
    •  & Matthias Nauck

  79. Division of Cancer Epidemiology and Genetics, National Cancer Institute, US National Institutes of Health, Bethesda, Maryland, USA.

    • Zhaoming Wang
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    • , Stephen J Chanock
    •  & Sonja I Berndt

  80. Core Genotyping Facility, SAIC-Frederick, Inc., NCI-Frederick, Frederick, Maryland, USA.

    • Zhaoming Wang
    •  & Kevin B Jacobs

  81. CNRS UMR 8199, Lille, France.

    • Loïc Yengo
    • , Elodie Eury
    • , Stéphane Lobbens
    • , Stéphane Cauchi
    •  & Philippe Froguel

  82. European Genomic Institute for Diabetes, Lille, France.

    • Loïc Yengo
    • , Elodie Eury
    • , Stéphane Lobbens
    • , Stéphane Cauchi
    •  & Philippe Froguel

  83. Université de Lille 2, Lille, France.

    • Loïc Yengo
    • , Elodie Eury
    • , Stéphane Lobbens
    • , Stéphane Cauchi
    •  & Philippe Froguel

  84. Ealing Hospital National Health Service (NHS) Trust, Middlesex, UK.

    • Weihua Zhang
    • , Uzma Afzal
    • , Laticia Oozageer
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    •  & John C Chambers

  85. Department of Epidemiology and Biostatistics, Imperial College London, London, UK.

    • Weihua Zhang
    • , Uzma Afzal
    • , Joban Sehmi
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    •  & John C Chambers

  86. School of Health and Social Studies, Dalarna University, Falun, Sweden.

    • Johan Ärnlöv

  87. PathWest Laboratory Medicine of Western Australia, Nedlands, Western Australia, Australia.

    • Gillian M Arscott
    • , John Beilby
    •  & Jennie Hui

  88. Geriatric Unit, Azienda Sanitaria Firenze (ASF), Florence, Italy.

    • Stefania Bandinelli

  89. Department of Genetics, Texas Biomedical Research Institute, San Antonio, Texas, USA.

    • Claire Bellis
    •  & John Blangero

  90. Department of Medical Sciences, Endocrinology, Diabetes and Metabolism, Uppsala University, Uppsala, Sweden.

    • Christian Berne

  91. Integrated Research and Treatment Center (IFB) Adiposity Diseases, University of Leipzig, Leipzig, Germany.

    • Matthias Blüher
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  92. Department of Medicine, University of Leipzig, Leipzig, Germany.

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  93. LifeLines, University Medical Center Groningen, University of Groningen, Groningen, the Netherlands.

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  94. Department of Pharmacology and Therapeutics, University College Cork, Cork, Ireland.

    • Brendan M Buckley

  95. Department of Statistics and Biostatistics, Rutgers University, Piscataway, New Jersy, USA.

    • Steven Buyske

  96. Department of Genetics, Rutgers University, Piscataway, New Jersey, USA.

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  97. Department of Biology, Norwegian University of Science and Technology, Trondheim, Norway.

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  98. Genome Technology Branch, National Human Genome Research Institute, US National Institutes of Health, Bethesda, Maryland, USA.

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    • Robert Clarke
    •  & Rory Collins

  100. Division of Endocrinology, Diabetes and Metabolism, Ulm University Medical Centre, Ulm, Germany.

    • Simone Claudi-Boehm
    • , Sigrun Merger
    •  & Bernhard O Boehm

  101. Department of Radiology, University Medical Center Utrecht, Utrecht, the Netherlands.

    • Pim A De Jong

  102. Department of Biomedical Informatics, Vanderbilt University, Nashville, Tennessee, USA.

    • Josh C Denny

  103. Department of Human Nutrition, Wageningen University, Wageningen, the Netherlands.

    • Rosalie Dhonukshe-Rutten
    •  & Lisette C P G M de Groot

  104. Department of Dietetics–Nutrition, Harokopio University, Athens, Greece.

    • Maria Dimitriou
    •  & George Dedoussis

  105. Medical Research Institute, University of Dundee, Ninewells Hospital and Medical School, Dundee, UK.

    • Alex S F Doney
    • , Andrew D Morris
    •  & Colin N A Palmer

  106. Department of Internal Medicine B, University Medicine Greifswald, Greifswald, Germany.

    • Marcus Dörr

  107. Institute for Molecular Medicine, University of Helsinki, Helsinki, Finland.

    • Niina Eklund
    • , Jaakko Kaprio
    • , Leif C Groop
    •  & Markus Perola

  108. Laboratory of Epidemiology and Population Sciences, National Institute on Aging, US National Institutes of Health, Bethesda, Maryland, USA.

    • Melissa E Garcia
    •  & Tamara B Harris

  109. Department of Public Health and Caring Sciences, Geriatrics, Uppsala University, Uppsala, Sweden.

    • Vilmantas Giedraitis

  110. Kaiser Permanente, Division of Research, Oakland, California, USA.

    • Alan S Go

  111. Research Unit of Molecular Epidemiology, Helmholtz Zentrum München–German Research Center for Environmental Health, Neuherberg, Germany.

    • Harald Grallert
    • , Thomas Illig
    • , Norman Klopp
    • , Melanie Waldenberger
    •  & Annette Peters

  112. German Center for Diabetes Research (DZD), Neuherberg, Germany.

    • Harald Grallert

  113. Department of Medicine III, University Hospital Carl Gustav Carus, Technische Universität Dresden, Dresden, Germany.

    • Jürgen Gräßler
    • , Stefan R Bornstein
    •  & Peter E H Schwarz

  114. Unit of Nutritional Research, Department of Public Health and Clinical Medicine, Umeå University, Umeå, Sweden.

    • Goran Hallmans

  115. Department of Psychiatry, University of Groningen, University Medical Center Groningen, Groningen, the Netherlands.

    • Catharina A Hartman
    •  & Albertine J Oldehinkel

  116. Kuopio Research Institute of Exercise Medicine, Kuopio, Finland.

    • Maija Hassinen
    • , Timo A Lakka
    •  & Rainer Rauramaa

  117. MRC Human Genetics Unit, Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, UK.

    • Caroline Hayward
    •  & Alan F Wright

  118. National Heart, Lung, and Blood Institute, Framingham Heart Study, Framingham, Massachusetts, USA.

    • Nancy L Heard-Costa
    • , L Adrienne Cupples
    •  & Caroline S Fox

  119. Department of Neurology, Boston University School of Medicine, Boston, Massachusetts, USA.

    • Nancy L Heard-Costa

  120. Department of Medical Statistics and Bioinformatics, Leiden University Medical Center, Leiden, the Netherlands.

    • Quinta Helmer
    •  & Jeanine J Houwing-Duistermaat

  121. Faculty of Psychology and Education, VU University Amsterdam, Amsterdam, the Netherlands.

    • Quinta Helmer

  122. Department of Epidemiology, University Medical Center Groningen, University of Groningen, Groningen, the Netherlands.

    • Hans L Hillege
    • , Ilja M Nolte
    • , Salome Scholtens
    • , Joanna Smolonska
    • , Judith M Vonk
    • , Ronald P Stolk
    •  & Harold Snieder

  123. Department of Medicine, Stanford University School of Medicine, Stanford, California, USA.

    • Mark A Hlatky
    • , Themistocles L Assimes
    • , Mads Melbye
    •  & Thomas Quertermous

  124. Institute for Community Medicine, University Medicine Greifswald, Greifswald, Germany.

    • Wolfgang Hoffmann
    •  & Henry Völzke

  125. Department of Biomedicine, Division of Medical Genetics, University of Basel, Basel, Switzerland.

    • Per Hoffmann

  126. Department of Genomics, Life and Brain Center, University of Bonn, Bonn, Germany.

    • Per Hoffmann
    •  & Markus M Nöthen

  127. Institute of Human Genetics, University of Bonn, Bonn, Germany.

    • Per Hoffmann
    •  & Markus M Nöthen

  128. Department of Public Health and General Practice, Norwegian University of Science and Technology, Trondheim, Norway.

    • Oddgeir Holmen
    •  & Kristian Hveem

  129. Hannover Unified Biobank, Hannover Medical School, Hannover, Germany.

    • Thomas Illig
    •  & Norman Klopp

  130. Center for Medical Systems Biology, Leiden, the Netherlands.

    • Aaron Isaacs
    • , Ben A Oostra
    •  & Cornelia M van Duijn

  131. Department of Pulmonary Physiology and Sleep Medicine, Nedlands, Western Australia, Australia.

    • Alan L James

  132. School of Medicine and Pharmacology, University of Western Australia, Crawley, Western Australia, Australia.

    • Alan L James

  133. Department of Immunology, Genetics and Pathology, SciLifeLab, Rudbeck Laboratory, Uppsala University, Uppsala, Sweden.

    • Åsa Johansson
    •  & Ulf Gyllensten

  134. Department of Haematology, University of Cambridge, Cambridge, UK.

    • Jennifer Jolley
    •  & Willem H Ouwehand

  135. NHS Blood and Transplant, Cambridge, UK.

    • Jennifer Jolley
    •  & Willem H Ouwehand

  136. Department of Medicine, University of Oulu, Oulu, Finland.

    • Juhani Junttila

  137. Department of Medicine, Northwestern University Feinberg School of Medicine, Chicago, Illinois, USA.

    • Abel N Kho
    •  & M Geoffrey Hayes

  138. Unit of Periodontology, Department of Restorative Dentistry, Periodontology and Endodontology, University Medicine Greifswald, Greifswald, Germany.

    • Thomas Kocher

  139. Department of Internal Medicine I, Ulm University Medical Centre, Ulm, Germany.

    • Wolfgang Kratzer
    •  & Thomas Seufferlein

  140. Institute of Human Genetics, Helmholtz Zentrum München–German Research Center for Environmental Health, Neuherberg, Germany.

    • Peter Lichtner

  141. Department of Medical Sciences, Cardiovascular Epidemiology, Uppsala University, Uppsala, Sweden.

    • Lars Lind
    •  & Johan Sundström

  142. Centre for Bone and Arthritis Research, Department of Internal Medicine and Clinical Nutrition, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, Gothenburg, Sweden.

    • Mattias Lorentzon
    •  & Claes Ohlsson

  143. Genetics of Obesity and Related Metabolic Traits Program, Icahn School of Medicine at Mount Sinai, New York, New York, USA.

    • Yingchang Lu
    •  & Ruth J F Loos

  144. Steno Diabetes Center A/S, Gentofte Denmark.

    • Valeriya Lyssenko

  145. Service of Nephrology, Department of Medicine, Lausanne University Hospital (CHUV), Lausanne, Switzerland.

    • Marc Maillard

  146. School of Social and Community Medicine, University of Bristol, Bristol, UK.

    • Wendy L McArdle

  147. Tropical Metabolism Research Unit, Tropical Medicine Research Institute, University of the West Indies, Mona, Kingston, Jamaica.

    • Colin A McKenzie
    •  & Terrence Forrester

  148. Global Health Institute, Department of Life Sciences, Ecole Polytechnique Fédérale de Lausanne, Lausanne, Switzerland.

    • Paul J McLaren

  149. Institute of Microbiology, University Hospital and University of Lausanne, Lausanne, Switzerland.

    • Paul J McLaren

  150. Center for Observational Research, Amgen, Inc., Thousand Oaks, California, USA.

    • Keri L Monda

  151. Center for Evidence-Based Healthcare, University Hospital Carl Gustav Carus, Technische Universität Dresden, Dresden, Germany.

    • Gabriele Müller

  152. Department of Medicine I, University Hospital Großhadern, Ludwig Maximilians Universität, Munich, Germany.

    • Martina Müller-Nurasyid

  153. Chair of Genetic Epidemiology, Institute of Medical Informatics, Biometry and Epidemiology, Ludwig Maximilians Universität, Neuherberg, Germany.

    • Martina Müller-Nurasyid
    •  & Konstantin Strauch

  154. DZHK (Deutsches Forschungszentrum für Herz-Kreislauferkrankungen–German Research Centre for Cardiovascular Research), Munich Heart Alliance, Munich, Germany.

    • Martina Müller-Nurasyid
    • , Heribert Schunkert
    • , Christian Hengstenberg
    • , Thomas Meitinger
    •  & Annette Peters

  155. Department of Respiratory Medicine, Sir Charles Gairdner Hospital, Nedlands, Western Australia, Australia.

    • Arthur W Musk

  156. Department of Epidemiology and Biostatistics, Institute for Research in Extramural Medicine (EMGO) Institute for Health and Care Research, VU University Medical Center, Amsterdam, the Netherlands.

    • Stefan Pilz
    •  & Natasja M van Schoor

  157. Department of Internal Medicine, Division of Endocrinology and Metabolism, Medical University of Graz, Graz, Austria.

    • Stefan Pilz

  158. Diabetology-Endocrinology-Nutrition, Public Hospital System of the City of Paris (AP-HP), Bichat Hospital, Paris, France.

    • Ronan Roussel

  159. INSERM U872, Centre de Recherche des Cordeliers, Paris, France.

    • Ronan Roussel

  160. Paris Diderot University, Paris, France.

    • Ronan Roussel

  161. Clinical Institute of Medical and Chemical Laboratory Diagnostics, Medical University of Graz, Graz, Austria.

    • Hubert Scharnagl
    •  & Winfried März

  162. Department of Preventive Medicine, Keck School of Medicine, University of Southern California, Los Angeles, California, USA.

    • Fredrick R Schumacher
    •  & Christopher A Haiman

  163. Deutsches Herzzentrum München, Technische Universität München, Munich, Germany.

    • Heribert Schunkert
    •  & Christian Hengstenberg

  164. National Cancer Institute, Bethesda, Maryland, USA.

    • Jianxin Shi

  165. Department of Sociology, University of Helsinki, Helsinki, Finland.

    • Karri Silventoinen

  166. EMGO Institute for Health and Care Research, VU University, Amsterdam, the Netherlands.

    • Johannes H Smit

  167. Department of Psychiatry, Neuroscience Campus, VU University Amsterdam, Amsterdam, the Netherlands.

    • Johannes H Smit

  168. Icelandic Heart Association, Kopavogur, Iceland.

    • Albert Vernon Smith
    •  & Vilmundur Gudnason

  169. University of Iceland, Reykjavik, Iceland.

    • Albert Vernon Smith
    •  & Vilmundur Gudnason

  170. Molecular and Cellular Therapeutics, Royal College of Surgeons in Ireland, Dublin, Ireland.

    • Alice V Stanton

  171. Institute of Cardiovascular and Medical Sciences, Faculty of Medicine, University of Glasgow, Glasgow, UK.

    • David J Stott

  172. Department of Medical Sciences, Molecular Medicine, Uppsala University, Uppsala, Sweden.

    • Ann-Christine Syvänen

  173. Department of Public Health Sciences, Stritch School of Medicine, Loyola University of Chicago, Maywood, Illinois, USA.

    • Bamidele O Tayo
    •  & Richard S Cooper

  174. deCODE Genetics, Amgen, Inc., Reykjavik, Iceland.

    • Gudmar Thorleifsson
    • , Valgerdur Steinthorsdottir
    • , Unnur Thorsteinsdottir
    •  & Kari Stefansson

  175. Department of Oncology, University of Cambridge, Cambridge, UK.

    • Jonathan P Tyrer

  176. Section of Geriatrics, Department of Internal Medicine, Academic Medical Center, Amsterdam, the Netherlands.

    • Nathalie van der Velde

  177. Department of Child and Adolescent Psychiatry, Psychology, Erasmus University Medical Center, Rotterdam, the Netherlands.

    • Floor V A van Oort

  178. Department for Health Evidence, Radboud University Medical Centre, Nijmegen, the Netherlands.

    • Sita H Vermeulen
    •  & Lambertus A Kiemeney

  179. Department of Genetics, Radboud University Medical Centre, Nijmegen, the Netherlands.

    • Sita H Vermeulen

  180. Department of Clinical Chemistry, Ulm University Medical Centre, Ulm, Germany.

    • Roman Wennauer

  181. DZHK (Deutsches Forschungszentrum für Herz-Kreislauferkrankungen–German Research Centre for Cardiovascular Research), partner site Hamburg-Lubeck-Kiel, Lubeck, Germany.

    • Christina Willenborg
    •  & Jeanette Erdmann

  182. Institut für Integrative und Experimentelle Genomik, Universität zu Lübeck, Lübeck, Germany.

    • Christina Willenborg
    •  & Jeanette Erdmann

  183. Department of Community Medicine, Faculty of Health Sciences, UiT The Arctic University of Tromsø, Tromsø, Norway.

    • Tom Wilsgaard
    •  & Inger Njølstad

  184. MRC Unit for Lifelong Health and Ageing at University College London, London, UK.

    • Andrew Wong
    • , Ken K Ong
    •  & Diana Kuh

  185. Department of Epidemiology and Public Health, University of Strasbourg, Faculty of Medicine, Strasbourg, France.

    • Dominique Arveiler

  186. Department of Internal Medicine, University Medical Center Groningen, University of Groningen, Groningen, the Netherlands.

    • Stephan J L Bakker
    •  & Ron T Gansevoort

  187. Pathology and Laboratory Medicine, University of Western Australia, Perth, Western Australia, Australia.

    • John Beilby
    •  & Jennie Hui

  188. Cedars-Sinai Diabetes and Obesity Research Institute, Los Angeles, California, USA.

    • Richard N Bergman

  189. Department of Prosthetic Dentistry, Gerostomatology and Dental Materials, University Medicine Greifswald, Greifswald, Germany.

    • Reiner Biffar

  190. Biological Psychology, VU University Amsterdam, Amsterdam, the Netherlands.

    • Dorret I Boomsma

  191. Institute of Social and Preventive Medicine (IUMSP), Centre Hospitalier Universitaire Vaudois (CHUV) and University of Lausanne, Lausanne, Switzerland.

    • Pascal Bovet
    •  & Murielle Bochud

  192. Ministry of Health, Victoria, Republic of Seychelles.

    • Pascal Bovet
    •  & Murielle Bochud

  193. Laboratory Medicine, Hospital of Desio, Department of Health Sciences, University of Milano, Bicocca, Italy.

    • Paolo Brambilla

  194. Clinical Pharmacology Unit, University of Cambridge, Addenbrooke's Hospital, Cambridge, UK.

    • Morris J Brown

  195. Clinical Pharmacology, William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London, UK.

    • Mark J Caulfield
    •  & Patricia B Munroe

  196. Barts and The London Genome Centre, William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London, UK.

    • Mark J Caulfield
    •  & Patricia B Munroe

  197. Center for Human Genetics Research, Vanderbilt University Medical Center, Nashville, Tennessee, USA.

    • Dana C Crawford

  198. Department of Molecular Physiology and Biophysics, Vanderbilt University, Nashville, Tennessee, USA.

    • Dana C Crawford

  199. Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts, USA.

    • L Adrienne Cupples

  200. Department of Public Health and Primary Care, University of Cambridge, Cambridge, UK.

    • John Danesh
    •  & Danish Saleheen

  201. Division of Cardiovascular Epidemiology, Institute of Environmental Medicine, Karolinska Institutet, Stockholm, Sweden.

    • Ulf de Faire

  202. Julius Center for Health Sciences and Primary Care, University Medical Center Utrecht, Utrecht, the Netherlands.

    • Hester M den Ruijter

  203. Clinic of Cardiology, West German Heart Centre, University Hospital Essen, Essen, Germany.

    • Raimund Erbel

  204. Department of General Practice and Primary Health Care, University of Helsinki, Helsinki, Finland.

    • Johan G Eriksson

  205. Unit of General Practice, Helsinki University Central Hospital, Helsinki, Finland.

    • Johan G Eriksson

  206. Department of Internal Medicine, University of Pisa, Pisa, Italy.

    • Ele Ferrannini

  207. National Research Council (CNR) Institute of Clinical Physiology, University of Pisa, Pisa, Italy.

    • Ele Ferrannini

  208. Department of Cardiology, Toulouse University School of Medicine, Rangueil Hospital, Toulouse, France.

    • Jean Ferrières

  209. Robertson Center for Biostatistics, University of Glasgow, Glasgow, UK.

    • Ian Ford

  210. NorthShore University HealthSystem, University of Chicago, Evanston, Illinois, USA.

    • Pablo V Gejman

  211. Service of Therapeutic Education for Diabetes, Obesity and Chronic Diseases, Geneva University Hospital, Geneva, Switzerland.

    • Alain Golay

  212. Department of Medicine, Pharmacology, Pathology, Microbiology and Immunology, Vanderbilt University School of Medicine, Nashville, Tennessee, USA.

    • David W Haas

  213. Leeds MRC Medical Bioinformatics Centre, University of Leeds, Leeds, UK.

    • Alistair S Hall

  214. Institute of Biomedical and Clinical Science, University of Exeter, Exeter, UK.

    • Andrew T Hattersley

  215. Department of Psychiatry, Washington University School of Medicine, St. Louis, Missouri, USA.

    • Andrew C Heath
    • , Pamela A F Madden
    • , D C Rao
    •  & Treva K Rice

  216. Center for Biomedicine, European Academy Bozen, Bolzano (EURAC), Bolzano, Italy (affiliated institute of the University of Lübeck, Lübeck, Germany).

    • Andrew A Hicks
    •  & Peter P Pramstaller

  217. Division of Genomic Medicine, National Human Genome Research Institute, US National Institutes of Health, Bethesda, Maryland, USA.

    • Lucia A Hindorff

  218. Institute of Cardiovascular Science, University College London, London, UK.

    • Aroon D Hingorani
    •  & Folkert W Asselbergs

  219. Department of Vascular Medicine, Academic Medical Center, Amsterdam, the Netherlands.

    • G Kees Hovingh
    •  & John J P Kastelein

  220. Centre for Cardiovascular Genetics, Institute of Cardiovascular Sciences, University College London, London, UK.

    • Steve E Humphries

  221. Cardiovascular Genetics Division, Department of Internal Medicine, University of Utah, Salt Lake City, Utah, USA.

    • Steven C Hunt

  222. School of Population Health, University of South Australia, Adelaide, South Australia, Australia.

    • Elina Hypponen
    •  & Jennie Hui

  223. Sansom Institute for Health Research, University of South Australia, Adelaide, South Australia, Australia.

    • Elina Hypponen
    •  & Jennie Hui

  224. South Australian Health and Medical Research Institute, Adelaide, South Australia, Australia.

    • Elina Hypponen

  225. Centre for Paediatric Epidemiology and Biostatistics, University College London Institute of Child Health, London, UK.

    • Elina Hypponen
    •  & Chris Power

  226. National Institute for Health and Welfare, Oulu, Finland.

    • Marjo-Riitta Jarvelin

  227. MRC Health Protection Agency (HPA) Centre for Environment and Health, School of Public Health, Imperial College London, London, UK.

    • Marjo-Riitta Jarvelin

  228. Unit of Primary Care, Oulu University Hospital, Oulu, Finland.

    • Marjo-Riitta Jarvelin

  229. Biocenter Oulu, University of Oulu, Oulu, Finland.

    • Marjo-Riitta Jarvelin

  230. Institute of Health Sciences, University of Oulu, Oulu, Finland.

    • Marjo-Riitta Jarvelin
    •  & Sylvain Sebert

  231. Hjelt Institute Department of Public Health, University of Helsinki, Helsinki, Finland.

    • Jaakko Kaprio

  232. Department of Forensic Molecular Biology, Erasmus Medical Center, Rotterdam, the Netherlands.

    • Manfred Kayser

  233. UK Clinical Research Collaboration Centre of Excellence for Public Health (Northern Ireland), Queens University of Belfast, Belfast, UK.

    • Frank Kee

  234. Faculty of Medicine, Institute of Health Sciences, University of Oulu, Oulu, Finland.

    • Sirkka M Keinanen-Kiukaanniemi

  235. Unit of General Practice, Oulu University Hospital, Oulu, Finland.

    • Sirkka M Keinanen-Kiukaanniemi

  236. Department of Urology, Radboud University Medical Centre, Nijmegen, the Netherlands.

    • Lambertus A Kiemeney

  237. Imperial College Healthcare NHS Trust, London, UK.

    • Jaspal S Kooner
    •  & John C Chambers

  238. National Heart and Lung Institute, Imperial College London, London, UK.

    • Jaspal S Kooner

  239. Department of Epidemiology and Public Health, University College London, London, UK.

    • Meena Kumari
    • , Claudia Langenberg
    •  & Mika Kivimaki

  240. Department of Medicine, Kuopio University Hospital and University of Eastern Finland, Kuopio, Finland.

    • Johanna Kuusisto
    •  & Markku Laakso

  241. Department of Physiology, Institute of Biomedicine, University of Eastern Finland, Kuopio Campus, Kuopio, Finland.

    • Timo A Lakka

  242. Department of Clinical Physiology and Nuclear Medicine, Kuopio University Hospital and University of Eastern Finland, Kuopio, Finland.

    • Timo A Lakka
    •  & Rainer Rauramaa

  243. Department of Clinical Chemistry, Fimlab Laboratories and School of Medicine, University of Tampere, Tampere, Finland.

    • Terho Lehtimäki

  244. Department of Health Sciences, University of Milano, Milan, Italy.

    • Sara Lupoli

  245. Fondazione Filarete, Milan, Italy.

    • Sara Lupoli

  246. Division of Nephrology and Dialysis, San Raffaele Scientific Institute, Milan, Italy.

    • Paolo Manunta

  247. Università Vita–Salute San Raffaele, Milan, Italy.

    • Paolo Manunta

  248. Institut Universitaire de Cardiologie et de Pneumologie de Québec, Faculty of Medicine, Laval University, Quebec City, Quebec, Canada.

    • André Marette

  249. Institute of Nutrition and Functional Foods, Laval University, Quebec City, Quebec, Canada.

    • André Marette
    • , Marie-Claude Vohl
    •  & Louis Pérusse

  250. Department of Biostatistics, University of Washington, Seattle, Washington, USA.

    • Barbara McKnight

  251. Department of Surgery, University Medical Center Utrecht, Utrecht, the Netherlands.

    • Frans L Moll

  252. Department of Biostatistics, University of Liverpool, Liverpool, UK.

    • Andrew P Morris

  253. Department of Pediatrics, University of Iowa, Iowa City, Iowa, USA.

    • Jeffrey C Murray

  254. Institute of Epidemiology II, Helmholtz Zentrum München–German Research Center for Environmental Health, Neuherberg, Germany.

    • Annette Peters

  255. Department of Neurology, General Central Hospital, Bolzano, Italy.

    • Peter P Pramstaller

  256. Department of Clinical Physiology and Nuclear Medicine, Turku University Hospital, Turku, Finland.

    • Olli T Raitakari

  257. Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku, Finland.

    • Olli T Raitakari

  258. Human Genomics Laboratory, Pennington Biomedical Research Center, Baton Rouge, Louisiana, USA.

    • Tuomo Rankinen
    • , Mark A Sarzynski
    •  & Claude Bouchard

  259. Center for Systems Genomics, Pennsylvania State University, University Park, Pennsylvania, USA.

    • Marylyn Ritchie

  260. Croatian Centre for Global Health, Faculty of Medicine, University of Split, Split, Croatia.

    • Igor Rudan

  261. Department of Cardiovascular Sciences, University of Leicester, Glenfield Hospital, Leicester, UK.

    • Nilesh J Samani

  262. National Institute for Health Research (NIHR) Leicester Cardiovascular Biomedical Research Unit, Glenfield Hospital, Leicester, UK.

    • Nilesh J Samani

  263. South Carelia Central Hospital, Lappeenranta, Finland.

    • Jouko Saramies

  264. Paul Langerhans Institute Dresden, German Center for Diabetes Research (DZD), Dresden, Germany.

    • Peter E H Schwarz

  265. International Centre for Circulatory Health, Imperial College London, London, UK.

    • Peter Sever

  266. Program for Personalized and Genomic Medicine, University of Maryland School of Medicine, Baltimore, Maryland, USA.

    • Alan R Shuldiner
    •  & Jeffrey R O'Connell

  267. Division of Endocrinology, Diabetes and Nutrition, University of Maryland School of Medicine, Baltimore, Maryland, USA.

    • Alan R Shuldiner
    •  & Jeffrey R O'Connell

  268. Geriatric Research and Education Clinical Center, Vetrans Administration Medical Center, Baltimore, Maryland, USA.

    • Alan R Shuldiner

  269. Helsinki University Central Hospital Heart and Lung Center, Department of Medicine, Helsinki University Central Hospital, Helsinki, Finland.

    • Juha Sinisalo

  270. Montreal Heart Institute, Université de Montréal, Montreal, Quebec, Canada.

    • Jean-Claude Tardif
    •  & Guillaume Lettre

  271. Department of Kinesiology, Laval University, Quebec City, Quebec, Canada.

    • Angelo Tremblay
    •  & Louis Pérusse

  272. Dipartimento di Scienze Farmacologiche e Biomolecolari, Università di Milano and Centro Cardiologico Monzino, Istituto di Ricovero e Cura a Carattere Scientifico (IRCCS), Milan, Italy.

    • Elena Tremoli

  273. Department of Food Science and Nutrition, Laval University, Quebec City, Quebec, Canada.

    • Marie-Claude Vohl

  274. Institut Pasteur de Lille, INSERM U744, Université de Lille 2, Lille, France.

    • Philippe Amouyel

  275. Department of Cardiology, Division of Heart and Lungs, University Medical Center Utrecht, Utrecht, the Netherlands.

    • Folkert W Asselbergs

  276. Durrer Center for Cardiogenetic Research, Interuniversity Cardiology Institute Netherlands–Netherlands Heart Institute, Utrecht, the Netherlands.

    • Folkert W Asselbergs
    • , J Wouter Jukema
    •  & Pim van der Harst

  277. Lee Kong Chian School of Medicine, Imperial College London and Nanyang Technological University, Singapore.

    • Bernhard O Boehm

  278. Health Science Center at Houston, University of Texas, Houston, Texas, USA.

    • Eric Boerwinkle

  279. Division of Genetics, Department of Medicine, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts, USA.

    • Paul I W de Bakker

  280. Department of Epidemiology, University Medical Center Utrecht, Utrecht, the Netherlands.

    • Paul I W de Bakker

  281. Lund University Diabetes Centre, Lund University, Malmö, Sweden.

    • Leif C Groop

  282. Diabetes and Endocrinology Unit, Department of Clinical Science, Lund University, Malmö, Sweden.

    • Leif C Groop

  283. Department of Epidemiology, Harvard School of Public Health, Harvard University, Boston, Massachusetts, USA.

    • David J Hunter
    •  & Alkes L Price

  284. Interuniversity Cardiology Institute of the Netherlands (ICIN), Utrecht, the Netherlands.

    • J Wouter Jukema

  285. Department of Epidemiology and Population Health, Albert Einstein College of Medicine. Belfer, New York, USA.

    • Robert C Kaplan

  286. Center for Human Genetics, Division of Public Health Sciences, Wake Forest School of Medicine, Winston-Salem, North Carolina, USA.

    • Yongmei Liu

  287. Synlab Academy, Synlab Services, Mannheim, Germany.

    • Winfried März

  288. Department of Clinical Genetics, Erasmus University Medical Center, Rotterdam, the Netherlands.

    • Ben A Oostra

  289. Harvard Medical School, Boston, Massachusetts, USA.

    • Paul M Ridker
    •  & Daniel I Chasman

  290. Institute for Translational Genomics and Population Sciences, Los Angeles BioMedical Research Institute at Harbor–University of California, Los Angeles Medical Center, Torrance, California, USA.

    • Jerome I Rotter

  291. Finnish Diabetes Association, Tampere, Finland.

    • Timo E Saaristo

  292. Pirkanmaa Hospital District, Tampere, Finland.

    • Timo E Saaristo

  293. Center for Non-Communicable Diseases, Karatchi, Pakistan.

    • Danish Saleheen

  294. Department of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, USA.

    • Danish Saleheen

  295. Laboratory of Genetics, National Institute on Aging, Baltimore, Maryland, USA.

    • David Schlessinger

  296. Instituto de Investigacion Sanitaria del Hospital Universario La Paz (IdiPAZ), Madrid, Spain.

    • Jaakko Tuomilehto

  297. Diabetes Research Group, King Abdulaziz University, Jeddah, Saudi Arabia.

    • Jaakko Tuomilehto

  298. Centre for Vascular Prevention, Danube University Krems, Krems, Austria.

    • Jaakko Tuomilehto

  299. Department of Public Health and Clinical Nutrition, University of Eastern Finland, Kuopio, Finland.

    • Matti Uusitupa

  300. Research Unit, Kuopio University Hospital, Kuopio, Finland.

    • Matti Uusitupa

  301. Institute of Cellular Medicine, Newcastle University, Newcastle, UK.

    • Mark Walker

  302. Chair of Epidemiology, Institute of Medical Informatics, Biometry and Epidemiology, Ludwig Maximilians Universität, Munich, Germany.

    • H-Erich Wichmann

  303. Klinikum Großhadern, Munich, Germany.

    • H-Erich Wichmann

  304. Institute of Epidemiology I, Helmholtz Zentrum München–German Research Center for Environmental Health, Neuherberg, Germany.

    • H-Erich Wichmann

  305. Department of Pulmonology, University Medical Center Utrecht, Utrecht, the Netherlands.

    • Pieter Zanen

  306. King Abdulaziz University, Jeddah, Saudi Arabia.

    • Panos Deloukas

  307. Department of Internal Medicine, Division of Gastroenterology, University of Michigan, Ann Arbor, Michigan, USA.

    • Elizabeth K Speliotes

  308. Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, Michigan, USA.

    • Elizabeth K Speliotes
    •  & Cristen J Willer

  309. Faculty of Medicine, University of Iceland, Reykjavik, Iceland.

    • Unnur Thorsteinsdottir
    •  & Kari Stefansson

  310. University of Cambridge Metabolic Research Laboratories, Institute of Metabolic Science, Addenbrooke's Hospital, Cambridge, UK.

    • Inês Barroso

  311. NIHR Cambridge Biomedical Research Centre, Institute of Metabolic Science, Addenbrooke's Hospital, Cambridge, UK.

    • Inês Barroso

  312. Carolina Center for Genome Sciences, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, USA.

    • Kari E North

  313. Division of Population Health Sciences and Education, St George's, University of London, London, UK.

    • David P Strachan

  314. Service of Medical Genetics, CHUV University Hospital, Lausanne, Switzerland.

    • Jacques S Beckmann

  315. Oxford NIHR Biomedical Research Centre, Oxford University Hospitals NHS Trust, Oxford, UK.

    • Mark I McCarthy

  316. Department of Internal Medicine, Division of Cardiovascular Medicine, University of Michigan, Ann Arbor, Michigan, USA.

    • Cristen J Willer

  317. Department of Human Genetics, University of Michigan, Ann Arbor, Michigan, USA.

    • Cristen J Willer

  318. Department of Biostatistics, Harvard School of Public Health, Boston, Massachusetts, USA.

    • Alkes L Price

  319. Mindich Child Health and Development Institute, Icahn School of Medicine at Mount Sinai, New York, New York, USA.

    • Ruth J F Loos

  320. Biosciences Research Division, Department of Primary Industries, Melbourne, Victoria, Australia.

    • Michael E Goddard

  321. Department of Food and Agricultural Systems, University of Melbourne, Melbourne, Victoria, Australia.

    • Michael E Goddard

Consortia

  1. The Electronic Medical Records and Genomics (eMERGE) Consortium

    A full list of members and affiliations appears in the Supplementary Note.

  2. The MIGen Consortium

    A full list of members and affiliations appears in the Supplementary Note.

  3. The PAGE Consortium

    A full list of members and affiliations appears in the Supplementary Note.

  4. The LifeLines Cohort Study

    A full list of members and affiliations appears in the Supplementary Note.

Authors

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Contributions

Steering committee overseeing the consortium: G.R.A., T.L.A., I.B., S.I.B., M. Boehnke, I.B.B., P.D., C.S.F., T.M.F., L.C.G., I.M.H., J.N.H., D.J.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 (wrote, edited and commented on the manuscript): S.I.B., D.I.C., A.Y.C., T.E., T.M.F., J.N.H., E.I., T.H.P., S.V., P.M.V., M.N.W., A.R.W. and J.Y. Data preparation group (checked and prepared data from contributing cohorts for meta-analyses): D.C.C.-C., F.R.D., T.E., T. Fall, T. Ferreira, S.G., I.M.H., Z.K., C.M.L., A.E.L., R.J.F.L., J. Luan, R.M., J.C.R., A. Scherag, E.K.S., S.V., T.W.W., A.R.W. and T. Workalemahu. Height meta-analyses group (GWAS and Metabochip) (analyses specific to the manuscript): T.E., T.M.F. (chair), S.V., P.M.V., A.R.W. (lead meta-analyses) and J.Y. (lead joint-effects and approximate conditional analyses). Mixed linear model analyses: J.S.B., M. Boehnke, D.I.C., A.Y.C., K.E., T.M.F. (chair), S.G., J.N.H., J.H.Z., E.I., A.U.J., Z.K., R.J.F.L., J. Luan, A. Metspalu, E.M., J.R.O., A.L.P., A.G.U., S.V., P.M.V., M.N.W., A.R.W. (lead) and J.Y. Large λ group: T.M.F., J.N.H., P.M.V., M.E. Goddard, A.L.P., M.N.W., J.Y. and G.R.A. Family transmission analyses: G.R.A., N.A., I.B.B., Y.D., C.M.v.D., J.N.H. (chair), E.I., J.R.O., E.P., S.V. (lead), P.M.V. and J.Y. Variance, heritability and prediction analyses: K.E., M.E. Goddard, M.I.M., A.A.E.V., P.M.V. (chair), M.N.W., A.R.W. and J.Y. (lead). Biological enrichment and pathway analyses: T.E. (lead biological enrichment analyses), J.N.H. (chair) and T.H.P. (lead pathway analyses). ENCODE working group: M.L.B., G.L. (chair) and K.S.L. Gene expression (eQTL) working group: J. Baron, T.E. (chair), L. Franke, J. Karjalainen, J.C.L., A. Metspalu, E.R., J.E.P. and H.-J.W. (lead). Other contributions: (DEPICT) R.F., L. Franke, J. Karjalainen and T.H.P.

Project design, management and coordination of contributing studies

Previous GWAS: (AGES) V. Gudnason, T.B.H.; (AMISH) A.R.S.; (ARIC) K.E.N.; (B58C T1D CONTROLS) D.P.S.; (B58C WTCCC) D.P.S.; (BRIGHT) M.J.B., N.J.S.; (CAPS) E.I.; (CHS) J.I.R.; (COLAUS) J.S.B., S. Bergmann; (CROATIA-Vis) I.R.; (deCODE) K. Stefansson, U.T.; (DGI) L.C.G.; (EGCUT) A. Metspalu; (EPIC-Norfolk) N.J.W.; (FENLAND) N.J.W.; (Finnish Twin Cohort) J. Kaprio, K. Silventoinen; (FRAM) L.A.C.; (FUSION) R.N.B., M. Boehnke; (GerMIFS I) J.E., C. Hengstenberg; (GerMIFS II) H. Schunkert; (H2000) S. Koskinen; (HFPS) D.J.H.; (KORA S4) C.G., A.P.; (MICROS) A.A.H., P.P.P.; (NFBC66) M.-R.J., S. Sebert; (NHS) D.J.H.; (NSPHS) U.G.; (NTRNESDA) D.I.B.; (ORCADES) H.C.; (PLCO) S.I.B., S.J.C.; (RS I) C.M.v.D., A. Hofman, M. Kayser, F. Rivadeneira, A.G.U.; (RUNMC) L.A.K.; (SardiNIA) G.R.A.; (SASBAC) E.I.; (SHIP) R.B., H.V.; (WGHS) P.M.R.; (WTCCC-CAD) A.S.H., N.J.S.; (WTCCC-T2D) C.M.L., M.I.M.; (Young Finns Study (YFS)) T.L., O.T.R.

New GWAS: (ASCOT) M.J.C., P.S.; (ATCG) P.I.W.d.B., D.W.H.; (Athero-Express Biobank Studies) F.W.A., H.M.d.R., F.L.M., G.P.; (B-PROOF) R.D.-R., L.C.P.G.M.d.G., N.M.v.S., N.v.d.V.; (BLSA) L. Ferrucci; (CLHNS) K.L. Mohlke; (COROGENE) M.P., J. Sinisalo; (DESIR) S.C., P.F.; (DNBS) M. Melbye, J.C.M.; (EGCUT) A. Metspalu; (eMERGE) M.G.H.; (ERF) B.A.O., C.M.v.D.; (FamHS) I.B.B.; (FINGESTURE) J.-C.T.; (GOOD) C.O.; (HBCS) J.G.E.; (Health ABC) T.B.H., Y. Liu; (HERITAGE Family Study) C. Bouchard, D.C.R., M.A. Sarzynski; (InCHIANTI) L. Ferrucci, T.M.F.; (IPM) E.P.B., R.J.F.L.; (LLS) P.E.S.; (LOLIPOP) J.C.C., J.S.K.; (MGS) P.V.G.; (NELSON) P.I.W.d.B., P.Z.; (PLCO2) S.I.B., S.J.C.; (PREVEND) P.v.d.H.; (PROCARDIS) H.W.; (PROSPER/PHASE) I.F., J.W.J.; (QFS) C. Bouchard, A. Marette, L.P., M.-C.V.; (QIMR) A.C.H., N.G.M., G.W.M.; (RISC) E.F., T.M.F., A. Golay, M. Walker; (RS II) A. Hofman, M. Kayser, F. Rivadeneira, A.G.U.; (RS III) A. Hofman, M. Kayser, F. Rivadeneira, A.G.U.; (SHIP-TREND) R.B., H.V.; (SORBS) A. Tönjes; (TRAILS) A.J.O., H. Snieder; (TWINGENE) E.I.; (TwinsUK) T.D.S.

Metabochip studies: (ADVANCE) T.L.A., T.Q.; (AMC-PAS) G.K.H., P.D.; (ARIC) E.B., K.E.N.; (B1958C) E.H., C.P.; (BHS) J. Beilby, J. Hui; (CARDIOGENICS) P.D., W.H.O., H. Schunkert; (DESIR) S.C., P.F.; (DGE DietGeneExpression) B.J.; (DIAGEN) S.R.B., P.E.H.S.; (DILGOM) P.J., A.M.J., S. Männistö, M.P., V. Salomaa; (DPS) M.U.; (DR's EXTRA) T.A.L., R. Rauramaa; (DUNDEE-GoDARTS) C.N.A.P.; (EAS) J.F.P.; (EGCUT) A. Metspalu; (EMIL (SWABIA)) B.O.B.; (FBPP) A.C., R.S.C., S.C.H.; (FIN-D2D 2007) S.M.K.-K., T.E.S.; (FUSION 2) F.S.C., J. Saramies, J.T.; (GLACIER) P.W.F.; (GxE) R.S.C., J.N.H., C.A.M.; (HNR) R.E., P. Hoffmann, S. Moebus; (HUNT 2) K.H.; (IMPROVE) U.d.F., A. Hamsten, S.E.H., E.T.; (KORA S3) T.M., H.-E.W.; (KORA S4) K. Strauch; (Leipzig) M.S.; (LURIC) W.M.; (MEC) C.A. Haiman, L.L.M.; (METSIM) J. Kuusisto, M. Laakso; (MORGAM) P.A., D. Arveiler, P. Brambilla, J.F., F.K., J.V.; (NSHD) D.K.; (PIVUS) E.I.; (PROMIS) J. Danesh, P.D., D. Saleheen; (ScarfSheep) A. Hamsten; (SPT) R.S.C., J.N.H., C.A.M.; (STR) E.I., (Tandem) M. Bochud, P. Bovet; (THISEAS) G. Dedoussis, P.D.; (Tromsø) I.N.; (ULSAM) E.I.; (WHI) T.C.M., C.K., U.P.; (Whitehall) A.D.H., M. Kivimaki, N.J.W.; (WTCCC-T2D) C.M.L., M.I.M.

Genotyping of contributing studies

Previous GWAS: (AGES) A.V. Smith; (B58C T1D CONTROLS) W.L.M.; (B58C WTCCC) W.L.M.; (CAPS) H. Grönberg; (CROATIA-Vis) C. Hayward; (EGCUT) M. Nelis; (EPIC-Norfolk) N.J.W.; (FENLAND) N.J.W.; (Finnish Twin Cohort) J. Kaprio; (KORA S3) T.I., M.M.-N.; (MICROS) A.A.H.; (NFBC66) M.-R.J.; (ORCADES) A.F.W.; (PLCO) S.J.C.; (RS I) K.E., C.M.-G., F. Rivadeneira, A.G.U.; (SASBAC) P. Hall; (SHIP) A. Hannemann, M. Nauck; (WGHS) D.I.C., L.M.R.; (WTCCC-CAD) A.S.H., N.J.S.; (WTCCC-T2D) A.T.H., M.I.M.; (Young Finns Study (YFS)) T.L., O.T.R.

New GWAS: (ASCOT) P.B.M.; (ATCG) P.I.W.d.B., D.W.H., P.J.M.; (Athero-Express Biobank Study) S.W.v.d.L.; (CLHNS) D.C.C.-C.; (DESIR) E.E., S. Lobbens; (EGCUT) T.E., L.M.; (eMERGE) D.C.C., M.G.H.; (ERF) A.I., B.A.O., C.M.v.D.; (FamHS) I.B.B., E.W.D., M.F.F., A.T.K., M.K.W., Q.Z.; (GOOD) C.O., M. Lorentzon; (Health ABC) Y. Liu; (HERITAGE Family Study) M.A. Sarzynski; (HYPERGENES) S. Lupoli.; (IPM) E.P.B.; (LifeLines) M.A. Swertz; (LLS) J. Deelen, Q.H.; (LOLIPOP) J.C.C., J.S.K.; (NELSON) J. Smolonska; (PLCO2) S.J.C., K.B.J., Z.W.; (PREVEND) P.v.d.H., I.M.L.; (PROCARDIS) M.F., A. Goel; (PROSPER/PHASE) J.W.J., D.J.S., S.T.; (QFS) C. Bellis, J. Blangero; (QIMR) A.K.H.; (SHIP-TREND) A. Hannemann, M. Nauck; (RS II) K.E., C.M.-G., F. Rivadeneira, A.G.U.; (RS III) K.E., C.M.-G., F. Rivadeneira, A.G.U.; (TRAILS) M. Bruinenberg, C.A. Hartman; (TWINGENE) A. Hamsten, N.L.P.; (TwinsUK) M. Mangino, A. Moayyeri; (WGHS) D.I.C., L.M.R.

Metabochip studies: (ADVANCE) D. Absher, T.L.A., T.Q.; (AMCPAS) K. Stirrups; (ARIC) E.B., K.E.N.; (B1958C) N.R.R., C.J.G., T.J.; (BHS) G.M.A., J. Hui; (CARDIOGENICS) K. Stirrups; (DESIR) E.E., S. Lobbens; (DGE DietGeneExpression) B.J.; (DIAGEN) M.A.M.; (DUNDEE-GoDARTS) A.J.B., C.N.A.P., N.W.R.; (EAS) J.F.W.; (EGCUT) T.E., L.M.; (ELY) N.G.F., C.L., R.J.F.L., K.K.O., R.A.S., N.J.W.; (EMIL (SWABIA)) B.O.B.; (EPIC-Norfolk) N.G.F., C.L., R.J.F.L., K.K.O., R.A.S., N.J.W.; (FBPP) A.C.; (FENLAND) N.G.F., C.L., R.J.F.L., K.K.O., R.A.S., N.J.W.; (FIN-D2D 2007) P.S.C.; (FUSION 2) L.K.; (GLACIER) I.B.; (HNR) M.M.N.; (HUNT 2) N.N.; (KORA S3) N.K., M. Waldenberger; (KORA S4) H. Grallert, P.L.; (Leipzig) Y.B., P.K.; (LURIC) M.E.K.; (MEC) C.A. Haiman, L.A.H.; (NSHD) D.K., K.K.O., A.W.; (PIVUS) E.I., C. Berne, L.L., J. Sundström; (PROMIS) K. Stirrups; (STR) N.L.P.; (Tandem) G.B.E., M. Maillard; (THISEAS) K. Stirrups; (Tromsø) P.S.C.; (ULSAM) J.Ä., E.I., A.-C.S.; (WHI) C.K., U.P.; (Whitehall) C.L.; (WTCCC-T2D) A.T.H., M.I.M.

Phenotype coordination of contributing studies

Previous GWAS: (AMISH) A.R.S.; (B58C T1D CONTROLS) D.P.S.; (B58C WTCCC) D.P.S.; (BRIGHT) M.J.B., N.J.S.; (CAPS) H. Grönberg; (CHS) R.C.K.; (CROATIA-Vis) I.R.; (DGI) V. Lyssenko; (EGCUT) A. Metspalu; (EPIC-Norfolk) N.J.W.; (FENLAND) N.J.W.; (Finnish Twin Cohort) J. Kaprio; (KORA S4) A.P.; (NFBC66) M.-R.J.; (NTRNESDA) J.H.S.; (ORCADES) A.F.W.; (PLCO) S.I.B.; (RS I) A. Hofman, F. Rivadeneira, A.G.U.; (SASBAC) P. Hall; (SHIP) M. Dörr, W.H., T.K.; (UKBS-CC) J. Jolley; (WGHS) D.I.C., L.M.R., A.Y.C.; (WTCCC-CAD) A.S.H., N.J.S.; (WTCCC-T2D) A.B., A.T.H.; (Young Finns Study (YFS)) T.L., O.T.R.

New GWAS: (ASCOT) M.J.C., P.S., A.V. Stanton; (ATCG) D.W.H.; (Athero-Express Biobank Study) F.L.M.; (BLSA) S. Bandinelli; (DESIR) R. Roussel; (DNBC) H.A.B., B.F., F.G.; (EGCUT) T.E., A. Metspalu; (eMERGE) J.C.D., A.N.K.; (ERF) B.A.O., C.M.v.D.; (FamHS) I.B.B., M.F.F.; (FINGESTURE) J. Junttila; (GOOD) C.O., M. Lorentzon; (HBCS) J.G.E.; (Health ABC) M.E. Garcia, T.B.H., M.A.N.; (HERITAGE Family Study) C. Bouchard; (HYPERGENES) P.M.; (InCHIANTI) S. Bandinelli, L. Ferrucci; (IPM) O.G.; (LifeLines) S. Scholtens, M.A. Swertz, J.M.V.; (LLS) D.v.H.; (LOLIPOP) J.C.C., J.S.K., U.A., L.O., J. Sehmi; (NELSON) P.A.D.J.; (PLCO2) S.I.B.; (PREVEND) S.J.L.B., R.T.G., H.L.H.; (PROCARDIS) R. Clarke, R. Collins, M.F., A. Hamsten; (PROSPER/PHASE) J.W.J., I.F., B.M.B.; (QFS) A. Tremblay; (QIMR) A.K.H., A.C.H., P.A.F.M., N.G.M., G.W.M.; (RS II) A. Hofman, F. Rivadeneira, A.G.U.; (RS III) A. Hofman, F. Rivadeneira, A.G.U.; (SORBS) A. Tönjes; (SHIP-TREND) M. Dörr, W.H., T.K.; (TRAILS) C.A. Hartman, R.P.S., F.V.A.v.O.; (TWINGENE) P.K.E.M., N.L.P.; (TwinsUK) M. Mangino, C.M.; (WGHS) D.I.C., L.M.R.

Metabochip studies: (ADVANCE) A.S.G., M.A.H.; (AMCPAS) J.J.P.K.; (ARIC) E.B.; (B1958C) E.H., C.P.; (BHS) A.L.J., A.W.M.; (DESIR) R. Roussel; (DGE DietGeneExpression) B.J., I.H.C.; (DIAGEN) J. Gräßler, G.M.; (DPS) J. Lindström; (DR's EXTRA) M.H.; (DUNDEE-GoDARTS) A.S.F.D., A.D.M., C.N.A.P.; (EAS) S. McLachlan; (EGCUT) T.E., A. Metspalu; (EMIL (SWABIA)) B.O.B., S.C.-B., W.K., S. Merger, T.S., R.W.; (FBPP) R.S.C., S.C.H.; (GLACIER) G. Hallmans; (GxE) T. Forrester, B.O.T.; (HNR) R.E., S. Moebus; (HUNT 2) O.H.; (KORA S3) H.-E.W.; (Leipzig) M. Blüher; (MEC) L.R.W.; (METSIM) H.M.S.; (NSHD) D.K.; (PIVUS) C. Berne, E.I., L.L., J. Sundström; (PROMIS) D. Saleheen; (SPT) T. Forrester, B.O.T.; (STR) N.L.P.; (Tandem) M. Bochud, P. Bovet; (THISEAS) S. Kanoni; (Tromsø) T. Wilsgaard; (ULSAM) J.Ä., V. Giedraitis, E.I.; (WHI) C.K., U.P.; (Whitehall) M. Kumari; (WTCCC-T2D) A.B., A.T.H.

Data analysis

Previous GWAS: (AGES) A.V. Smith; (ARIC) K.L. Monda, K.E.N.; (B58C T1D CONTROLS) D.P.S.; (B58C WTCCC) D.P.S.; (CAPS) E.I.; (CHS) R.C.K., B.M.; (COLAUS) S. Bergmann, Z.K.; (CROATIA-Vis) C. Hayward; (deCODE) V. Steinthorsdottir, G.T.; (EGCUT) M. Nelis; (EPIC-Norfolk) J.H.Z.; (FENLAND) J. Luan; (FRAM) L.A.C., N.L.H.-C.; (FUSION) C.J.W.; (GerMIFS II) C.W.; (H2000) N.E.; (HPFS) L.Q.; (NHS) L.Q.; (NSPHS) Å.J.; (PLCO) S.I.B.; (RS I) K.E., C.M.-G., F. Rivadeneira, A.G.U.; (RUNMC) S.H.V.; (SardiNIA) S. Sanna; (SASBAC) E.I.; (SEARCH) J.P.T.; (SHIP) A. Teumer; (WGHS) D.I.C., L.M.R., A.Y.C.; (WTCCC-T2D) A.P.M., T. Ferreira, A. Mahajan, R.M.

New GWAS: (ATCG) P.I.W.d.B., P.J.M., S.R.; (Athero-Express Biobank Studies) S.W.v.d.L.; (B-PROOF) S.v.D.; (BHS) M.C.; (BLSA) T.T.; (CLHNS) D.C.C.-C.; (DESIR) S.C., L.Y.; (DNBC) B.F., F.G.; (EGCUT) T.E., K.F., T.H., R.M.; (eMERGE) M.G.H.; (ERF) N.A., A.D.; (FamHS) M.F.F.; (GOOD) C.O., M. Lorentzon; (HBCS) N.E.; (Health ABC) M.A.N.; (HERITAGE Family Study) C. Bouchard, M.A. Sarzynski, D.C.R., T.R., T.K.R., Y.J.S.; (HYPERGENES) S. Lupoli; (InCHIANTI) D.P., T.T., A.R.W.; (IPM) J. Jeff, V. Lotay, Y. Lu; (LifeLines) I.M.N., J.V.V.V.-O.; (LLS) M. Beekman, J.J.H.-D.; (LOLIPOP) W.Z.; (MGS) J. Shi, (NELSON) S.R., J.v.S.; (PLCO2) S.I.B., Z.W.; (PREVEND) P.v.d.H., I.M.L., N.V.; (PROCARDIS) A. Goel; (PROSPER/PHASE) I.F., B.M.B., S.T.; (QFS) J. Blangero, L.P.; (QIMR) G. Hemani, D.R.N., J.E.P.; (RISC) D.P., A.R.W.; (RS II) K.E., C.M.-G., F. Rivadeneira, A.G.U.; (RS III) K.E., C.M.-G., F. Rivadeneira, A.G.U.; (SHIP-TREND) A. Teumer; (SORBS) R.M.; (TRAILS) H. Snieder; (TWINGENE) E.I., S.G.; (TwinsUK) M. Mangino; (WGHS) D.I.C., L.M.R.

Metabochip studies: (ADVANCE) D. Absher, T.L.A., L.L.W.; (AMCPAS) S. Kanoni; (ARIC) S. Buyske, A.E.J., K.E.N.; (B1958C) T. Ferreira; (BHS) D. Anderson; (CARDIOGENICS) S. Kanoni; (DESIR) S.C., L.Y.; (DGE DietGeneExpression) I.H.C.; (DIAGEN) A.U.J., G.M.; (DILGOM) K.K.; (DUNDEE) T. Ferreira; (EAS) J.L.B., R.M.F.; (EGCUT) T.E., K.F., E.M.; (ELY) J. Luan; (EMIL (SWABIA)) B.O.B.; (EPIC-Norfolk) J. Luan; (FBPP) A.C., G.B.E.; (FENLAND) J. Luan; (GLACIER) F. Renstrom, D. Shungin; (GxE) C.D.P.; (HNR) S. Pechlivanis, A. Scherag; (IMPROVE) L. Folkersen, R.J.S.; (KORA S3) J.S.R.; (KORA S4) E.A.; (Leipzig) A. Mahajan, I.P.; (LURIC) G. Delgado, T.B.G., M.E.K., S. Pilz, H. Scharnag; (MEC) U.L., F.R.S.; (METSIM) A. Stancáková; (NSHD) A.W., J. Luan; (PIVUS) S.G., E.I.; (PROMIS) S. Kanoni; (ScarfSheep) R.J.S.; (SPT) C.D.P.; (STR) E.I., S.G.; (TANDEM) G.B.E.; (THISEAS) M. Dimitriou; (ULSAM) S.G., E.I.; (WHI) J. Gong, J. Haessler, M.R.; (Whitehall) J. Luan; (WTCCC-T2D) A.P.M., T. Ferreira, A. Mahajan, R.M.

Competing interests

The authors declare no competing financial interests.

Corresponding authors

Correspondence to Peter M Visscher or Joel N Hirschhorn or Timothy M Frayling.

Integrated supplementary information

Supplementary figures

  1. 1.

    Overview of the analysis strategy implemented.

  2. 2.

    Quantile-quantile plot showing the P-value distribution of GWAS meta-analysis results after applying a single genomic control.

  3. 3.

    Manhattan plots.

  4. 4.

    Examples of loci with multiple signals that cluster around the same gene or are in close proximity to others.

  5. 5.

    Partitioning the variance in the SNP-derived genetic predictor using a within-family analysis.

  6. 6.

    Variance explained by the SNPs at known loci.

  7. 7.

    Results from GRAIL analysis.

  8. 8.

    DEPICT cell type enrichment analysis.

  9. 9.

    Partitioning the variance explained by chromosome length.

Supplementary information

PDF files

  1. 1.

    Supplementary Text and Figures

    Supplementary Figures 1–9, Supplementary Tables 2, 4–6, 9–12, 15 and 17–22, and Supplementary Note.

Excel files

  1. 1.

    Supplementary Table 1

    SNPs representing signals identified through meta-analysis and approximate conditional analysis.

  2. 2.

    Supplementary Table 3

    Results from Metabochip replication analysis.

  3. 3.

    Supplementary Table 7

    Nonsynonymous variant enrichment analysis.

  4. 4.

    Supplementary Table 8

    Cis-regulatory variant enrichment analysis.

  5. 5.

    Supplementary Table 13

    Results from GRAIL text-mining analysis.

  6. 6.

    Supplementary Table 14

    Pathways and protein complexes prioritized by DEPICT.

  7. 7.

    Supplementary Table 16

    Results from DEPICT gene prioritization analysis.

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https://doi.org/10.1038/ng.3097

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