Article

Rare and low-frequency coding variants alter human adult height

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Abstract

Height is a highly heritable, classic polygenic trait with approximately 700 common associated variants identified through genome-wide association studies so far. Here, we report 83 height-associated coding variants with lower minor-allele frequencies (in the range of 0.1–4.8%) and effects of up to 2 centimetres per allele (such as those in IHH, STC2, AR and CRISPLD2), greater than ten times the average effect of common variants. In functional follow-up studies, rare height-increasing alleles of STC2 (giving an increase of 1–2 centimetres per allele) compromised proteolytic inhibition of PAPP-A and increased cleavage of IGFBP-4 in vitro, resulting in higher bioavailability of insulin-like growth factors. These 83 height-associated variants overlap genes that are mutated in monogenic growth disorders and highlight new biological candidates (such as ADAMTS3, IL11RA and NOX4) and pathways (such as proteoglycan and glycosaminoglycan synthesis) involved in growth. Our results demonstrate that sufficiently large sample sizes can uncover rare and low-frequency variants of moderate-to-large effect associated with polygenic human phenotypes, and that these variants implicate relevant genes and pathways.

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Acknowledgements

A full list of acknowledgments appears in the Supplementary Information. Part of this work was conducted using the UK Biobank resource.

Author information

Author notes

    • Eirini Marouli
    • , Mariaelisa Graff
    • , Carolina Medina-Gomez
    • , Ken Sin Lo
    • , Andrew R. Wood
    • , Troels R. Kjaer
    • , Rebecca S. Fine
    •  & Yingchang Lu

    These authors contributed equally to this work.

    • Claus Oxvig
    • , Zoltán Kutalik
    • , Fernando Rivadeneira
    • , Ruth J. F. Loos
    • , Timothy M. Frayling
    • , Joel N. Hirschhorn
    • , Panos Deloukas
    •  & Guillaume Lettre

    These authors jointly supervised this work.

Affiliations

  1. William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London EC1M 6BQ, UK

    • Eirini Marouli
    • , Kathleen E. Stirrups
    • , Mark J. Caulfield
    • , Stavroula Kanoni
    • , Patricia B. Munroe
    • , Ioanna Ntalla
    • , Helen R. Warren
    •  & Panos Deloukas
  2. Department of Epidemiology, University of North Carolina, Chapel Hill, North Carolina 27514, USA

    • Mariaelisa Graff
    • , Heather M. Highland
    • , Anne E. Justice
    • , Kristin L. Young
    •  & Angela L. Mazul
  3. Department of Epidemiology, Erasmus Medical Center, Rotterdam, 3015 GE, The Netherlands

    • Carolina Medina-Gomez
    • , M. Arfan Ikram
    • , André G. Uitterlinden
    • , Cornelia M. van Duijn
    •  & Fernando Rivadeneira
  4. Department of Internal Medicine, Erasmus Medical Center, Rotterdam, 3015 GE, The Netherlands

    • Carolina Medina-Gomez
    • , Linda Broer
    • , André G. Uitterlinden
    •  & Fernando Rivadeneira
  5. Montreal Heart Institute, Montreal, Quebec H1T 1C8, Canada

    • Ken Sin Lo
    • , Valérie Turcot
    • , Simon de Denus
    • , John D. Rioux
    • , Jean-Claude Tardif
    •  & Guillaume Lettre
  6. Genetics of Complex Traits, University of Exeter Medical School, University of Exeter, Exeter EX2 5DW, UK

    • Andrew R. Wood
    • , Katherine S. Ruth
    • , Hanieh Yaghootkar
    •  & Timothy M. Frayling
  7. Department of Molecular Biology and Genetics, Aarhus University, Aarhus, 8000, Denmark

    • Troels R. Kjaer
    •  & Claus Oxvig
  8. Broad Institute of MIT and Harvard, Cambridge, Massachusetts 02142, USA

    • Rebecca S. Fine
    • , Tõnu Esko
    • , Manuel A. Rivas
    • , Sailaja Vedantam
    • , Daniel I. Chasman
    • , Aniruddh P. Patel
    • , Sekar Kathiresan
    •  & Joel N. Hirschhorn
  9. Department of Genetics, Harvard Medical School, Boston, Massachusetts 02115, USA

    • Rebecca S. Fine
    •  & Sailaja Vedantam
  10. Division of Endocrinology and Center for Basic and Translational Obesity Research, Boston Children’s Hospital, Boston, Massachusetts 02115, USA

    • Rebecca S. Fine
    • , Tõnu Esko
    • , Sailaja Vedantam
    •  & Joel N. Hirschhorn
  11. Division of Epidemiology, Department of Medicine, Vanderbilt-Ingram Cancer Center, Vanderbilt Epidemiology Center, Vanderbilt University School of Medicine, Nashville, Tennessee 37203, USA

    • Yingchang Lu
  12. The Charles Bronfman Institute for Personalized Medicine, Icahn School of Medicine at Mount Sinai, New York, New York 10029, USA

    • Yingchang Lu
    • , Claudia Schurmann
    • , Erwin P. Bottinger
    •  & Ruth J. F. Loos
  13. The Genetics of Obesity and Related Metabolic Traits Program, Ichan School of Medicine at Mount Sinai, New York, New York 10069, USA

    • Yingchang Lu
    • , Claudia Schurmann
    •  & Ruth J. F. Loos
  14. Human Genetics Center, The University of Texas School of Public Health, The University of Texas Graduate School of Biomedical Sciences at Houston, The University of Texas Health Science Center at Houston, Houston, Texas 77030, USA

    • Heather M. Highland
  15. Institute of Social and Preventive Medicine, Lausanne University Hospital, Lausanne, 1010, Switzerland

    • Sina Rüeger
    •  & Zoltán Kutalik
  16. Swiss Institute of Bioinformatics, Lausanne, 1015, Switzerland

    • Sina Rüeger
    • , David Lamparter
    • , Sven Bergmann
    •  & Zoltán Kutalik
  17. deCODE Genetics/Amgen inc., Reykjavik, 101, Iceland

    • Gudmar Thorleifsson
    • , Valgerdur Steinthorsdottir
    • , Unnur Thorsteinsdottir
    •  & Kari Stefansson
  18. Department of Computational Biology, University of Lausanne, Lausanne, 1011, Switzerland

    • David Lamparter
    •  & Sven Bergmann
  19. Department of Haematology, University of Cambridge, Cambridge CB2 0PT, UK

    • Kathleen E. Stirrups
  20. Department of Genetic Epidemiology, University of Regensburg, Regensburg, D-93051, Germany

    • Thomas W. Winkler
    • , Mathias Gorski
    •  & Iris M. Heid
  21. Estonian Genome Center, University of Tartu, Tartu, 51010, Estonia

    • Tõnu Esko
    •  & Andres Metspalu
  22. Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford OX3 7BN, UK

    • Tugce Karaderi
    • , Anubha Mahajan
    • , Wei Gan
    • , Hidetoshi Kitajima
    • , Andrew P. Morris
    • , Neil Robertson
    • , Lorraine Southam
    • , Mark I. McCarthy
    •  & Cecilia M. Lindgren
  23. Department of Biostatistics and Center for Statistical Genetics, University of Michigan, Ann Arbor, Michigan 48109, USA

    • Adam E. Locke
    • , Goncalo Abecasis
    • , Shuang Feng
    • , Anne U. Jackson
    • , Xueling Sim
    • , Heather M. Stringham
    •  & Michael Boehnke
  24. McDonnell Genome Institute, Washington University School of Medicine, Saint Louis, Missouri 63108, USA

    • Adam E. Locke
  25. Department of Cardiovascular Sciences, Univeristy of Leicester, Glenfield Hospital, Leicester LE3 9QP, UK

    • Nicholas G. D. Masca
    • , Christopher P. Nelson
    •  & Nilesh J. Samani
  26. NIHR Leicester Cardiovascular Biomedical Research Unit, Glenfield Hospital, Leicester LE3 9QP, UK

    • Nicholas G. D. Masca
    • , Christopher P. Nelson
    •  & Nilesh J. Samani
  27. Center for Diabetes Research, Wake Forest School of Medicine, Winston-Salem, North Carolina 27157, USA

    • Maggie C. Y. Ng
    • , Poorva Mudgal
    • , Donald W. Bowden
    •  & Amanda J. Cox
  28. Center for Genomics and Personalized Medicine Research, Wake Forest School of Medicine, Winston-Salem, North Carolina 27157, USA

    • Maggie C. Y. Ng
    • , Donald W. Bowden
    •  & Amanda J. Cox
  29. Nuffield Department of Clinical Medicine, Oxford OX3 7BN, UK

    • Manuel A. Rivas
  30. Institute for Translational Genomics and Population Sciences, LABioMed at Harbor-UCLA Medical Center, Torrance, California 90502, USA

    • Xiuqing Guo
    • , Yii-Der Ida Chen
    • , Yucheng Jia
    • , Yeheng Liu
    • , Kevin Sandow
    • , Kent D. Taylor
    • , Jie Yao
    •  & Jerome I Rotter
  31. Netherlands Comprehensive Cancer Organisation, Utrecht, 3501 DB, The Netherlands

    • Katja K. Aben
  32. Radboud University Medical Center, Nijmegen, 6500 HB, The Netherlands

    • Katja K. Aben
    • , Tessel E. Galesloot
    • , Lambertus A. Kiemeney
    • , Kirsten B. Kluivers
    •  & Sita H. Vermeulen
  33. Department of Nutrition, University of North Carolina, Chapel Hill, North Carolina 27599, USA

    • Linda S. Adair
  34. Centre for Control of Chronic Diseases (CCCD), Dhaka, 1212, Bangladesh

    • Dewan S. Alam
  35. Institute of Genetic Epidemiology, Helmholtz Zentrum München - German Research Center for Environmental Health, Neuherberg, D-85764, Germany

    • Eva Albrecht
    • , Iris M. Heid
    •  & Martina Müller-Nurasyid
  36. The Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, 2100, Denmark

    • Kristine H. Allin
    • , Emil V. Appel
    • , Jette Bork-Jensen
    • , Niels Grarup
    • , Torben Hansen
    • , Mette Hollensted
    • , Oluf Pedersen
    • , Henrik Vestergaard
    •  & Tune H. Pers
  37. Department of Family Medicine & Public Health, University of California, San Diego, La Jolla, California 92093, USA

    • Matthew Allison
  38. INSERM U1167, Lille, F-59019, France

    • Philippe Amouyel
  39. Institut Pasteur de Lille, U1167, Lille, F-59019, France

    • Philippe Amouyel
  40. Universite de Lille, U1167 - RID-AGE - Risk factors and molecular determinants of aging-related diseases, Lille, F-59019, France

    • Philippe Amouyel
  41. Department of Epidemiology and Public Health, University of Strasbourg, Strasbourg, F-67085, France

    • Dominique Arveiler
  42. Department of Public Health, University Hospital of Strasbourg, Strasbourg, 67081, France

    • Dominique Arveiler
  43. Department of Cardiology, Division Heart & Lungs, University Medical Center Utrecht, Utrecht, The Netherlands

    • Folkert W. Asselbergs
    •  & Jessica van Setten
  44. Durrer Center for Cardiogenetic Research, ICIN-Netherlands Heart Institute, Utrecht, The Netherlands

    • Folkert W. Asselbergs
  45. Institute of Cardiovascular Science, Faculty of Population Health Sciences, University College London, London, UK

    • Folkert W. Asselbergs
  46. Zilber School of Public Health, University of Wisconsin-Milwaukee, Milwaukee, Wisconsin 53201, USA

    • Paul L. Auer
  47. INSERM U1018, Centre de recherche en Épidemiologie et Sante des Populations (CESP), Villejuif, France

    • Beverley Balkau
  48. Department of Nephrology, University Hospital Regensburg, Regensburg, 93042, Germany

    • Bernhard Banas
    • , Carsten A. Böger
    •  & Mathias Gorski
  49. Department of Cardiology, Rigshospitalet, Copenhagen University Hospital, Copenhagen, 2100, Denmark

    • Lia E. Bang
  50. Department of Clinical Biochemistry, Herlev and Gentofte Hospital, Copenhagen University Hospital, Herlev, 2730, Denmark

    • Marianne Benn
    • , Pia R. Kamstrup
    • , Sune F. Nielsen
    • , Børge G. Nordestgaard
    •  & Anette Varbo
  51. Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, 2200, Denmark

    • Marianne Benn
    • , Ruth Frikke-Schmidt
    • , Torben Jørgensen
    • , Allan Linneberg
    • , Sune F. Nielsen
    • , Børge G. Nordestgaard
    • , Anne Tybjaerg-Hansen
    •  & Anette Varbo
  52. Department of Epidemiology, School of Public Health, University of Michigan, Ann Arbor, Michigan 48109, USA

    • Lawrence F. Bielak
    • , Min A. Jhun
    • , Sharon L. R. Kardia
    • , Jennifer A. Smith
    •  & Wei Zhao
  53. IFB Adiposity Diseases, University of Leipzig, Leipzig, 04103, Germany

    • Matthias Blüher
    • , Peter Kovacs
    •  & Michael Stumvoll
  54. University of Leipzig, Department of Medicine, Leipzig, 04103, Germany

    • Matthias Blüher
    •  & Michael Stumvoll
  55. Department of Epidemiology, German Institute of Human Nutrition Potsdam-Rehbruecke (DIfE), Nuthetal, 14558, Germany

    • Heiner Boeing
  56. School of Public Health, Human Genetics Center, The University of Texas Health Science Center at Houston, Houston, Texas 77030, USA

    • Eric Boerwinkle
    •  & Megan L. Grove
  57. Human Genome Sequencing Center, Baylor College of Medicine, Houston, Texas 77030, USA

    • Eric Boerwinkle
  58. Medical Genomics and Metabolic Genetics Branch, National Human Genome Research Institute, National Institutes of Health, Bethesda, Maryland 20892, USA

    • Lori L. Bonnycastle
    •  & Francis S. Collins
  59. Julius Center for Health Sciences and Primary Care, University Medical Center Utrecht, Utrecht, The Netherlands

    • Michiel L. Bots
    •  & Paul I. W. de Bakker
  60. Department of Biochemistry, Wake Forest School of Medicine, Winston-Salem, North Carolina 27157, USA

    • Donald W. Bowden
  61. Department of Clinical Biochemistry, Lillebaelt Hospital, Vejle, 7100, Denmark

    • Ivan Brandslund
  62. Institute of Regional Health Research, University of Southern Denmark, Odense, 5000, Denmark

    • Ivan Brandslund
  63. MRC Social Genetic and Developmental Psychiatry Centre, Institute of Psychiatry, Psychology and Neuroscience, King’s College London & NIHR Biomedical Research Centre for Mental Health at the Maudsley, London, SE5 8AF, UK

    • Gerome Breen
  64. Marshfield Clinic Research Foundation, Marshfield, Wisconsin 54449, USA

    • Murray H. Brilliant
    • , Peggy L. Peissig
    •  & Thomas N. Person
  65. Department of Medicine, University of Washington, Seattle, Washington 98195, USA

    • Amber A. Burt
    • , Gail P. Jarvik
    •  & Eric B. Larson
  66. MRC/BHF Cardiovascular Epidemiology Unit, Department of Public Health and Primary Care, University of Cambridge, Cambridge CB1 8RN, UK

    • Adam S. Butterworth
    • , Rajiv Chowdhury
    • , John Danesh
    • , Emanuele Di Angelantonio
    • , Joanna M. M. Howson
    • , Praveen Surendran
    •  & Robin Young
  67. NIHR Blood and Transplant Research Unit in Donor Health and Genomics, University of Cambridge, Cambridge CB1 8RN, UK

    • Adam S. Butterworth
    • , John Danesh
    •  & Emanuele Di Angelantonio
  68. The Sigfried and Janet Weis Center for Research, Danville, Pennsylvania 17822, USA

    • David J. Carey
    • , Helena Kuivaniemi
    •  & Gerard Tromp
  69. NIHR Barts Cardiovascular Research Unit, Barts and The London School of Medicine & Dentistry, Queen Mary University, London EC1M 6BQ, UK

    • Mark J. Caulfield
    • , Patricia B. Munroe
    •  & Helen R. Warren
  70. Department of Cardiology, London North West Healthcare NHS Trust, Ealing Hospital, Middlesex UB1 3HW, UK

    • John C. Chambers
    • , Jaspal S. Kooner
    •  & Weihua Zhang
  71. Department of Epidemiology and Biostatistics, School of Public Health, Imperial College London, London W2 1PG, UK

    • John C. Chambers
    • , Evangelos Evangelou
    • , Marcelo P. Segura-Lepe
    •  & Weihua Zhang
  72. Imperial College Healthcare NHS Trust, London W12 0HS, UK

    • John C. Chambers
    •  & Jaspal S. Kooner
  73. Division of Genetics, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts 02115, USA

    • Daniel I. Chasman
  74. Division of Preventive Medicine, Brigham and Women's and Harvard Medical School, Boston, Massachusetts 02215, USA

    • Daniel I. Chasman
    • , Audrey Y. Chu
    •  & Paul M. Ridker
  75. Harvard Medical School, Boston, Massachusetts 02115, USA

    • Daniel I. Chasman
    • , Aniruddh P. Patel
    • , Paul M. Ridker
    •  & Sekar Kathiresan
  76. Medical Department, Lillebaelt Hospital, Vejle, 7100, Denmark

    • Cramer Christensen
  77. NHLBI Framingham Heart Study, Framingham, Massachusetts 01702, USA

    • Audrey Y. Chu
    •  & Nancy L. Heard-Costa
  78. Department of Medical, Surgical and Health Sciences, University of Trieste, Trieste, 34100, Italy

    • Massimiliano Cocca
  79. Department of Biostatistics, University of Liverpool, Liverpool L69 3GL, UK

    • James P. Cook
    •  & Andrew P. Morris
  80. Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh EH8 9JZ, UK

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    • , Gail Davies
    • , Ian J. Deary
    •  & John M. Starr
  81. Department of Psychology, University of Edinburgh, Edinburgh EH8 9JZ, UK

    • Janie Corley
    • , Gail Davies
    • , Ian J. Deary
    •  & Alison Pattie
  82. Department of Human Genetics, Radboud University Medical Center, Nijmegen, 6500 HB, The Netherlands

    • Jordi Corominas Galbany
  83. Menzies Health Institute Queensland, Griffith University, Southport, Queensland, Australia

    • Amanda J. Cox
  84. Diamantina Institute, University of Qeensland, Brisbane, Queensland, 4072, Australia

    • Gabriel Cuellar-Partida
  85. QIMR Berghofer Medical Research Institute, Brisbane, Queensland, 4006, Australia

    • Gabriel Cuellar-Partida
    • , Scott D. Gordon
    • , Nicholas G. Martin
    •  & Yadav Sapkota
  86. Wellcome Trust Sanger Institute, Wellcome Genome Campus, Hinxton, Cambridge CB10 1SA, UK

    • John Danesh
    • , Gaëlle Marenne
    • , Lorraine Southam
    •  & Eleftheria Zeggini
  87. British Heart Foundation, Cambridge Centre of Excellence, Department of Medicine, University of Cambridge, Cambridge CB2 0QQ, UK

    • John Danesh
  88. Department of Genetics, Center for Molecular Medicine, University Medical Center Utrecht, Utrecht, 3584 CX, The Netherlands

    • Paul I. W. de Bakker
  89. Department of Vascular Surgery, Division of Surgical Specialties, University Medical Center Utrecht, Utrecht, 3584 CX, The Netherlands

    • Gert J. de Borst
  90. Faculty of Pharmacy, Université de Montréal, Montreal, Quebec, H3T 1J4, Canada

    • Simon de Denus
  91. Department of Clinical Chemistry and Haematology, Division of Laboratory and Pharmacy, University Medical Center Utrecht, Utrecht, 3508 GA, The Netherlands

    • Mark C. H. de Groot
  92. Utrecht Institute for Pharmaceutical Sciences, Division Pharmacoepidemiology & Clinical Pharmacology, Utrecht University, Utrecht, 3508 TB, The Netherlands

    • Mark C. H. de Groot
  93. Department of Clinical Epidemiology, Leiden University Medical Center, Leiden, 2300 RC, The Netherlands

    • Renée de Mutsert
    • , Ruifang Li-Gao
    •  & Dennis O. Mook-Kanamori
  94. Department of Nutrition and Dietetics, School of Health Science and Education, Harokopio University, Athens, 17671, Greece

    • George Dedoussis
    •  & Aliki-Eleni Farmaki
  95. Division of Epidemiology & Community Health, School of Public Health, University of Minnesota, Minneapolis, Minnesota 55454, USA

    • Ellen W. Demerath
  96. Department of Ophthalmology, Radboud University Medical Center, Nijmegen, 6500 HB, The Netherlands

    • Anneke I. den Hollander
    •  & Carel B. Hoyng
  97. Centre for Cancer Genetic Epidemiology, Department of Public Health and Primary Care, University of Cambridge, Cambridge CB1 8RN, UK

    • Joe G. Dennis
    • , Douglas F. Easton
    •  & Deborah J. Thompson
  98. Institute of Cardiovascular Science, University College London, London WC1E 6JF, UK

    • Fotios Drenos
  99. MRC Integrative Epidemiology Unit, School of Social & Community Medicine, University of Bristol, Bristol BS8 2BN, UK

    • Fotios Drenos
  100. Fred Hutchinson Cancer Research Center, Public Health Sciences Division, Seattle, Washington 98109, USA

    • Mengmeng Du
  101. Memorial Sloan Kettering Cancer Center, Department of Epidemiology and Biostatistics, New York, New York 10017, USA

    • Mengmeng Du
  102. Centre for Cancer Genetic Epidemiology, Department of Oncology, University of Cambridge, Cambridge CB1 8RN, UK

    • Alison M. Dunning
    • , Douglas F. Easton
    • , Ailith Pirie
    •  & Jonathan P. Tyrer
  103. Department of Medicine, Oulu University Hospital, Oulu, 90029, Finland

    • Tapani Ebeling
  104. Research Unit of Internal Medicine, University of Oulu, Oulu, FI-90014, Finland

    • Tapani Ebeling
  105. Division of Epidemiology, Department of Medicine, Institute for Medicine and Public Health, Vanderbilt Genetics Institute, Vanderbilt University, Nashville, Tennessee 37203, USA

    • Todd L. Edwards
    •  & Ayush Giri
  106. Massachusetts General Hospital, Boston, Massachusetts 02114, USA

    • Patrick T. Ellinor
    • , Jose C. Florez
    • , Paul L. Huang
    • , Steven A. Lubitz
    • , Alisa K. Manning
    • , Aniruddh P. Patel
    • , Gina M. Peloso
    • , Suthesh Sivapalaratnam
    •  & Sekar Kathiresan
  107. Medical and Population Genetics Program, Broad Institute, Cambridge, Massachusetts 02141, USA

    • Patrick T. Ellinor
    • , Jose C. Florez
    • , Steven A. Lubitz
    • , Alisa K. Manning
    •  & Gina M. Peloso
  108. Department of Epidemiology and Biostatistics, MRC-PHE Centre for Environment and Health, School of Public Health, Imperial College London, London W2 1PG, UK

    • Paul Elliott
  109. Department of Hygiene and Epidemiology, University of Ioannina Medical School, Ioannina, 45110, Greece

    • Evangelos Evangelou
  110. Survey Research Center, Institute for Social Research, University of Michigan, Ann Arbor, Michigan 48104, USA

    • Jessica D. Faul
  111. Division of Statistical Genomics, Department of Genetics, Washington University School of Medicine, StLouis, Missouri 63108, USA

    • Mary F. Feitosa
    •  & Ingrid B. Borecki
  112. CNR Institute of Clinical Physiology, Pisa, Italy

    • Ele Ferrannini
  113. Department of Clinical & Experimental Medicine, University of Pisa, Pisa, Italy

    • Ele Ferrannini
  114. Research Center on Epidemiology and Preventive Medicine, Department of Clinical and Experimental Medicine, University of Insubria, Varese, 21100, Italy

    • Marco M. Ferrario
  115. Toulouse University School of Medicine, Toulouse, TSA 50032 31059, France

    • Jean Ferrieres
  116. Department of Medicine, Harvard University Medical School, Boston, Massachusetts 02115, USA

    • Jose C. Florez
    •  & Alisa K. Manning
  117. University of Glasgow, Glasgow G12 8QQ, UK

    • Ian Ford
    • , Chris J. Packard
    • , Sandosh Padmanabhan
    • , Naveed Sattar
    •  & Robin Young
  118. Institute of Molecular Medicine, The University of Texas Health Science Center at Houston, Houston, Texas 77030, USA

    • Myriam Fornage
    •  & Li-An Lin
  119. Department of Clinical Sciences, Genetic and Molecular Epidemiology Unit, Lund University, Malmö, SE-20502, Sweden

    • Paul W. Franks
    • , Frida Renström
    •  & Tibor V. Varga
  120. Department of Nutrition, Harvard School of Public Health, Boston, Massachusetts 02115, USA

    • Paul W. Franks
  121. Department of Public Health and Clinical Medicine, Unit of Medicine, Umeå University, Umeå, 901 87, Sweden

    • Paul W. Franks
  122. Department of Clinical Biochemistry, Rigshospitalet, Copenhagen University Hospital, Copenhagen, 2100, Denmark

    • Ruth Frikke-Schmidt
    •  & Anne Tybjaerg-Hansen
  123. Department of Medical Sciences, University of Trieste, Trieste, 34137, Italy

    • Ilaria Gandin
    • , Paolo Gasparini
    • , Giorgia Girotto
    •  & Anna Morgan
  124. Division of Experimental Genetics, Sidra Medical and Research Center, Doha, 26999, Qatar

    • Paolo Gasparini
    • , Giorgia Girotto
    •  & Diego Vozzi
  125. Geriatrics, Department of Public Health, Uppsala University, Uppsala, 751 85, Sweden

    • Vilmantas Giedraitis
  126. Carolina Population Center, University of North Carolina, Chapel Hill, North Carolina 27514, USA

    • Penny Gordon-Larsen
    •  & Kathleen Mullan Harris
  127. Department of Nutrition, Gillings School of Global Public Health, University of North Carolina, Chapel Hill, North Carolina 27514, USA

    • Penny Gordon-Larsen
  128. Faculty of Medicine, University of Iceland, Reykjavik, 101, Iceland

    • Vilmundur Gudnason
    • , Albert Vernon Smith
    • , Unnur Thorsteinsdottir
    •  & Kari Stefansson
  129. Icelandic Heart Association, Kopavogur, 201, Iceland

    • Vilmundur Gudnason
    •  & Albert Vernon Smith
  130. Department of Medical Sciences, Molecular Epidemiology and Science for Life Laboratory, Uppsala University, Uppsala, 751 41, Sweden

    • Stefan Gustafsson
    •  & Erik Ingelsson
  131. Department of Sociology, University of North Carolina, Chapel Hill, North Carolina 27514, USA

    • Kathleen Mullan Harris
  132. Laboratory of Epidemiology and Population Sciences, National Institute on Aging, Intramural Research Program, National Institutes of Health, Bethesda, Maryland 20892, USA

    • Tamara B. Harris
  133. University of Exeter Medical School, University of Exeter, Exeter EX2 5DW, UK

    • Andrew T. Hattersley
  134. MRCHGU, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh EH4 2XU, UK

    • Caroline Hayward
    • , Jonathan Marten
    •  & Veronique Vitart
  135. Biodemography of Aging Research Unit, Social Science Research Institute, Duke University, Durham, North Carolina 27708, USA

    • Liang He
  136. Department of Public Health, University of Helsinki, Helsinki, FI-00014, Finland

    • Liang He
    • , Kauko Heikkilä
    •  & Jaakko Kaprio
  137. Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, FI-00014, Finland

    • Kauko Heikkilä
    •  & Jaakko Kaprio
  138. Department of Pediatrics, Haukeland University Hospital, Bergen, 5021, Norway

    • Øyvind Helgeland
    •  & Pål R. Njølstad
  139. KG Jebsen Center for Diabetes Research, Department of Clinical Science, University of Bergen, Bergen, 5020, Norway

    • Øyvind Helgeland
    • , Stefan Johansson
    •  & Pål R. Njølstad
  140. Department of Cardiology, Heart Center, Tampere University Hospital, Tampere, 33521, Finland

    • Jussi Hernesniemi
  141. Department of Clinical Chemistry, Fimlab Laboratories, Tampere, 33520, Finland

    • Jussi Hernesniemi
    • , Terho Lehtimäki
    •  & Leo-Pekka Lyytikäinen
  142. Department of Clinical Chemistry, University of Tampere School of Medicine, Tampere, 33014, Finland

    • Jussi Hernesniemi
    • , Terho Lehtimäki
    •  & Leo-Pekka Lyytikäinen
  143. Centre for Eye Research Australia, Royal Victorian Eye and Ear Hospital, University of Melbourne, Melbourne, Victoria, 3002, Australia

    • Alex W. Hewitt
  144. Centre for Ophthalmology and Vision Science, Lions Eye Institute, University of Western Australia, Perth, Western Australia, 6009, Australia

    • Alex W. Hewitt
    •  & David A. Mackey
  145. Menzies Research Institute Tasmania, University of Tasmania, Hobart, Tasmania, 7000, Australia

    • Alex W. Hewitt
  146. Generation Scotland, Centre for Genomic and Experimental Medicine, University of Edinburgh, Edinburgh EH4 2XU, UK

    • Lynne J. Hocking
  147. Musculoskeletal Research Programme, Division of Applied Medicine, University of Aberdeen, Aberdeen, AB25 2ZD, UK

    • Lynne J. Hocking
  148. K.G. Jebsen Center for Genetic Epidemiology, Department of Public Health, NTNU, Norwegian University of Science and Technology, Trondheim, 7600, Norway

    • Oddgeir L. Holmen
  149. AMC, Department of Vascular Medicine, Amsterdam, 1105 AZ, The Netherlands

    • G. Kees Hovingh
  150. HUNT Research Centre, Department of Public Health and General Practice, Norwegian University of Science and Technology, Levanger, 7600, Norway

    • Kristian Hveem
  151. Department of Neurology, Erasmus Medical Center, Rotterdam, 3015 GE, The Netherlands

    • M. Arfan Ikram
  152. Department of Radiology, Erasmus Medical Center, Rotterdam, 3015 GE, The Netherlands

    • M. Arfan Ikram
  153. Department of Medicine, Division of Cardiovascular Medicine, Stanford University School of Medicine, Stanford, California 94305, USA

    • Erik Ingelsson
  154. Department of Public Health & Clinical Medicine, Umeå University, Umeå, SE-90185, Sweden

    • Jan-Håkan Jansson
    •  & Olov Rolandsson
  155. Research Unit Skellefteå, Skellefteå, SE-93141, Sweden

    • Jan-Håkan Jansson
  156. Department of Genome Sciences, University of Washington, Seattle, Washington 98195, USA

    • Gail P. Jarvik
  157. The Copenhagen City Heart Study, Frederiksberg Hospital, Frederiksberg, 2000, Denmark

    • Gorm B. Jensen
  158. Department of Preventive Medicine, Keck School of Medicine of the University of California, Los Angeles, California 90089, USA

    • Xuejuan Jiang
  159. USC Roski Eye Institute, Department of Ophthalmology, Keck School of Medicine of the University of Southern California, Los Angeles, California 90089, USA

    • Xuejuan Jiang
    •  & Rohit Varma
  160. Center for Medical Genetics and Molecular Medicine, Haukeland University Hospital, Bergen, 5021, Norway

    • Stefan Johansson
  161. National Institute of Public Health, University of Southern Denmark, Copenhagen, 1353, Denmark

    • Marit E. Jørgensen
  162. Steno Diabetes Center, Gentofte, 2820, Denmark

    • Marit E. Jørgensen
  163. Aalborg University, Aalborg, DK-9000, Denmark

    • Torben Jørgensen
  164. Research Center for Prevention and Health, Capital Region of Denmark, Glostrup, DK-2600, Denmark

    • Torben Jørgensen
    • , Allan Linneberg
    •  & Betina H. Thuesen
  165. National Institute for Health and Welfare, Helsinki, FI-00271, Finland

    • Pekka Jousilahti
    • , Jaakko Kaprio
    • , Jukka Kontto
    • , Jaana Lindström
    • , Satu Männistö
    • , Markus Perola
    • , Veikko Salomaa
    •  & Jaakko Tuomilehto
  166. Department of Cardiology, Leiden University Medical Center, Leiden, 2333, The Netherlands

    • J. Wouter Jukema
    •  & Stella Trompet
  167. The Interuniversity Cardiology Institute of the Netherlands, Utrecht, 2333, The Netherlands

    • J. Wouter Jukema
  168. Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, Michigan 48109, USA

    • Bratati Kahali
    • , Elizabeth K. Speliotes
    • , Wei Zhou
    •  & Cristen J. Willer
  169. Department of Internal Medicine, University of Michigan, Ann Arbor, Michigan 48109, USA

    • Bratati Kahali
    • , Elizabeth K. Speliotes
    • , Wei Zhou
    •  & Cristen J. Willer
  170. Division of Gastroenterology, University of Michigan, Ann Arbor, Michigan 48109, USA

    • Bratati Kahali
    •  & Elizabeth K. Speliotes
  171. Department of Psychiatry, Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht, 3584 CG, The Netherlands

    • René S. Kahn
    •  & Roel A. Ophoff
  172. Department of Clinical Physiology, University of Tampere School of Medicine, Tampere, 33014, Finland

    • Mika Kähönen
  173. Echinos Medical Centre, Echinos, Greece

    • Maria Karaleftheri
  174. Oxford Centre for Diabetes, Endocrinology and Metabolism, Radcliffe Department of Medicine, University of Oxford, Oxford OX3 7LE, UK

    • Fredrik Karpe
    • , Matt Neville
    • , Katharine R. Owen
    • , Neil Robertson
    •  & Mark I. McCarthy
  175. Oxford NIHR Biomedical Research Centre, Oxford University Hospitals Trust, Oxford OX3 7LE, UK

    • Fredrik Karpe
    • , Matt Neville
    • , Katharine R. Owen
    •  & Mark I. McCarthy
  176. UKCRC Centre of Excellence for Public Health Research, Queens University Belfast, Belfast BT12 6BJ, UK

    • Frank Kee
  177. Netherlands Cancer Institute - Antoni van Leeuwenhoek hospital, Amsterdam, 1066 CX, The Netherlands

    • Renske Keeman
    •  & Marjanka K. Schmidt
  178. Department of Restorative Dentistry, Periodontology and Endodontology, University Medicine Greifswald, Greifswald, 17475, Germany

    • Thomas Kocher
  179. Foundation for Research in Health Exercise and Nutrition, Kuopio Research Institute of Exercise Medicine, Kuopio, 70100, Finland

    • Pirjo Komulainen
    • , Timo A. Lakka
    •  & Rainer Rauramaa
  180. National Heart and Lung Institute, Imperial College London, Hammersmith Hospital Campus, London W12 0NN, UK

    • Jaspal S. Kooner
  181. Division of Public Health Sciences, Fred Hutchinson Cancer Research Center, Seattle, Washington 98109, USA

    • Charles Kooperberg
    •  & Alex P. Reiner
  182. German Center for Diabetes Research, München-Neuherberg, 85764, Germany

    • Jennifer Kriebel
    • , Karina Meidtner
    •  & Matthias B. Schulze
  183. Institute of Epidemiology II, Helmholtz Zentrum München - German Research Center for Environmental Health, Neuherberg, D-85764, Germany

    • Jennifer Kriebel
  184. Research Unit of Molecular Epidemiology, Helmholtz Zentrum München—German Research Center for Environmental Health, Neuherberg, D-85764, Germany

    • Jennifer Kriebel
  185. Department of Psychiatry, and Division of Molecular Biology and Human Genetics, Department of Biomedical Sciences, Faculty of Medicine and Health Sciences, Stellenbosch University, Tygerberg, Western Cape, 7505, South Africa

    • Helena Kuivaniemi
  186. CHU Nantes, Service de Génétique Médicale, Nantes, 44093, France

    • Sébastien Küry
  187. Institute of Clinical Medicine, Internal Medicine, University of Eastern Finland and Kuopio University Hospital, Kuopio, 70210, Finland

    • Johanna Kuusisto
    •  & Markku Laakso
  188. Institute for Maternal and Child Health - IRCCS ‘Burlo Garofolo’, Trieste, 34137, Italy

    • Martina La Bianca
    • , Antonietta Robino
    •  & Sheila Ulivi
  189. Institute of Biomedicine & Physiology, University of Eastern Finland, Kuopio, 70210, Finland

    • Timo A. Lakka
  190. Department of Genetics, University of North Carolina, Chapel Hill, North Carolina 27514, USA

    • Ethan M. Lange
    • , Leslie A. Lange
    • , Karen L. Mohlke
    •  & Ying Wu
  191. Department of Biostatistical Sciences and Center for Public Health Genomics, Wake Forest School of Medicine, Winston-Salem, North Carolina 27157, USA

    • Carl D. Langefeld
  192. MRC Epidemiology Unit, University of Cambridge School of Clinical Medicine, Institute of Metabolic Science, Cambridge CB2 0QQ, UK

    • Claudia Langenberg
    • , Jian'an Luan
    • , John R. B. Perry
    • , Robert A. Scott
    • , Nicholas J. Wareham
    • , Sara M. Willems
    •  & Jing Hua Zhao
  193. Group Health Research Institute, Seattle, Washington 98101, USA

    • Eric B. Larson
  194. Department of Health Services, University of Washington, Seattle, Washington 98101, USA

    • Eric B. Larson
  195. Division of Endocrinology and Metabolism, Department of Internal Medicine, Taichung Veterans General Hospital, Taichung 407, Taiwan

    • I-Te Lee
  196. School of Medicine, National Yang-Ming University, Taipei 112, Taiwan

    • I-Te Lee
  197. School of Medicine, Chung Shan Medical University, Taichung 402, Taiwan

    • I-Te Lee
  198. Division of Preventive Medicine University of Alabama at Birmingham, Birmingham, Alabama 35205, USA

    • Cora E. Lewis
  199. Key Laboratory of Nutrition and Metabolism, Institute for Nutritional Sciences, Shanghai Institutes for Biological Sciences, Chinese Academy of Sciences, University of the Chinese Academy of Sciences, Shanghai, People’s Republic of China, Shanghai, 200031, China

    • Huaixing Li
    • , Xu Lin
    • , Feijie Wang
    • , Yiqin Wang
    • , Pang Yao
    •  & He Zheng
  200. Department of Medicine, Division of Cardiovascular Medicine, Stanford University School of Medicine, Palo Alto, California 94304, USA

    • Jin Li
  201. Department of Medicine, Boston University School of Medicine, Boston, Massachusetts 02118, USA

    • Honghuang Lin
  202. Uppsala University, Uppsala, 75185, Sweden

    • Lars Lind
  203. Department of Experimental Medicine, Rigshospitalet, Copenhagen, DK-2200, Denmark

    • Allan Linneberg
  204. Division of Public Health Sciences, Wake Forest School of Medicine, Winston-Salem, North Carolina 27157, USA

    • Yongmei Liu
  205. Division of Health Sciences, Warwick Medical School, Warwick University, Coventry CV4 7AL, UK

    • Artitaya Lophatananon
  206. Department of Psychiatry, Washington University, Saint Louis, Missouri 63110, USA

    • Pamela A. F. Madden
  207. Department of Molecular Epidemiology, German Institute of Human Nutrition Potsdam-Rehbruecke (DIfE), Nuthetal, 14558, Germany

    • Karina Meidtner
    •  & Matthias B. Schulze
  208. Westmead Millennium Institute of Medical Research, Centre for Vision Research and Department of Ophthalmology, University of Sydney, Sydney, New South Wales, 2022, Australia

    • Paul Mitchell
  209. Department of Public Health and Primary Care, Leiden University Medical Center, Leiden, 2300 RC, The Netherlands

    • Dennis O. Mook-Kanamori
  210. Centre for Global Health Research, Usher Institute of Population Health Sciences and Informatics, University of Edinburgh, Edinburgh EH8 9AG, UK

    • Andrew D. Morris
    • , Ozren Polasek
    •  & Igor Rudan
  211. Department of Medicine I, Ludwig-Maximilians-Universität, Munich, 81377, Germany

    • Martina Müller-Nurasyid
  212. DZHK (German Centre for Cardiovascular Research), partner site Munich Heart Alliance, Munich, 80802, Germany

    • Martina Müller-Nurasyid
  213. Laboratory of Neurogenetics, National Institute on Aging, NIH, Bethesda, Maryland 20892, USA

    • Mike A. Nalls
  214. DZHK (German Centre for Cardiovascular Research), partner site Greifswald, Greifswald, 17475, Germany

    • Matthias Nauck
  215. Institute of Clinical Chemistry and Laboratory Medicine, University Medicine Greifswald, Greifswald, 17475, Germany

    • Matthias Nauck
  216. Department of Cardiology, Heart Center, Tampere University Hospital and School of Medicine, University of Tampere, Tampere, 33521, Finland

    • Kjell Nikus
  217. Program in Personalized Medicine, Department of Medicine, University of Maryland School of Medicine, Baltimore, Maryland 21201, USA

    • Jeffrey R. O'Connel
    • , James A. Perry
    •  & Laura M. Yerges-Armstrong
  218. Department of Medicine, Tampere University Hospital, Tampere, 33521, Finland

    • Heikki Oksa
  219. Center for Neurobehavioral Genetics, UCLA, Los Angeles, California 90095, USA

    • Loes M. Olde Loohuis
    •  & Roel A. Ophoff
  220. Pat Macpherson Centre for Pharmacogenetics and Pharmacogenomics, Medical Research Institute, Ninewells Hospital and Medical School, Dundee DD1 9SY, UK

    • Colin N. A. Palmer
  221. Laboratory of Clinical Chemistry and Hematology, Division Laboratories and Pharmacy, University Medical Center Utrecht, Utrecht, 3584 CX, The Netherlands

    • Gerard Pasterkamp
  222. Laboratory of Experimental Cardiology, Division Heart & Lungs, University Medical Center Utrecht, Utrecht, 3584 CX, The Netherlands

    • Gerard Pasterkamp
    •  & Sander W. van der Laan
  223. School of Women’s and Infants’ Health, The University of Western Australia, Perth, Western Australia, 6009, Australia

    • Craig E. Pennell
    •  & Carol A. Wang
  224. University of Helsinki, Institute for Molecular Medicine (FIMM) and Diabetes and Obesity Research Program, Helsinki, FI00014, Finland

    • Markus Perola
  225. University of Tartu, Estonian Genome Center, Tartu, Estonia, Tartu, 51010, Estonia

    • Markus Perola
  226. School of Medicine, University of Split, Split, 21000, Croatia

    • Ozren Polasek
  227. Center for Neurogenomics and Cognitive Research, Department Complex Trait Genetics, VU University, Amsterdam, 1081 HV, The Netherlands

    • Danielle Posthuma
  228. Neuroscience Campus Amsterdam, Department Clinical Genetics, VU Medical Center, Amsterdam, 1081 HV, The Netherlands

    • Danielle Posthuma
  229. Department of Clinical Physiology and Nuclear Medicine, Turku University Hospital, Turku, 20521, Finland

    • Olli T. Raitakari
  230. Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku, 20520, Finland

    • Olli T. Raitakari
  231. Centre for Non-Communicable Diseases, Karachi, Pakistan

    • Asif Rasheed
    •  & Danish Saleheen
  232. Department of Clinical Physiology and Nuclear Medicine, Kuopio University Hospital, Kuopio, 70210, Finland

    • Rainer Rauramaa
  233. MRL, Merck & Co., Inc., Genetics and Pharmacogenomics, Boston, Massachusetts 02115, USA

    • Dermot F. Reilly
  234. Department of Epidemiology, University of Washington, Seattle, Washington 98195, USA

    • Alex P. Reiner
  235. Department of Biobank Research, Umeå University, Umeå, SE-90187, Sweden

    • Frida Renström
  236. Division of Cardiovascular Medicine, Brigham and Women’s Hospital and Harvard Medical School, Boston, Massachusetts 02115, USA

    • Paul M. Ridker
  237. Department of Medicine, Faculty of Medicine, Université de Montréal, Montreal, Quebec, H3T 1J4, Canada

    • John D. Rioux
    • , Jean-Claude Tardif
    •  & Guillaume Lettre
  238. Department of Public Health and Clinical Medicine, Unit of Family Medicine, Umeå University, Umeå, 90185, Sweden

    • Olov Rolandsson
  239. Department of Biostatistics and Epidemiology, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania 19104, USA

    • Danish Saleheen
    •  & Wei Zhao
  240. Division of Epidemiology & Community Health University of Minnesota, Minneapolis, Minnesota 55454, USA

    • Pamela J. Schreiner
  241. Duke University, Durham, North Carolina 27703, USA

    • Svati Shah
  242. Saw Swee Hock School of Public Health, National University of Singapore, National University Health System, Singapore, Singapore

    • Xueling Sim
  243. Departement of Haematology, University of Cambridge, Cambridge CB2 OPT, UK

    • Suthesh Sivapalaratnam
  244. Department of Vascular Medicine, AMC, Amsterdam, 1105 AZ, The Netherlands

    • Suthesh Sivapalaratnam
  245. Department of Twin Research and Genetic Epidemiology, Kingís College London, London SE1 7EH, UK

    • Kerrin S. Small
    •  & Timothy D. Spector
  246. Alzheimer Scotland Dementia Research Centre, University of Edinburgh, Edinburgh EH8 9JZ, UK

    • John M. Starr
  247. Department of Epidemiology and Biostatistics, VU University Medical Center, Amsterdam, 1007MB, The Netherlands

    • Leen M. ‘t Hart
    •  & Natasja M. van Schoor
  248. Department of Molecular Cell Biology, Leiden University Medical Center, Leiden, 2333ZC, The Netherlands

    • Leen M. ‘t Hart
  249. Department of Molecular Epidemiology, Leiden University Medical Center, Leiden, 2333ZC, The Netherlands

    • Leen M. ‘t Hart
  250. College of Biomedical and Life Sciences, Cardiff University, Cardiff, CF14 4EP, UK

    • Katherine E. Tansey
  251. MRC Integrative Epidemiology Unit, School of Social and Community Medicine, University of Bristol, Bristol BS8 2BN, UK

    • Katherine E. Tansey
  252. Institute for Community Medicine, University Medicine Greifswald, Greifswald, 17475, Germany

    • Alexander Teumer
  253. Center for Pediatric Research, Department for Women’s and Child Health, University of Leipzig, Leipzig, 04103, Germany

    • Anke Tönjes
  254. Division of Molecular Biology and Human Genetics, Department of Biomedical Sciences, Faculty of Medicine and Health Sciences, Stellenbosch University, Tygerberg, Western Cape, 7505, South Africa

    • Gerard Tromp
  255. Department of Gerontology and Geriatrics, Leiden University Medical Center, Leiden, 2333, The Netherlands

    • Stella Trompet
  256. Anogia Medical Centre, Anogia, Greece

    • Emmanouil Tsafantakis
  257. Centre for Vascular Prevention, Danube-University Krems, Krems, 3500, Austria

    • Jaakko Tuomilehto
  258. Dasman Diabetes Institute, Dasman, 15462, Kuwait

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

    • Jaakko Tuomilehto
  260. Department of Psychiatry, Dalhousie University, Halifax B3H 4R2, Canada

    • Rudolf Uher
  261. University of Amsterdam, Department of Brain & Cognition, Amsterdam, 1018 WS, The Netherlands

    • Andries R. Van Der Leij
  262. Department of Obstetrics and Gynecology, Institute for Medicine and Public Health, Vanderbilt Genetics Institute, Vanderbilt University, Nashville, Tennessee 37203, USA

    • Digna R. Velez Edwards
  263. MRL, Merck & Co., Inc., Cardiometabolic Disease, Kenilworth, New Jersey 07033, USA

    • Thomas F. Vogt
  264. Institute of Cellular Medicine, The Medical School, Newcastle University, Newcastle NE2 4HH, UK

    • Mark Walker
  265. Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts 02118, USA

    • Shuai Wang
  266. Departments of Epidemiology & Medicine, Diabetes Translational Research Center, Fairbanks School of Public Health & School of Medicine, Indiana University, Indiana 46202, USA

    • Jennifer Wessel
  267. Department of Physiology and Biophysics, University of Mississippi Medical Center, Jackson, Mississippi 39216, USA

    • James G. Wilson
  268. Danish Diabetes Academy, Odense, 5000, Denmark

    • Daniel R. Witte
  269. Department of Public Health, Aarhus University, Aarhus, 8000, Denmark

    • Daniel R. Witte
  270. Memorial University, Faculty of Medicine, Discipline of Genetics, St. John’s, Newfoundland A1B 3V6, Canada

    • Michael O. Woods
  271. GlaxoSmithKlein, King of Prussia, Pennsylvania 19406, USA

    • Laura M. Yerges-Armstrong
  272. Department of Clinical Sciences, Quantitative Biomedical Research Center, Center for the Genetics of Host Defense, University of Texas Southwestern Medical Center, Dallas, Texas 75390, USA

    • Xiaowei Zhan
  273. Department of Human Genetics, University of Michigan, Ann Arbor, Michigan 48109, USA

    • Cristen J. Willer
  274. Department of Public Health Sciences, Institute for Personalized Medicine, the Pennsylvania State University College of Medicine, Hershey, Pennsylvania 17033, USA

    • Dajiang J. Liu
  275. Department of Epidemiology and Carolina Center of Genome Sciences, Chapel Hill, North Carolina 27514, USA

    • Kari E. North
  276. Department of Neurology, Boston University School of Medicine, Boston, Massachusetts 02118, USA

    • Nancy L. Heard-Costa
  277. Department of Epidemiology Research, Statens Serum Institut, Copenhagen, 2200, Denmark

    • Tune H. Pers
  278. Li Ka Shing Centre for Health Information and Discovery, The Big Data Institute, University of Oxford, Oxford OX3 7BN, UK

    • Cecilia M. Lindgren
  279. The Mindich Child Health and Development Institute, Ichan School of Medicine at Mount Sinai, New York, New York 10069, USA

    • Ruth J. F. Loos
  280. Departments of Pediatrics and Genetics, Harvard Medical School, Boston, Massachusetts 02115, USA

    • Joel N. Hirschhorn
  281. Princess Al-Jawhara Al-Brahim Centre of Excellence in Research of Hereditary Disorders (PACER-HD), King Abdulaziz University, Jeddah, 21589, Saudi Arabia

    • Panos Deloukas

Consortia

  1. The EPIC-InterAct Consortium

  2. CHD Exome+ Consortium

  3. ExomeBP Consortium

  4. T2D-Genes Consortium

  5. GoT2D Genes Consortium

  6. Global Lipids Genetics Consortium

  7. ReproGen Consortium

  8. MAGIC Investigators

    A list of members and their affiliations appears in the Supplementary Information.

Authors

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Contributions

Writing group (wrote and edited manuscript): P.D., T.M.F., M.Gr., J.N.H., G.L., K.S.L., Y.Lu., E.M., C.M.-G., F.Ri. All authors contributed and discussed the results, and commented on the manuscript. Data preparation group (checked and prepared data from contributing cohorts for meta-analyses and replication): T.Es., M.Gr., H.M.H., A.E.J., T.Ka., K.S.L., A.E.L., Y.Lu., E.M., N.G.D.M., C. M.-G., P.Mu., M.C.Y.N., M.A.R., C.S., K.St., V.T., S.V., T.W.W., K.L.Y. This work was done under the auspices of the GIANT, CHARGE, BBMRI, UK ExomeChip, and GOT2D consortia. Height meta-analyses (discovery and replication, single-variant and gene-based): P.D., T.M.F., M.Gr., J.N.H., G.L., D.J.L., K.S.L., Y.Lu, E.M., C.M.-G., F.Ri., A.R.W. UK Biobank-based integration of height association signals group and heritability analyses: P.D., T.M.F., G.L., Z.K., K.S.L., E.M., S.R., A.R.W. Pleiotropy working group: G.A., M. Bo., J.P.C., P.D., F.D., J.C.F., H.M.H., S. Kat., C.M.L., D.J.L., R.J.F.L., A.Ma., E.M., M.I.M., P.B.M., G.M.P., J.R.B.P., K.S.R., C.J.W. Biological and clinical enrichment and pathway analyses: R.S.F., J.N.H., Z.K., D.L., G.L., K.S.L., T.H.P. Functional characterization of STC2: T.R.K., C.O.

Competing interests

The authors declare no competing financial interests.

Corresponding authors

Correspondence to Joel N. Hirschhorn or Panos Deloukas or Guillaume Lettre.

Reviewer Information Nature thanks J. Barrett, D. Hinds and D. Hunter for their contribution to the peer review of this work.

Extended data

Supplementary information

PDF files

  1. 1.

    Supplementary Information

    This file contains Supplementary Text and Data and additional references.

Excel files

  1. 1.

    Supplementary Tables

    This file contains Supplementary Tables 1-24.

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