Article | Published:

Genome-wide trans-ancestry meta-analysis provides insight into the genetic architecture of type 2 diabetes susceptibility

Nature Genetics volume 46, pages 234244 (2014) | Download Citation

Abstract

To further understanding of the genetic basis of type 2 diabetes (T2D) susceptibility, we aggregated published meta-analyses of genome-wide association studies (GWAS), including 26,488 cases and 83,964 controls of European, east Asian, south Asian and Mexican and Mexican American ancestry. We observed a significant excess in the directional consistency of T2D risk alleles across ancestry groups, even at SNPs demonstrating only weak evidence of association. By following up the strongest signals of association from the trans-ethnic meta-analysis in an additional 21,491 cases and 55,647 controls of European ancestry, we identified seven new T2D susceptibility loci. Furthermore, we observed considerable improvements in the fine-mapping resolution of common variant association signals at several T2D susceptibility loci. These observations highlight the benefits of trans-ethnic GWAS for the discovery and characterization of complex trait loci and emphasize an exciting opportunity to extend insight into the genetic architecture and pathogenesis of human diseases across populations of diverse ancestry.

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Acknowledgements

Funding for the research undertaken in this study has been received from the following: the Canadian Institutes of Health Research; the European Commission (ENGAGE FP7 HEALTH-F4-2007- 201413); the Medical Research Council UK (G0601261); the Mexico Convocatoria (SSA/IMMS/ISSSTE-CONACYT 2012-2, clave 150352, IMSS R-2011-785-018 and CONACYT Salud-2007-C01-71068); the US National Institutes of Health (DK062370, HG000376, DK085584, DK085545, DK073541 and DK085501); and the Wellcome Trust (WT098017, WT090532, WT090367, WT098381, WT081682 and WT085475). We acknowledge the many colleagues who contributed to collection and phenotypic characterization of the clinical samples and the genotyping and analysis of the GWAS data, full details of which are provided in the contributing consortia papers5,11,13,15. We also thank those individuals who agreed to participate in this study.

Author information

Author notes

    • Leena Peltonen
    •  & Steven Wiltshire

    Deceased.

    • Anubha Mahajan
    • , Min Jin Go
    • , Weihua Zhang
    • , Jennifer E Below
    •  & Kyle J Gaulton

    These authors contributed equally to this work.

    • David Altshuler
    • , Donald W Bowden
    • , Yoon Shin Cho
    • , Nancy J Cox
    • , Miguel Cruz
    • , Craig L Hanis
    • , Jaspal Kooner
    • , Jong-Young Lee
    • , Mark Seielstad
    • , Yik Ying Teo
    • , Michael Boehnke
    • , Esteban J Parra
    • , John C Chambers
    • , E Shyong Tai
    • , Mark I McCarthy
    •  & Andrew P Morris

    These authors jointly directed this work.

Affiliations

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

    • Anubha Mahajan
    • , Kyle J Gaulton
    • , Teresa Ferreira
    • , Momoko Horikoshi
    • , Inga Prokopenko
    • , Antigone S Dimas
    • , Peter J Donnelly
    • , Asa K Hedman
    • , Erik Ingelsson
    • , Ashish Kumar
    • , Vasiliki Lagou
    • , Cecilia M Lindgren
    • , Ghazala Mirza
    • , John R B Perry
    • , N William Rayner
    • , Neil Robertson
    • , Joseph Trakalo
    • , Steven Wiltshire
    • , Mark I McCarthy
    •  & Andrew P Morris
  2. Center for Genome Science, National Institute of Health, Osong Health Technology Administration Complex, Chungcheongbuk-do, Cheongwon-gun, Gangoe-myeon, Yeonje-ri, Korea.

    • Min Jin Go
    • , Bok-Ghee Han
    • , Young Jin Kim
    •  & Jong-Young Lee
  3. Epidemiology and Biostatistics, Imperial College London, London, UK.

    • Weihua Zhang
    • , Paul Elliott
    •  & John C Chambers
  4. Department of Genome Sciences, University of Washington, Seattle, Washington, USA.

    • Jennifer E Below
  5. Oxford Centre for Diabetes, Endocrinology and Metabolism, University of Oxford, UK.

    • Momoko Horikoshi
    • , Inga Prokopenko
    • , Christopher J Groves
    • , Neelam Hassanali
    • , Prasad Katulanda
    • , David R Matthews
    • , Katharine R Owen
    • , N William Rayner
    • , Neil Robertson
    •  & Mark I McCarthy
  6. National Heart, Lung and Blood Institute's Framingham Heart Study, Framingham, Massachusetts, USA.

    • Andrew D Johnson
    • , Josée Dupuis
    •  & Caroline Fox
  7. Center for Genomics and Personalized Medicine Research, Wake Forest School of Medicine, Winston-Salem, North Carolina, USA.

    • Maggie C Y Ng
    •  & Donald W Bowden
  8. Center for Diabetes Research, Wake Forest School of Medicine, Winston-Salem, North Carolina, USA.

    • Maggie C Y Ng
    •  & Donald W Bowden
  9. Genomics of Common Disease, Imperial College London, Hammersmith Hospital, London, UK.

    • Inga Prokopenko
    •  & Philippe Froguel
  10. Department of Public Health and Primary Care, University of Cambridge, Cambridge, UK.

    • Danish Saleheen
    • , John Danesh
    • , Hassan Khan
    • , Kay-Tee Khaw
    •  & Robin Young
  11. Center for Non-Communicable Diseases Pakistan, Karachi, Pakistan.

    • Danish Saleheen
    • , Philippe Frossard
    • , Asif Rasheed
    • , Maria Samuel
    •  & Nabi Shah
  12. Department of Epidemiology and Public Health, National University of Singapore, Singapore.

    • Xu Wang
    • , Kee-Seng Chia
    • , Jen-Mai Lee
    • , Peng-Keat Ng
    • , Twee Hee Ong
    • , Yik Ying Teo
    •  & E Shyong Tai
  13. Wellcome Trust Sanger Institute, Hinxton, Cambridge, UK.

    • Eleftheria Zeggini
    • , Ines Barroso
    • , Panos Deloukas
    • , Sarah Edkins
    • , Sarah E Hunt
    • , Stavroula Kanoni
    • , Cordelia Langford
    • , Leena Peltonen
    • , Simon Potter
    • , Samuli Ripatti
    • , Kerrin S Small
    •  & Kathleen Stirrups
  14. Department of Biostatistics, University of Michigan, Ann Arbor, Michigan, USA.

    • Goncalo R Abecasis
    • , Anne U Jackson
    • , Hyun Min Kang
    • , Laura J Scott
    • , Xueling Sim
    • , Heather M Stringham
    • , Tanya M Teslovich
    •  & Michael Boehnke
  15. Department of Nutrition, University of North Carolina, Chapel Hill, North Carolina, USA.

    • Linda S Adair
  16. Lund University Diabetes Centre, Department of Clinical Science Malmö, Scania University Hospital, Lund University, Malmö, Sweden.

    • Peter Almgren
    • , Mozhgan Dorkhan
    • , Leif C Groop
    • , Anna Jonsson
    • , Jasmina Kravic
    • , Eero Lindholm
    • , Valeriya Lyssenko
    • , Olle Melander
    •  & Peter M Nilsson
  17. Institute of Biomedicine, University of Eastern Finland, Kuopio Campus, Kuopio, Finland.

    • Mustafa Atalay
    •  & Timo A Lakka
  18. Singapore Eye Research Institute, Singapore National Eye Centre, Singapore.

    • Tin Aung
    • , Chiea-Chuen Khor
    • , Wan Ting Tay
    •  & Tien Yin Wong
  19. Department of Ophthalmology, National University of Singapore, Singapore.

    • Tin Aung
    •  & Tien Yin Wong
  20. Centro Cardiologico Monzino, Istituto di Ricovero e Cura a Carattere Scientifico (IRCCS), Milan, Italy.

    • Damiano Baldassarre
    • , Elena Tremoli
    •  & Fabrizio Veglia
  21. Department of Pharmacological Sciences, University of Milan, Milan, Italy.

    • Damiano Baldassarre
    •  & Elena Tremoli
  22. Institut National de la Santé et de la Recherche Médicale (INSERM) Centre de recherche en Epidémiologie et Santé des Populations (CESP) U1018, Villejuif, France.

    • Beverley Balkau
  23. University Paris Sud 11, UMRS 1018, Villejuif, France.

    • Beverley Balkau
  24. Shanghai Diabetes Institute, Shanghai Key Laboratory of Diabetes Mellitus, Department of Endocrinology and Metabolism, Shanghai Jiao Tong University Affiliated Sixth People's Hospital, Shanghai, China.

    • Yuqian Bao
    • , Cheng Hu
    • , Weiping Jia
    • , Congrong Wang
    •  & Rong Zhang
  25. College of Medical and Dental Sciences, University of Birmingham, Birmingham, UK.

    • Anthony H Barnett
    • , Ann M Kelly
    •  & Simon D Rees
  26. BioMedical Research Centre, Heart of England National Health Service (NHS) Foundation Trust, Birmingham, UK.

    • Anthony H Barnett
    • , Ann M Kelly
    •  & Simon D Rees
  27. University of Cambridge Metabolic Research Laboratories, Institute of Metabolic Science, Addenbrooke's Hospital, Cambridge, UK.

    • Ines Barroso
  28. National Institute for Health Research (NIHR) Cambridge Biomedical Research Centre, Institute of Metabolic Science, Addenbrooke's Hospital, Cambridge, UK.

    • Ines Barroso
  29. Baqai Institute of Diabetology and Endocrinology, Karachi, Pakistan.

    • Abdul Basit
    •  & Zafar I Hydrie
  30. Department of Pediatrics, Section of Genetics, College of Medicine, University of Oklahoma Health Sciences Center, Oklahoma City, Oklahoma, USA.

    • Latonya F Been
    •  & Dharambir K Sanghera
  31. Busselton Population Medical Research Institute, Sir Charles Gairdner Hospital, Nedlands, Western Australia, Australia.

    • John Beilby
    • , Jennie Hui
    • , Alan James
    •  & Bill Musk
  32. PathWest Laboratory Medicine of Western Australia, Queen Elizabeth II Medical Centre, Nedlands, Western Australia, Australia.

    • John Beilby
  33. School of Pathology and Laboratory Medicine, The University of Western Australia, Nedlands, Western Australia, Australia.

    • John Beilby
  34. Department of Medicine, University of Chicago, Chicago, Illinois, USA.

    • Graeme I Bell
    •  & Nancy J Cox
  35. Department of Human Genetics, University of Chicago, Chicago, Illinois, USA.

    • Graeme I Bell
    •  & Nancy J Cox
  36. Faculty of Medicine, University of Iceland, Reykjavík, Iceland.

    • Rafn Benediktsson
    • , Kari Stefansson
    •  & Unnur Thorsteinsdottir
  37. Landspitali University Hospital, Reykjavík, Iceland.

    • Rafn Benediktsson
    • , Astradur B Hreidarsson
    •  & Gunnar Sigurðsson
  38. Diabetes and Obesity Research Institute, Cedars-Sinai Medical Center, Los Angeles, California, USA.

    • Richard N Bergman
  39. Department of Internal Medicine, Division of Endocrinology and Diabetes, University Medical Centre Ulm, Ulm, Germany.

    • Bernhard O Boehm
  40. Lee Kong Chian (LKC) School of Medicine, Nanyang Technological University, Singapore and Imperial College London, London, UK.

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

    • Eric Boerwinkle
  42. Human Genome Sequencing Center at Baylor College of Medicine, Houston, Texas, USA.

    • Eric Boerwinkle
  43. National Human Genome Research Institute, National Institutes of Health, Bethesda, Maryland, USA.

    • Lori L Bonnycastle
    • , Peter S Chines
    •  & Francis S Collins
  44. Broad Institute of Harvard and Massachusetts Institute of Technology (MIT), Cambridge, Massachusetts, USA.

    • Noël Burtt
    • , Jason Carey
    • , Andrew T Crenshaw
    • , Pierre Fontanillas
    • , George B Grant
    • , Candace Guiducci
    • , Sekar Kathiresan
    • , Cecilia M Lindgren
    • , Melissa Parkin
    • , Leena Peltonen
    • , Soumya Raychaudhuri
    • , Elizabeth J Rossin
    • , Ayellet V Segrè
    • , Benjamin F Voight
    • , Wendy Winckler
    •  & David Altshuler
  45. Department of Medicine, Vanderbilt Epidemiology Center, Vanderbilt-Ingram Cancer Center, Vanderbilt University School of Medicine, Nashville, Tennessee, USA.

    • Qiuyin Cai
    • , Jirong Long
    • , Xiao Ou Shu
    •  & Wei Zheng
  46. Centre for Population Health Sciences, University of Edinburgh, Teviot Place, Edinburgh, UK.

    • Harry Campbell
    • , Ross M Fraser
    • , Jackie F Price
    • , Igor Rudan
    •  & James F Wilson
  47. Medical Research Council (MRC) Institute of Genetics and Molecular Medicine at the University of Edinburgh, Western General Hospital, Edinburgh, UK.

    • Harry Campbell
    • , Caroline Hayward
    • , Pau Navarro
    •  & James F Wilson
  48. Centre National de la Recherche Scientifique (CNRS)–Unité Mixte de Recherche (UMR)-8199, Institute of Biology and Lille 2 University, Pasteur Institute, Lille, France.

    • Stephane Cauchi
    • , Elodie Eury
    • , Philippe Froguel
    • , Stéphane Lobbens
    •  & Loic Yengo
  49. Clinical Pharmacology and Barts and the London Genome Centre, William Harvey Research Institute, Barts and the London School of Medicine, Queen Mary University of London, London, UK.

    • Mark Caulfield
  50. Department of Medicine and Therapeutics, Chinese University of Hong Kong, Prince of Wales Hospital, Hong Kong, China.

    • Juliana C N Chan
    • , Ronald C W Ma
    •  & Wing Yee So
  51. Institute of Biomedical Sciences, Academia Sinica, Nankang, Taipei, Taiwan.

    • Li-Ching Chang
    • , Chien-Hsiun Chen
    • , Yuan-Tsong Chen
    •  & Jer-Yuarn Wu
  52. Department of Internal Medicine, National Taiwan University Hospital, Taipei, Taiwan.

    • Tien-Jyun Chang
    • , Yi-Cheng Chang
    •  & Lee-Ming Chuang
  53. Endocrinology-Diabetology Unit, Corbeil-Essonnes Hospital, Corbeil-Essonnes, France.

    • Guillaume Charpentier
  54. School of Chinese Medicine, China Medical University, Taichung, Taiwan.

    • Chien-Hsiun Chen
    • , Fuu Jen Tsai
    •  & Jer-Yuarn Wu
  55. Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts, USA.

    • Han Chen
    • , Josée Dupuis
    •  & Ching-Ti Liu
  56. Centre for Molecular Epidemiology, National University of Singapore, Singapore.

    • Kee-Seng Chia
    • , Chiea-Chuen Khor
    • , Wei-Yen Lim
    • , Xueling Sim
    • , Chen Suo
    •  & Yik Ying Teo
  57. Department of Molecular Genetics, Madras Diabetes Research Foundation–Indian Council of Medical Research (ICMR) Advanced Centre for Genomics of Diabetes, Chennai, India.

    • Manickam Chidambaram
    • , Samuel Liju
    • , Viswanathan Mohan
    •  & Venkatesan Radha
  58. Department of Preventive Medicine, Ajou University School of Medicine, Suwon, Korea.

    • Nam H Cho
  59. Department of Internal Medicine, Seoul National University College of Medicine, Seoul, Korea.

    • Young Min Cho
    • , Soo Heon Kwak
    •  & Kyong Soo Park
  60. Graduate Institute of Clinical Medicine, National Taiwan University School of Medicine, Taipei, Taiwan.

    • Lee-Ming Chuang
    •  & Shiro Maeda
  61. Department of Nutrition and Epidemiology, Harvard School of Public Health, Boston, Massachusetts, USA.

    • Marilyn C Cornelis
    • , Rob M van Dam
    • , Frank B Hu
    • , David J Hunter
    • , Peter Kraft
    • , Lu Qi
    •  & Qi Sun
  62. Collaborative Studies Coordinating Center, Department of Biostatistics, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, USA.

    • David J Couper
  63. Saw Swee Hock School of Public Health, National University of Singapore, Singapore.

    • Rob M van Dam
  64. Ealing Hospital NHS Trust, Middlesex, UK.

    • Debashish Das
    • , Joban Sehmi
    • , Jaspal Kooner
    •  & John C Chambers
  65. Division of Cardiovascular Epidemiology, Institute of Environmental Medicine, Karolinska Institutet, Stockholm, Sweden.

    • Ulf de Faire
    • , Bruna Gigante
    •  & Karin Leander
  66. Department of Dietetics-Nutrition, Harokopio University, Athens, Greece.

    • George Dedoussis
  67. Department of Genetic Medicine and Development, University of Geneva Medical School, Geneva, Switzerland.

    • Antigone S Dimas
    •  & Alexandra C Nica
  68. Biomedical Sciences Research Center 'Alexander Fleming', Vari, Greece.

    • Antigone S Dimas
  69. INSERM UMR 1087, Nantes, France.

    • Christian Dina
  70. CNRS UMR 6291, Nantes, France.

    • Christian Dina
  71. Nantes University, Nantes, France.

    • Christian Dina
  72. Diabetes Research Centre, Biomedical Research Institute, University of Dundee, Ninewells Hospital, Dundee, UK.

    • Alex S F Doney
    • , Andrew D Morris
    •  & Colin N A Palmer
  73. Pharmacogenomics Centre, Biomedical Research Institute, University of Dundee, Ninewells Hospital, Dundee, UK.

    • Alex S F Doney
    • , Andrew D Morris
    •  & Colin N A Palmer
  74. Department of Statistics, University of Oxford, Oxford, UK.

    • Peter J Donnelly
  75. Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands.

    • Cornelia van Duijn
    • , Albert Hofman
    •  & Andre G Uitterlinden
  76. Netherland Genomics Initiative, Netherlands Consortium for Healthy Ageing and Centre for Medical Systems Biology, Rotterdam, The Netherlands.

    • Cornelia van Duijn
    •  & Andre G Uitterlinden
  77. MRC–Health Protection Agency (HPA) Centre for Environment and Health, Imperial College London, London, UK.

    • Paul Elliott
  78. Icelandic Heart Association, Kopavogur, Iceland.

    • Valur Emilsson
    •  & Gunnar Sigurðsson
  79. Clinic of Cardiology, West German Heart Centre, University Hospital of Essen, University Duisburg-Essen, Essen, Germany.

    • Raimund Erbel
  80. Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, Finland.

    • Johan G Eriksson
    • , Satu Männistö
    • , Leena Peltonen
    • , Samuli Ripatti
    • , Veikko Salomaa
    •  & Emmi Tikkanen
  81. Department of General Practice and Primary Health Care, University of Helsinki, Helsinki, Finland.

    • Johan G Eriksson
    •  & Tom Forsen
  82. Unit of General Practice, Helsinki University General Hospital, Helsinki, Finland.

    • Johan G Eriksson
  83. Folkhälsan Research Center, Helsinki, Finland.

    • Johan G Eriksson
    • , Bo Isomaa
    •  & Tiinamaija Tuomi
  84. Unidad de Investigacion en Epidemiologia Clinica, Hospital General Regional I, Dr. Carlos MacGregor, Instituto Mexicano del Seguro Social (IMSS), Mexico City, Mexico.

    • Jorge Escobedo
  85. Estonian Genome Center, University of Tartu, Tartu, Estonia.

    • Tonu Esko
    • , Reedik Mägi
    • , Andres Metspalu
    •  & Evelin Mihailov
  86. Institute of Molecular and Cell Biology, University of Tartu, Tartu, Estonia.

    • Tonu Esko
    •  & Andres Metspalu
  87. Program in Medical and Population Genetics, Broad Institute, Cambridge, Massachusetts, USA.

    • Tonu Esko
    • , Jose C Florez
    • , Elizabeth J Rossin
    •  & Sailaja Vedantam
  88. Division of Genetics and Endocrinology, Children's Hospital, Boston, Massachusetts, USA.

    • Tonu Esko
    • , Jose C Florez
    •  & Sailaja Vedantam
  89. Department of Medicine, Harvard Medical School, Boston, Massachusetts, USA.

    • Jose C Florez
    • , James B Meigs
    • , Ayellet V Segrè
    •  & David Altshuler
  90. Center for Human Genetic Research, Massachusetts General Hospital, Boston, Massachusetts, USA.

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    • , Sekar Kathiresan
    • , Ayellet V Segrè
    •  & David Altshuler
  91. Diabetes Research Center, Diabetes Unit, Massachusetts General Hospital, Boston, Massachusetts, USA.

    • Jose C Florez
  92. MRC Epidemiology Unit, Institute of Metabolic Science, Addenbrooke's Hospital, Cambridge, UK.

    • Nita G Forouhi
    • , Claudia Langenberg
    • , Ruth J F Loos
    • , Jian'an Luan
    • , Robert A Scott
    •  & Nicholas J Wareham
  93. Vaasa Health Care Centre, Vaasa, Finland.

    • Tom Forsen
  94. Division of Endocrinology and Metabolism, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts, USA.

    • Caroline Fox
  95. Genetics of Complex Traits, Institute of Biomedical and Clinical Science, Peninsula Medical School, University of Exeter, Exeter, UK.

    • Timothy M Frayling
    • , John R B Perry
    •  & Andrew R Wood
  96. Department of Epidemiology, Shanghai Cancer Institute, Shanghai, China.

    • Yutang Gao
  97. Atherosclerosis Research Unit, Department of Medicine Solna, Karolinska Institutet, Stockholm, Sweden.

    • Karl Gertow
    • , Anders Hamsten
    • , Bengt Sennblad
    • , Angela Silveira
    •  & Rona J Strawbridge
  98. Center for Molecular Medicine, Karolinska University Hospital Solna, Stockholm, Sweden.

    • Karl Gertow
    • , Anders Hamsten
    • , Bengt Sennblad
    • , Angela Silveira
    •  & Rona J Strawbridge
  99. Institute of Genetic Epidemiology, Helmholtz Zentrum Muenchen, Neuherberg, Germany.

    • Christian Gieger
    • , Julia Meyer
    •  & Martina Müller-Nurasyid
  100. Research Unit of Molecular Epidemiology, Helmholtz Zentrum Muenchen, Neuherberg, Germany.

    • Harald Grallert
    • , Thomas Illig
    •  & Norman Klopp
  101. Clinical Cooperation Group Diabetes, Ludwig-Maximilians-Universität Muenchen and Helmholtz Zentrum Muenchen, Muenchen, Germany.

    • Harald Grallert
  102. Clinical Cooperation Group Nutrigenomics and Type 2 Diabetes, Technical University Muenchen and Helmholtz Zentrum Muenchen, Muenchen, Germany.

    • Harald Grallert
  103. German Center for Diabetes Research (DZD), Neuherberg, Germany.

    • Harald Grallert
  104. Department of Twin Research and Genetic Epidemiology, King's College London, London, UK.

    • Elin Grundberg
    •  & Kerrin S Small
  105. Department of Diabetes and Metabolic Diseases, Graduate School of Medicine, The University of Tokyo, Tokyo, Japan.

    • Kazuo Hara
    • , Takashi Kadowaki
    •  & Toshimasa Yamauchi
  106. Diabetes Genetics, Institute of Biomedical and Clinical Science, Peninsula Medical School, University of Exeter, Exeter, UK.

    • Andrew T Hattersley
  107. Institute for Clinical Diabetology, German Diabetes Center, Leibniz Center for Diabetes Research at Heinrich Heine University Düsseldorf, Düsseldorf, Germany.

    • Christian Herder
    •  & Michael Roden
  108. HUNT Research Centre, Department of Public Health and General Practice, Norwegian University of Science and Technology, Levanger, Norway.

    • Oddgeir L Holmen
    • , Kristian Hveem
    •  & Carl G P Platou
  109. Department of Vascular Medicine, Academic Medical Center, University of Amsterdam, Amsterdam, The Netherlands.

    • Kees Hovingh
    • , Suthesh Sivapalaratnam
    •  & Mieke D Trip
  110. Channing Laboratory, Department of Medicine, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts, USA.

    • Frank B Hu
    • , David J Hunter
    • , Lu Qi
    •  & Qi Sun
  111. School of Population Health, The University of Western Australia, Nedlands, Western Australia, Australia.

    • Jennie Hui
    •  & Bill Musk
  112. Institute of Cardiovascular Science, University College London, London, UK.

    • Steve E Humphries
  113. Program in Molecular and Genetic Epidemiology, Harvard School of Public Health, Boston, Massachusetts, USA.

    • David J Hunter
    •  & Peter Kraft
  114. Department of Endocrinology, Metabolism and Diabetes, Kinki University School of Medicine, Osaka-sayama, Osaka, Japan.

    • Hiroshi Ikegami
  115. Hannover Unified Biobank, Hannover Medical School, Hannover, Germany.

    • Thomas Illig
    •  & Norman Klopp
  116. Department of Medical Sciences, Molecular Epidemiology and Science for Life Laboratory, Uppsala University Hospital, Uppsala, Sweden.

    • Erik Ingelsson
  117. Department of Community Health Sciences, Aga Khan University, Karachi, Pakistan.

    • Muhammed Islam
    •  & Tazeen Jafar
  118. Department of Social Services and Health Care, Jakobstad, Finland.

    • Bo Isomaa
  119. Department of Medicine, Aga Khan University, Karachi, Pakistan.

    • Tazeen Jafar
  120. Department of Pulmonary Physiology and Sleep Medicine, West Australian Sleep Disorders Research Institute, Queen Elizabeth II Medical Centre, Hospital Avenue, Nedlands, Western Australia, Australia.

    • Alan James
  121. School of Medicine and Pharmacology, University of Western Australia, Nedlands, Western Australia, Australia.

    • Alan James
    •  & Bill Musk
  122. Institute for Medical Informatics, Biometry and Epidemiology, University Hospital of Essen, Essen, Germany.

    • Karl-Heinz Jöckel
    • , Susanne Moebus
    •  & Sonali Pechlivanis
  123. Baker International Diabetes Institute (IDI) Heart and Diabetes Institute, Melbourne, Victoria, Australia.

    • Jeremy B M Jowett
    • , Malene M Kristensen
    •  & Paul Z Zimmet
  124. Department of Epidemiology, Johns Hopkins Bloomberg School of Public Health, Baltimore, Maryland, USA.

    • Wen Hong L Kao
    •  & Man Li
  125. Cardiovascular Research Center, Massachusetts General Hospital, Boston, Massachusetts, USA.

    • Sekar Kathiresan
  126. Research Institute, National Center for Global Health and Medicine, Shinjuku-ku, Tokyo, Japan.

    • Norihiro Kato
    •  & Fumihiko Takeuchi
  127. Diabetes Research Unit, Department of Clinical Medicine, University of Colombo, Colombo, Sri Lanka.

    • Prasad Katulanda
  128. Faculty of Medicine, Institute of Health Sciences, University of Oulu, Oulu, Finland.

    • Sirkka M Keinanen-Kiukaanniemi
  129. Unit of General Practice, Oulu University Hospital, Oulu, Finland.

    • Sirkka M Keinanen-Kiukaanniemi
  130. Genome Institute of Singapore, Agency for Science, Technology and Research, Singapore.

    • Chiea-Chuen Khor
    • , Jian Jun Liu
    • , Mark Seielstad
    •  & Yik Ying Teo
  131. Department of Biochemistry, School of Medicine, Ewha Womans University, Seoul, Korea.

    • Hyung-Lae Kim
  132. School of Systems Biomedical Science, Soongsil University, Dongjak-gu, Seoul, Korea.

    • Sangsoo Kim
  133. Diabetes Prevention Unit, National Institute for Health and Welfare, Helsinki, Finland.

    • Leena Kinnunen
    •  & Jaakko Tuomilehto
  134. deCODE Genetics, Reykjavik, Iceland.

    • Augustine Kong
    • , Kari Stefansson
    • , Valgerdur Steinthorsdottir
    • , Gudmar Thorleifsson
    •  & Unnur Thorsteinsdottir
  135. South Ostrobothnia Central Hospital, Seinäjoki, Finland.

    • Eeva Korpi-Hyövälti
  136. Ministry of Health, Port Louis, Mauritius.

    • Sudhir Kowlessur
  137. Department of Anthropology, University of Toronto at Mississauga, Mississauga, Ontario, Canada.

    • S Krithika
    •  & Esteban J Parra
  138. Fundacion IMSS, Mz. Poniente, Col. Juarez, Deleg. Cuauhtemoc, Mexico City, Mexico.

    • Jesus Kumate
  139. Department of Medicine, University of Eastern Finland and Kuopio University Hospital, Kuopio, Finland.

    • Johanna Kuusisto
    • , Markku Laakso
    •  & Alena Stančáková
  140. Kuopio Research Institute of Exercise Medicine, Kuopio, Finland.

    • Timo A Lakka
    •  & Rainer Rauramaa
  141. Centre for Genetic Epidemiology and Biostatistics, The University of Western Australia, Nedlands, Western Australia, Australia.

    • Robert Lawrence
  142. Office of Population Studies Foundation Inc., University of San Carlos, Cebu City, Philippines.

    • Nanette R Lee
  143. National Heart and Lung Institute (NHLI), Imperial College London, Hammersmith Hospital, London, UK.

    • Xinzhong Li
    • , James Scott
    • , Joban Sehmi
    •  & Jaspal Kooner
  144. Department of Genetics, University of North Carolina, Chapel Hill, North Carolina, USA.

    • Yun Li
    • , Karen L Mohlke
    •  & Ying Wu
  145. Department of Biostatistics, University of North Carolina, Chapel Hill, North Carolina, USA.

    • Yun Li
  146. Beijing Genomics Institute, Shenzhen, China.

    • Junbin Liang
    • , Mingyu Yang
    •  & Fan Zhang
  147. Department of Medical Sciences, Uppsala University, Akademiska Sjukhuset, Uppsala, Sweden.

    • Lars Lind
  148. Charles R. Bronfman Institute for Personalized Medicine, Mount Sinai School of Medicine, New York, New York, USA.

    • Ruth J F Loos
  149. Child Health and Development Institute, Mount Sinai School of Medicine, New York, New York, USA.

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

    • Ruth J F Loos
  151. Shanghai Institute of Preventive Medicine, Shanghai, China.

    • Wei Lu
  152. General Medicine Division, Massachusetts General Hospital, Boston, Massachusetts, USA.

    • James B Meigs
  153. Dr Mohan's Diabetes Specialties Centre, Chennai, India.

    • Viswanathan Mohan
  154. Institute of Human Genetics, University of Bonn, Bonn, Germany.

    • Thomas W Mühleisen
    •  & Markus M Nöthen
  155. Department of Genomics, Life and Brain Center, University of Bonn, Bonn, Germany.

    • Thomas W Mühleisen
    •  & Markus M Nöthen
  156. Institute of Medical Informatics, Biometry and Epidemiology, Chair of Genetic Epidemiology, Ludwig-Maximilians-Universität, Munich, Germany.

    • Martina Müller-Nurasyid
  157. Department of Medicine I, University Hospital Grosshadern, Ludwig-Maximilians-University, Munich, Germany.

    • Martina Müller-Nurasyid
  158. Respiratory Medicine, Sir Charles Gairdner Hospital, Nedlands, Western Australia, Australia.

    • Bill Musk
  159. Department of Internal Medicine, Division of Endocrinology and Diabetes, Nagoya University Graduate School of Medicine, Nagoya, Japan.

    • Jiro Nakamura
    •  & Eitaro Nakashima
  160. Department of Diabetes and Endocrinology, Chubu Rosai Hospital, Nagoya, Japan.

    • Eitaro Nakashima
  161. Department of Community Medicine, Faculty of Health Sciences, University of Tromsø, Tromsø, Norway.

    • Inger Njølstad
    •  & Tom Wilsgaard
  162. Department of Geriatric Medicine, Graduate School of Medical Sciences, Kyushu University, Higashi-ku, Fukuoka, Japan.

    • Keizo Ohnaka
  163. Oxford National Institute for Health Research Biomedical Research Centre, Churchill Hospital, Oxford, UK.

    • Katharine R Owen
    •  & Mark I McCarthy
  164. Division of Epidemiology and Community Health, University of Minnesota, Minneapolis, Minnesota, USA.

    • James S Pankow
  165. World Class University Program, Department of Molecular Medicine and Biopharmaceutical Sciences, Graduate School of Convergence Science and Technology and College of Medicine, Seoul National University, Seoul, Korea.

    • Kyong Soo Park
  166. Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden.

    • Nancy L Pedersen
    •  & Emil Rehnberg
  167. Institute for Molecular Medicine Finland (FIMM), Helsinki, Finland.

    • Leena Peltonen
    • , Samuli Ripatti
    •  & Emmi Tikkanen
  168. Institute of Epidemiology II, Helmholtz Zentrum Muenchen, Neuherberg, Germany.

    • Annette Peters
    •  & Barbara Thorand
  169. Department of Public Health, Faculty of Medicine, University of Kelaniya, Ragama, Sri Lanka.

    • Janani M Pinidiyapathirage
    •  & Ananda R Wickremasinghe
  170. Department of Internal Medicine, Levanger Hospital, Nord-Trøndelag Health Trust, Levanger, Norway.

    • Carl G P Platou
  171. University General Hospital Attikon, 1 Rimini, Athens, Greece.

    • Loukianos Rallidis
  172. Institute of Biometrics and Epidemiology, German Diabetes Center, Leibniz Center for Diabetes Research at Heinrich Heine University Düsseldorf, Düsseldorf, Germany.

    • Wolfgang Rathmann
  173. Department of Clinical Physiology and Nuclear Medicine, Kuopio University Hospital, Kuopio, Finland.

    • Rainer Rauramaa
  174. Division of Rheumatology, Immunology and Allergy, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts, USA.

    • Soumya Raychaudhuri
  175. Partners Center for Personalized Genomic Medicine, Boston, Massachusetts, USA.

    • Soumya Raychaudhuri
  176. Department of Endocrinology and Diabetology, Heinrich Heine University Düsseldorf, Düsseldorf, Germany.

    • Michael Roden
  177. Department of Metabolic Diseases, Heinrich Heine University Düsseldorf, Düsseldorf, Germany.

    • Michael Roden
  178. Health Science and Technology MD Program, Harvard University and Massachusetts Institute of Technology, Boston, Massachusetts, USA.

    • Elizabeth J Rossin
  179. Harvard Biological and Biomedical Sciences Program, Harvard University, Boston, Massachusetts, USA.

    • Elizabeth J Rossin
  180. Boston University Data Coordinating Center, Boston, Massachusetts, USA.

    • Denis Rybin
  181. Finnish Diabetes Association, Tampere, Finland.

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

    • Timo E Saaristo
  183. Department of Medicine, Central Finland Central Hospital, Jyväskylä, Finland.

    • Juha Saltevo
  184. South Karelia Central Hospital, Lappeenranta, Finland.

    • Jouko Saramies
  185. University College London Genetics Institute, Department of Genetics, Evolution and Environment, University College London, London, UK.

    • Sonia Shah
    •  & Delilah Zabaneh
  186. Diabetic Association Pakistan, Karachi, Pakistan.

    • A Samad Shera
  187. Division of Endocrinology, Diabetes and Nutrition, University of Maryland School of Medicine, Baltimore, Maryland, USA.

    • Alan R Shuldiner
  188. Geriatric Research Education and Clinical Center, Baltimore Veterans Administration Medical Center, Baltimore, Maryland, USA.

    • Alan R Shuldiner
  189. Program in Personalised and Genomic Medicine, University of Maryland School of Medicine, Baltimore, Maryland, USA.

    • Alan R Shuldiner
  190. Department of Internal Medicine, Erasmus University Medical Center, Rotterdam, The Netherlands.

    • Eric Sijbrands
    • , Andre G Uitterlinden
    •  & Roman Wennauer
  191. Department of Clinical Chemistry and Central Laboratory, University of Ulm, Ulm, Germany.

    • Gerald Steinbach
  192. Molecular Medicine, Department of Medical Sciences, Uppsala University, Uppsala, Sweden.

    • Ann-Christine Syvänen
  193. Department of Internal Medicine and Bioregulatory Science, Graduate School of Medical Sciences, Kyushu University, Higashi-ku, Fukuoka, Japan.

    • Ryoichi Takayanagi
  194. Department of Medicine, Helsinki University Hospital, University of Helsinki, Helsinki, Finland.

    • Tiinamaija Tuomi
  195. Instituto de Investigacion Sanitaria del Hospital Universario LaPaz (IdiPAZ), Madrid, Spain.

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

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

    • Jaakko Tuomilehto
  198. Unidad de Investigacion Medica en Bioquimica, Hospital de Especialidades, Centro Medico Nacional Siglo XXI, IMSS, Av. Cuauhtemoc 330, Col. Doctores, Mexico City, Mexico.

    • Adan Valladares-Salgado
    •  & Miguel Cruz
  199. University of Pennsylvania–Perelman School of Medicine, Department of Pharmacology, Philadelphia, Pennsylvania, USA.

    • Benjamin F Voight
  200. Centre for Eye Research Australia, University of Melbourne, East Melbourne, Victoria, Australia.

    • Tien Yin Wong
  201. Division of Genome Analysis, Research Center for Genetic Information, Medical Institute of Bioregulation, Kyushu University, Higashi-ku, Fukuoka, Japan.

    • Ken Yamamoto
  202. University Lille 1, Laboratory of Mathematics, CNRS-UMR 8524, MODAL team, INRIA Lille Nord-Europe, Villeneuve-d'Ascq, France.

    • Loic Yengo
  203. Department of Genome Science, Aichi-Gakuin University, School of Dentistry, Nagoya, Japan.

    • Mitsuhiro Yokota
  204. Department of Genetics, Harvard Medical School, Boston, Massachusetts, USA.

    • David Altshuler
  205. Department of Molecular Biology, Harvard Medical School, Boston, Massachusetts, USA.

    • David Altshuler
  206. Diabetes Unit, Massachusetts General Hospital, Boston, Massachusetts, USA.

    • David Altshuler
  207. Department of Biochemistry, Wake Forest School of Medicine, Winston-Salem, North Carolina, USA.

    • Donald W Bowden
  208. Department of Internal Medicine, Wake Forest School of Medicine, Winston-Salem, North Carolina, USA.

    • Donald W Bowden
  209. Department of Biomedical Science, Hallym University, Chuncheon, Gangwon-do, Korea.

    • Yoon Shin Cho
  210. Human Genetics Center, University of Texas Health Science Center at Houston, Houston, Texas, USA.

    • Craig L Hanis
  211. Imperial College Healthcare NHS Trust, London, UK.

    • Jaspal Kooner
    •  & John C Chambers
  212. Institute for Human Genetics, University of California, San Francisco, San Francisco, California, USA.

    • Mark Seielstad
  213. Blood Systems Research Institute, San Francisco, California, USA.

    • Mark Seielstad
  214. Graduate School for Integrative Science and Engineering, National University of Singapore, Singapore.

    • Yik Ying Teo
  215. Department of Statistics and Applied Probability, National University of Singapore, Singapore.

    • Yik Ying Teo
  216. Department of Medicine, National University of Singapore, Singapore.

    • E Shyong Tai
  217. Duke–National University of Singapore Graduate Medical School, Singapore.

    • E Shyong Tai
  218. Department of Biostatistics, University of Liverpool, Liverpool, UK.

    • Andrew P Morris

Consortia

  1. DIAbetes Genetics Replication And Meta-analysis (DIAGRAM) Consortium

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

  2. Asian Genetic Epidemiology Network Type 2 Diabetes (AGEN-T2D) Consortium

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

  3. South Asian Type 2 Diabetes (SAT2D) Consortium

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

  4. Mexican American Type 2 Diabetes (MAT2D) Consortium

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

  5. Type 2 Diabetes Genetic Exploration by Next-generation sequencing in multi-Ethnic Samples (T2D-GENES) Consortium

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

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Contributions

Writing group: A. Mahajan, M.J.G., W. Zhang, J.E.B., K.J.G., M.H., A.D.J., I.P., E.Z., Y.Y.T., M.B., E.J.P., J.C.C., E.S.T., M.I.M. and A.P.M.

Analysis group: A. Mahajan, M.J.G., W. Zhang, J.E.B., K.J.G., T. Ferreira, M.H., A.D.J., M.C.Y.N., I.P., D.S., X.W., E.Z., Y.Y.T., M.B., E.J.P., M.I.M. and A.P.M.

DIAGRAM Consortium samples, genotyping, analysis and management: A. Mahajan, M.H., I.P., D.S., E.Z., G.R.A., P.A., M.A., D.B., B.B., I.B., J.B., R.B., R.N.B., B.O.B., E.B., L.L.B., N.B., H. Campbell, J.C., S.C., G.C., H. Chen, P.S.C., F.S.C., M.C.C., D.J.C., A.T.C., R.M.v.D., J. Danesh, U.d.F., G.D., P.D., A.S.D., C.D., A.S.F.D., P.J.D., M.D., C.v.D., J. Dupuis, S.E., V.E., R.E., J.G.E., T.E., E.E., T. Ferreira, J.C.F., P. Fontanillas, N.G.F., T. Forsen, C.F., R.M.F., T.M.F., P. Froguel, K.G., C. Gieger, B.G., H.G., G.B.G., L.C.G., C.J.G., C. Guiducci, A. Hamsten, A.T.H., C. Hayward, C. Herder, A. Hofman, O.L.H., K. Hovingh, A.B.H., F.B.H., J.H., S.E. Humphries, S.E. Hunt, D.J.H., K. Hveem, T.I., E.I., B.I., A.U.J., A. James, K.-H.J., A. Jonsson, H.M.K., S. Kanoni, W.H.L.K., S. Kathiresan, S.M.K.-K., H.K., K.-T.K., L.K., N. Klopp, A. Kong, E.K.-H., P. Kraft, J. Kravic, A. Kumar, J. Kuusisto, M. Laakso, V. Lagou, T.A.L., C. Langenberg, C. Langford, R.L., K.L., M. Li, L.L., C.M.L., E.L., C.-T.L., S. Lobbens, R.J.F.L., J. Luan, V. Lyssenko, R.M., S. Männistö, J.B.M., O.M., A. Metspalu, J.M., G.M., E.M., S. Moebus, K.L.M., A.D.M., T.W.M., M.M.-N., B.M., P.N., P.M.N., I.N., M.M.N., K.R.O., C.N.A.P., J.S.P., M.P., S. Pechlivanis, N.L.P., L.P., J.R.B.P., A.P., C.G.P.P., S. Potter, J.F.P., L.Q., L.R., W.R., R.R., S. Raychaudhuri, N.W.R., E.R., S. Ripatti, N.R., M.R., E.J.R., I.R., D.R., T.E.S., V. Salomaa, J. Saltevo, J. Saramies, L.J.S., R.A.S., A.V.S., B.S., S. Shah, A.R.S., G. Sigurðsson, E.S., A. Silveira, S. Sivapalaratnam, A. Stančáková, K. Stefansson, G. Steinbach, V. Steinthorsdottir, K. Stirrups, R.J.S., H.M.S., Q.S., A.-C.S., T.M.T., B.T., G.T., U.T., E. Tikkanen, J. Trakalo, E. Tremoli, M.D.T., T.T., J. Tuomilehto, A.G.U., S.V., F.V., B.F.V., N.J.W., R.W., T.W., J.F.W., S.W., W.W., A.R. Wood, L.Y., D.Z., D.A., M.B., M.I.M. and A.P.M.

AGEN-T2D Consortium samples, genotyping, analysis and management: M.J.G., X.W., L.S.A., T.A., Y.B., Q.C., J.C.N.C., L.-C.C., T.-J.C., Y.-C.C., C.-H.C., Y.-T.C., N.H.C., Y.M.C., L.-M.C., Y.G., B.-G.H., K. Hara, A.K.H., C. Hu, F.B.H., H.I., W.J., T.K., N. Kato, H.-L.K., S. Kim, Y.J.K., S.H.K., J.-M.L., N.R.L., Y.L., J.J.L., J. Long, W.L., R.C.W.M., S. Maeda, K.L.M., J.N., E.N., P.-K.N., K.O., T.H.O., K.S.P., X.O.S., X.S., W.Y.S., R.T., W.T.T., F.J.T., C.W., T.Y.W., J.-Y.W., Y.W., K.Y., T.Y., M. Yokota, R.Z., W. Zheng, Y.S.C., J.-Y.L., M. Seielstad, Y.Y.T., E.S.T. and M.I.M.

SAT2D Consortium samples, genotyping, analysis and management. W. Zhang, I.P., D.S., G.R.A., T.A., A.H.B., A.B., L.F.B., M. Caulfield, K.-S.C., M. Chidambaram, J. Danesh, D.D., P.D., A.S.D., P.E., T.M.F., P. Froguel, P. Frossard, E.G., N.H., A.K.H., Z.I.H., M.I., T.J., J.B.M.J., N. Kato, P. Katulanda, A.M.K., C.-C.K., S. Kowlessur, M.M.K., X.L., J. Liang, S. Liju, W.-Y.L., J.J.L., D.R.M., V.M., A.C.N., J.M.P., V.R., A.R., S.D.R., M. Samuel, D.K.S., J. Scott, J. Sehmi, N.S., A.S.S., X.S., K.S.S., C.S., R.T., F.T., A.R. Wickremasinghe, T.Y.W., M. Yang, R.Y., F.Z., P.Z.Z., J. Kooner, M. Seielstad, Y.Y.T., J.C.C., E.S.T. and M.I.M.

MAT2D Consortium samples, genotyping, analysis and management: J.E.B., G.I.B., J.E., S. Krithika, J. Kumate, A.V.-S., N.J.C., M. Cruz, C.L.H. and E.J.P.

Project management: D.A., D.W.B., Y.S.C., N.J.C., M. Cruz, C.L.H., J. Kooner, J.-Y.L., M. Seielstad, Y.Y.T., M.B., E.J.P., J.C.C., E.S.T., M.I.M. and A.P.M.

Competing interests

K. Stefansson, V. Steinthorsdottir, G.T. and U.T. are employed by deCODE Genetics/Amgen inc. I.B. and spouse own stock in GlaxoSmithKline and Incyte.

Corresponding authors

Correspondence to Michael Boehnke or Esteban J Parra or John C Chambers or E Shyong Tai or Mark I McCarthy or Andrew P Morris.

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https://doi.org/10.1038/ng.2897

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