Article

Meta-analysis identifies 13 new loci associated with waist-hip ratio and reveals sexual dimorphism in the genetic basis of fat distribution

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Abstract

Waist-hip ratio (WHR) is a measure of body fat distribution and a predictor of metabolic consequences independent of overall adiposity. WHR is heritable, but few genetic variants influencing this trait have been identified. We conducted a meta-analysis of 32 genome-wide association studies for WHR adjusted for body mass index (comprising up to 77,167 participants), following up 16 loci in an additional 29 studies (comprising up to 113,636 subjects). We identified 13 new loci in or near RSPO3, VEGFA, TBX15-WARS2, NFE2L3, GRB14, DNM3-PIGC, ITPR2-SSPN, LY86, HOXC13, ADAMTS9, ZNRF3-KREMEN1, NISCH-STAB1 and CPEB4 (P = 1.9 × 10−9 to P = 1.8 × 10−40) and the known signal at LYPLAL1. Seven of these loci exhibited marked sexual dimorphism, all with a stronger effect on WHR in women than men (P for sex difference = 1.9 × 10−3 to P = 1.2 × 10−13). These findings provide evidence for multiple loci that modulate body fat distribution independent of overall adiposity and reveal strong gene-by-sex interactions.

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Change history

  • Corrected online 12 October 2011

    In the version of this article initially published, there were errors in Table 1. Specifically, for eight SNPs, the effect allele frequencies were reversed. The correct effect allele frequencies for rs9491696, rs984222, rs4846567, rs1011731, rs718314, rs1294421, rs6795735 and rs2076529 are 0.480, 0.635, 0.717, 0.428, 0.259, 0.613, 0.594 and 0.430, respectively. These errors have been corrected in the HTML and PDF versions of the article.

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Acknowledgements

Funding for this study was provided by the Academy of Finland (grants 104781, 120315, 129269, 117797, 121584, 126925, 129418, 129568, 77299, 124243, 213506, 129680, 129494, 10404, 213506, 129680, 114382, 126775, 127437, 129255, 129306, 130326, 209072, 210595, 213225 and 216374); an ADA Mentor-Based Postdoctoral Fellowship grant; Affymetrix, Inc., for genotyping services (N02-HL-6-4278); ALF/LUA Gothenburg; Althingi (the Icelandic Parliament); Amgen; AstraZeneca AB; Augustinus Foundation; Becket Foundation; Biocentrum Helsinki; Biomedicum Helsinki Foundation; Boston Obesity Nutrition Research Center (DK46200); British Diabetes Association (1192); British Diabetic Association Research; British Heart Foundation (97020, PG/02/128); Busselton Population Medical Research Foundation; Cambridge NIHR Comprehensive Biomedical Research Centre; CamStrad; Chief Scientist Office of the Scottish Government; Contrat Plan Etat Région de France; Danish Centre for Health Technology Assessment; Danish Diabetes Association; Danish Ministry of Internal Affairs and Health; Danish Heart Foundation; Danish Pharmaceutical Association; Danish Research Council; DIAB Core (German Network of Diabetes); Diabetes UK; Donald W. Reynolds Foundation; Dresden University of Technology Funding Grant, Med Drive; EMGO+ institute; Emil and Vera Cornell Foundation; Erasmus Medical Center and Erasmus University, Rotterdam, The Netherlands; Estonian Government SF0180142s08; European Commission (2004310, 212111, 205419, 245536, DG XII, HEALTH-F4-2007-201413, FP7/2007-2013, QLG1-CT-2000-01643, QLG2-CT-2002-01254, LSHG-CT-2006-018947, LSHG-CT-2006-01947, LSHG-CT-2004-512066, LSHM-CT-2007-037273, EU/WLRT-2001-01254, LSHG-CT-2004-518153, SOC 95201408 05F02, Marie Curie Intra-European Fellowship); Federal Ministry of Education and Research, Germany (01ZZ9603, 01ZZ0103, 01ZZ0403, 03ZIK012, 01 EA 9401); Federal State of Mecklenburg-West Pomerania; Finnish Diabetes Research Foundation; Finnish Diabetes Research Society; Finnish Foundation for Pediatric Research; Finnish Foundation of Cardiovascular Research; Finnish Medical Society; Finska Läkaresällskapet; Finnish Ministry of Education; Folkhälsan Research Foundation; Fond Européen pour le Développement Régional; Fondation LeDucq; Foundation for Life and Health in Finland; GEN-AU 'GOLD' from Austria; German Bundesministerium fuer Forschung und Technology (# 01 AK 803 A-H, # 01 IG 07015 G); German National Genome Research Net NGFN2 and NGFNplus (01GS0823, FKZ 01GS0823); German Research Council (KFO-152); GlaxoSmithKline; Göteborg Medical Society; Gyllenberg Foundation; Health Care Centers in Vasa, Närpes and Korsholm; Healthway, Western Australia; Helmholtz Center Munich; Helsinki University Central Hospital; Hjartavernd (the Icelandic Heart Association); Ib Henriksen Foundation; IZKF (B27); Jalmari and Rauha Ahokas Foundation; Juho Vainio Foundation; Juvenile Diabetes Research Foundation International (JDRF); Karolinska Institute and the Stockholm County Council (560183); Knut and Alice Wallenberg Foundation; Lundbeck Foundation Centre of Applied Medical Genomics for Personalized Disease Prediction, Prevention and Care; Knut Krohn, Microarray Core Facility of the Interdisciplinary Centre for Clinical Research (IZKF), University of Leipzig, Germany; Lundberg Foundation; MC Health; Ministry of Cultural Affairs of the Federal State of Mecklenburg-West Pomerania, Germany; South Tyrol Ministry of Health; Ministry of Science, Education and Sport of the Republic of Croatia (216-1080315-0302); Medical Research Council UK (G0000649, G0601261, G9521010D, G0000934, G0500539, G0600331, PrevMetSyn); Montreal Heart Institute Foundation; MRC Centre for Obesity-Related Metabolic Disease; Municipal Health Care Center and Hospital in Jakobstad; Municipality of Rotterdam; Närpes Health Care Foundation; National Health and Medical Research Council of Australia and the Great Wine Estates Auctions; Netherlands Centre for Medical Systems Biology (SPI 56-464-1419); Netherlands Ministry for Health, Welfare and Sports; Netherlands Ministry of Education, Culture and Science; Netherlands Genomics Initiative; Netherlands Consortium for Healthy Aging (050-060-810); Netherlands Organisation of Scientific Research Netherlandse Organisatie voor Wetenschappelijk Onderzoek (NWO) Investments (175.010.2005.011, 911-03-012, 904-61-090, 904-61-193, 480-04-004, 400-05-717); National Institute on Aging Intramural Research Program; US National Institutes of Health (CA047988, CA65725, CA87969, CA49449, CA67262, CA50385, DK075787, DK062370, DK58845, DK072193, K23-DK080145, K99HL094535, N01-HC85079 through N01-HC85086, N01-HG-65403, N01-AG-12100, N01-HC-25195, N01-HC35129, N01-HC15103, N01-HC55222, N01-HC75150, N01-HC45133, N01-HC55015, N01-HC55016, N01-HC-55018, N01-HC-55019, N01-HC-55020, N01-HC-55021, N01-HC-55022, NO1-AG-1-2109, HL71981, HG005581, HG002651, HL084729, HL043851, HHSN268200625226C, K23-DK080145, MH084698, P30-DK072488, R01-DK075787, R01 HL087652, R01-HL087641, R01-HL59367, R01-HL086694, R01-HL087647, R01-HL087679, R01-HL087700, R01-AG031890, R01-HL088119, R01-DK068336, R01-DK075681, R01-DK-073490, R01-DK075787, R01-MH63706, U01-HL72515, U01-GM074518, U01-HL084756, U01-HG004399, UO1-CA098233, UL1-RR025005, UL1-RR025005, U01-HG004402, U01-DK062418, U01 HL080295, T32-HG00040, 263-MA-410953, 1RL1-MH083268-01, intramural project 1Z01-HG000024); National Institute for Health Research (NIHR); Neuroscience Campus Amsterdam; Novo Nordisk Foundation; Novo Nordisk Inc., Research Foundation of Copenhagen County; Ollqvist Foundation; Paavo Nurmi Foundation; Päivikki and Sakari Sohlberg Foundation; Pew Scholarship for the Biomedical Sciences; Perklén Foundation; Petrus and Augusta Hedlunds Foundation; Research Institute for Diseases in the Elderly (014-93-015, RIDE, RIDE2); Sahlgrenska Center for Cardiovascular and Metabolic Research (CMR, A305:188); Siemens Healthcare, Erlangen, Germany; Signe and Ane Gyllenberg Foundation; Sigrid Juselius Foundation; Social Insurance Institution of Finland; Social Ministry of the Federal State of Mecklenburg-West Pomerania, Germany; South Tyrolean Sparkasse Foundation; State of Bavaria, Germany; Support for Science Funding programme; Swedish Cultural Foundation in Finland; Swedish Foundation for Strategic Research (SSF); Swedish Heart-Lung Foundation; Swedish Medical Research Council (8691, K2007-66X-20270-01-3, K2010-54X-09894-19-3); Swedish Society of Medicine; Swiss National Science Foundation (33CSCO-122661); the Royal Society; the Royal Swedish Academy of Science; Torsten and Ragnar Söderberg's Foundation; Turku University Hospitals; UK Department of Health Policy Research Programme; University and Research of the Autonomous Province of Bolzano; University Hospital Medical funds to Tampere; University Hospital Oulu, Biocenter, University of Oulu, Finland (75617); Västra Götaland Foundation; Wellcome Trust (077016/Z/05/Z, 068545/Z/02, 072960, 076113, 083270, 085301, 079557, 081682, 075491, 076113/B/04/Z, 091746/Z/10/Z, 079895, WT086596/Z/08/Z, WT Research Career Development Fellowship; WT Career Development Award); Western Australian Genetic Epidemiology Resource and the Western Australian DNA Bank (both National Health and Medical Research Council of Australia Enabling Facilities); Yrjö Jahnsson Foundation.

Author information

Author notes

    • Iris M Heid
    • , Anne U Jackson
    • , Joshua C Randall
    • , Thomas W Winkler
    • , Lu Qi
    • , Valgerdur Steinthorsdottir
    •  & Gudmar Thorleifsson

    These authors contributed equally to this work.

    • Kari Stefansson
    • , L Adrienne Cupples
    • , Ruth J F Loos
    • , Inês Barroso
    • , Mark I McCarthy
    • , Caroline S Fox
    • , Karen L Mohlke
    •  & Cecilia M Lindgren

    These authors jointly directed this work.

Affiliations

  1. Regensburg University Medical Center, Department of Epidemiology and Preventive Medicine, Regensburg, Germany.

    • Iris M Heid
    • , Thomas W Winkler
    •  & Michael F Leitzmann
  2. Institute of Epidemiology, Helmholtz Zentrum München-German Research Center for Environmental Health, Neuherberg, Germany.

    • Iris M Heid
    • , Claudia Lamina
    • , Harald Grallert
    • , Thomas Illig
    •  & H-Erich Wichmann
  3. Department of Biostatistics, Center for Statistical Genetics, University of Michigan, Ann Arbor, Michigan, USA.

    • Anne U Jackson
    • , Cristen J Willer
    • , Robert J Weyant
    • , Laura J Scott
    • , Tanya M Teslovich
    • , Ryan Welch
    • , Heather M Stringham
    • , Gonçalo R Abecasis
    •  & Michael Boehnke
  4. Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, UK.

    • Joshua C Randall
    • , Reedik Mägi
    • , Teresa Ferreira
    • , Andrew P Morris
    • , Inga Prokopenko
    • , Nigel W Rayner
    • , Neil R Robertson
    • , John F Peden
    • , Mark I McCarthy
    •  & Cecilia M Lindgren
  5. Department of Nutrition, Harvard School of Public Health, Boston, Massachusetts, USA.

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

    • Lu Qi
    • , Frank B Hu
    •  & David J Hunter
  7. deCODE Genetics, Reykjavik, Iceland.

    • Valgerdur Steinthorsdottir
    • , Gudmar Thorleifsson
    • , G Bragi Walters
    • , Unnur Thorsteinsdottir
    •  & Kari Stefansson
  8. Department of Internal Medicine, Erasmus Medical Center (MC), Rotterdam, The Netherlands.

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

    • M Carola Zillikens
    • , Karol Estrada
    • , Marjolein J Peters
    • , Fernando Rivadeneira
    • , Joyce B J van Meurs
    • , Albert Hofman
    • , André Uitterlinden
    • , Jacqueline C Witteman
    •  & Cornelia M van Duijn
  10. Metabolism Initiative and Program in Medical and Population Genetics, Broad Institute, Cambridge, Massachusetts, USA.

    • Elizabeth K Speliotes
    • , Sailaja Vedantam
    • , Candace Guiducci
    • , Brian Thomson
    •  & Joel N Hirschhorn
  11. Division of Gastroenterology, Massachusetts General Hospital, Boston, Massachusetts, USA.

    • Elizabeth K Speliotes
    •  & Lee M Kaplan
  12. Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts, USA.

    • Charles C White
    •  & L Adrienne Cupples
  13. Centre National de la Recherche Scientifique (CNRS), UMR8199-IBL-Institut Pasteur de Lille, Lille, France.

    • Nabila Bouatia-Naji
    • , Christine Cavalcanti-Proença
    • , Cecile Lecoeur
    • , Stefan Gaget
    • , Vincent Vatin
    •  & Philippe Froguel
  14. University Lille Nord de France, Lille, France.

    • Nabila Bouatia-Naji
    • , Christine Cavalcanti-Proença
    • , Cecile Lecoeur
    • , Stefan Gaget
    • , Vincent Vatin
    •  & Philippe Froguel
  15. Laboratory of Epidemiology, Demography, Biometry, National Institute on Aging, National Institutes of Health, Bethesda, Maryland, USA.

    • Tamara B Harris
    •  & Lenore J Launer
  16. Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Department of Health and Human Services, Bethesda, Maryland, USA.

    • Sonja I Berndt
  17. Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden.

    • Erik Ingelsson
  18. Genetics of Complex Traits, Peninsula College of Medicine and Dentistry, University of Exeter, Exeter, UK.

    • Michael N Weedon
    • , Hana Lango Allen
    • , Andrew R Wood
    • , John R B Perry
    • , Andrew T Hattersley
    •  & Timothy M Frayling
  19. Medical Research Council (MRC) Epidemiology Unit, Institute of Metabolic Science, Addenbrooke's Hospital, Cambridge, UK.

    • Jian'an Luan
    • , Tuomas O Kilpeläinen
    • , Shengxu Li
    • , Ken K Ong
    • , Jing Hua Zhao
    • , Nicholas J Wareham
    •  & Ruth J F Loos
  20. Divisions of Genetics and Endocrinology and Program in Genomics, Children's Hospital, Boston, Massachusetts, USA.

    • Sailaja Vedantam
    •  & Joel N Hirschhorn
  21. Estonian Genome Center, University of Tartu, Tartu, Estonia.

    • Tõnu Esko
    • , Mari-Liis Tammesoo
    • , Helene Alavere
    • , Mari Nelis
    •  & Andres Metspalu
  22. Estonian Biocenter, Tartu, Estonia.

    • Tõnu Esko
    • , Mari Nelis
    • , Maris Teder-Laving
    •  & Andres Metspalu
  23. Institute of Molecular and Cell Biology, University of Tartu, Tartu, Estonia.

    • Tõnu Esko
    • , Mari Nelis
    • , Maris Teder-Laving
    •  & Andres Metspalu
  24. Department of Medical Genetics, University of Lausanne, Lausanne, Switzerland.

    • Zoltán Kutalik
    • , Toby Johnson
    • , Karen Kapur
    • , Jacques S Beckmann
    •  & Sven Bergmann
  25. Swiss Institute of Bioinformatics, Lausanne, Switzerland.

    • Zoltán Kutalik
    • , Toby Johnson
    • , Karen Kapur
    •  & Sven Bergmann
  26. Department of Epidemiology, School of Public Health, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, USA.

    • Keri L Monda
    •  & Kari E North
  27. Department of Pharmacy and Pharmacology, University of Bath, Bath, UK.

    • Anna L Dixon
  28. MRC Harwell, Harwell Science and Innovation Campus, Oxfordshire, UK.

    • Christopher C Holmes
  29. Department of Statistics, University of Oxford, Oxford, UK.

    • Christopher C Holmes
    •  & George Nicholson
  30. Harvard Medical School, Boston, Massachusetts, USA.

    • Lee M Kaplan
    • , Daniel I Chasman
    •  & Paul M Ridker
  31. Massachusetts General Hospital (MGH) Weight Center, Massachusetts General Hospital, Boston, Massachusetts, USA.

    • Lee M Kaplan
  32. Department of Epidemiology, Harvard School of Public Health, Boston, Massachusetts, USA.

    • Liming Liang
    • , Peter Kraft
    • , Frank B Hu
    •  & David J Hunter
  33. Department of Biostatistics, Harvard School of Public Health, Boston, Massachusetts, USA.

    • Liming Liang
    •  & Peter Kraft
  34. Human Genetics, Leiden University Medical Center, Leiden, The Netherlands.

    • Josine L Min
  35. National Heart and Lung Institute, Imperial College London, London, UK.

    • Miriam F Moffatt
  36. Merck Research Laboratories, Merck & Co., Inc., Boston, Massachusetts, USA.

    • Cliona Molony
  37. Pacific Biosciences, Menlo Park, California, USA.

    • Eric E Schadt
  38. Sage Bionetworks, Seattle, Washington, USA.

    • Eric E Schadt
  39. Genetic and Genomic Epidemiology Unit, Wellcome Trust Centre for Human Genetics, Oxford, UK.

    • Krina T Zondervan
  40. Department of Genetics, Washington University School of Medicine, St. Louis, Missouri, USA.

    • Mary F Feitosa
    • , Shamika Ketkar
    • , Aldi T Kraja
    •  & Ingrid B Borecki
  41. Wellcome Trust Sanger Institute, Hinxton, Cambridge, UK.

    • Eleanor Wheeler
    • , Nicole Soranzo
    • , Panos Deloukas
    • , Leena Peltonen
    •  & Inês Barroso
  42. Department of Epidemiology, Erasmus MC, Rotterdam, The Netherlands.

    • Karol Estrada
    • , Najaf Amin
    • , Fernando Rivadeneira
    • , Sophie van Wingerden
    • , Joyce B J van Meurs
    • , Albert Hofman
    • , André Uitterlinden
    • , Jacqueline C Witteman
    •  & Cornelia M van Duijn
  43. University of Melbourne, Parkville, Australia.

    • Michael E Goddard
  44. Department of Primary Industries, Melbourne, Victoria, Australia.

    • Michael E Goddard
  45. Montreal Heart Institute, Montreal, Quebec, Canada.

    • Guillaume Lettre
  46. Department of Medicine, Université de Montréal, Montreal, Quebec, Canada.

    • Guillaume Lettre
  47. Department of Twin Research and Genetic Epidemiology, King's College London, London, UK.

    • Massimo Mangino
    • , Nicole Soranzo
    •  & Timothy D Spector
  48. Neurogenetics Laboratory, Queensland Institute of Medical Research, Queensland, Australia.

    • Dale R Nyholt
  49. Center for Human Genetic Research, Massachusetts General Hospital, Boston, Massachusetts, USA.

    • Shaun Purcell
    • , Steven A McCarroll
    •  & Benjamin F Voight
  50. The Broad Institute of Harvard and Massachusetts Institute of Technology (MIT), Cambridge, Massachusetts, USA.

    • Shaun Purcell
    •  & Leena Peltonen
  51. Department of Psychiatry, Harvard Medical School, Boston, Massachusetts, USA.

    • Shaun Purcell
  52. Icelandic Heart Association, Kopavogur, Iceland.

    • Albert Vernon Smith
    • , Thor Aspelund
    • , Gudny Eiriksdottir
    •  & Vilmundur Gudnason
  53. University of Iceland, Reykjavik, Iceland.

    • Albert Vernon Smith
    • , Thor Aspelund
    •  & Vilmundur Gudnason
  54. Queensland Statistical Genetics Laboratory, Queensland Institute of Medical Research, Queensland, Australia.

    • Peter M Visscher
    •  & Jian Yang
  55. Program in Medical and Population Genetics, Broad Institute of Harvard and MIT, Cambridge, Massachusetts, USA.

    • Steven A McCarroll
    • , James Nemesh
    •  & Benjamin F Voight
  56. Department of Molecular Biology, Massachusetts General Hospital, Boston, Massachusetts, USA.

    • Steven A McCarroll
    •  & Benjamin F Voight
  57. Hudson Alpha Institute for Biotechnology, Huntsville, Alabama, USA.

    • Devin Absher
  58. Department of Epidemiology and Biostatistics, School of Public Health, Faculty of Medicine, Imperial College London, London, UK.

    • Lachlan Coin
    • , Ulla Sovio
    • , Paul Elliott
    •  & Marjo-Riitta Jarvelin
  59. Department of Medicine, University of Washington, Seattle, Washington, USA.

    • Nicole L Glazer
  60. Cardiovascular Health Research Unit, University of Washington, Seattle, Washington, USA.

    • Nicole L Glazer
  61. MRC Human Genetics Unit, Institute for Genetics and Molecular Medicine, Western General Hospital, Edinburgh, Scotland, UK.

    • Caroline Hayward
    • , Veronique Vitart
    •  & Alan F Wright
  62. Department of Neurology, Boston University School of Medicine, Boston, Massachusetts, USA.

    • Nancy L Heard-Costa
    •  & Larry D Atwood
  63. Department of Biological Psychology, Vrije Universiteit (VU) University Amsterdam, Amsterdam, The Netherlands.

    • Jouke-Jan Hottenga
    • , Eco J C Geus
    • , Gonneke Willemsen
    •  & Dorret I Boomsma
  64. Department of Genetics and Pathology, Rudbeck Laboratory, University of Uppsala, Uppsala, Sweden.

    • Åsa Johansson
    • , Wilmar Igl
    •  & Ulf Gyllensten
  65. Department of Cancer Research and Molecular Medicine, Faculty of Medicine, Norwegian University of Science and Technology (NTNU), Trondheim, Norway.

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

    • Toby Johnson
  67. Clinical Pharmacology and 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.

    • Toby Johnson
    • , Mark J Caulfield
    •  & Patricia B Munroe
  68. Institute of Health Sciences, University of Oulu, Oulu, Finland.

    • Marika Kaakinen
    •  & Marjo-Riitta Jarvelin
  69. Biocenter Oulu, University of Oulu, Oulu, Finland.

    • Marika Kaakinen
    • , Karl-Heinz Herzig
    •  & Marjo-Riitta Jarvelin
  70. Department of Medicine, Stanford University School of Medicine, Stanford, California, USA.

    • Joshua W Knowles
    • , Thomas Quertermous
    •  & Themistocles L Assimes
  71. Division of Genetic Epidemiology, Department of Medical Genetics, Molecular and Clinical Pharmacology, Innsbruck Medical University, Innsbruck, Austria.

    • Claudia Lamina
  72. Department of Biostatistics, University of Washington, Seattle, Washington, USA.

    • Barbara McKnight
    •  & Alice M Arnold
  73. Andrija Stampar School of Public Health, Medical School, University of Zagreb, Zagreb, Croatia.

    • Ozren Polasek
    • , Ivana Kolcic
    •  & Lina Zgaga
  74. Gen-Info Ltd, Zagreb, Croatia.

    • Ozren Polasek
  75. Oxford Centre for Diabetes, Endocrinology and Metabolism, University of Oxford, Oxford, UK.

    • Inga Prokopenko
    • , Nigel W Rayner
    • , Neil R Robertson
    • , Amanda J Bennett
    • , Christopher J Groves
    • , Neelam Hassanali
    • , Matt J Neville
    • , Fredrik Karpe
    • , Mark I McCarthy
    •  & Cecilia M Lindgren
  76. Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland.

    • Samuli Ripatti
    • , Ida Surakka
    • , Kaisa Silander
    • , Johannes Kettunen
    • , Niina Pellikka
    • , Markus Perola
    • , Elisabeth Widen
    • , Jaakko Kaprio
    •  & Leena Peltonen
  77. National Institute for Health and Welfare, Department of Chronic Disease Prevention, Unit of Public Health Genomics, Helsinki, Finland.

    • Samuli Ripatti
    • , Ida Surakka
    • , Kaisa Silander
    • , Johannes Kettunen
    • , Niina Pellikka
    •  & Markus Perola
  78. Istituto di Neurogenetica e Neurofarmacologia del CNR, Monserrato, Cagliari, Italy.

    • Serena Sanna
    • , Mariano Dei
    • , Gianluca Usala
    •  & Manuela Uda
  79. Interfaculty Institute for Genetics and Functional Genomics, Ernst-Moritz-Arndt-University Greifswald, Greifswald, Germany.

    • Alexander Teumer
  80. National Human Genome Research Institute, National Institutes of Health, Bethesda, Maryland, USA.

    • Peter S Chines
    • , Narisu Narisu
    • , Lori L Bonnycastle
    • , Michael R Erdos
    • , Mario A Morken
    • , Amy J Swift
    •  & Francis S Collins
  81. Department of Epidemiology, German Institute of Human Nutrition Potsdam-Rehbruecke, Nuthetal, Germany.

    • Eva Fisher
    •  & Heiner Boeing
  82. Department of Genetics, University of North Carolina, Chapel Hill, North Carolina, USA.

    • Jennifer R Kulzer
    •  & Karen L Mohlke
  83. Hagedorn Research Institute, Gentofte, Denmark.

    • Camilla Sandholt
    • , Anette P Gjesing
    • , Torben Hansen
    •  & Oluf Pedersen
  84. Regensburg University Medical Center, Clinic and Policlinic for Internal Medicine II, Regensburg, Germany.

    • Klaus Stark
  85. MRC Centre for Causal Analyses in Translational Epidemiology, Department of Social Medicine, Oakfield House, Bristol, UK.

    • Nicholas John Timpson
    • , Ian N M Day
    • , George Davey Smith
    •  & Debbie A Lawlor
  86. Department of Physiology and Biophysics, Keck School of Medicine, University of Southern California, Los Angeles, California, USA.

    • Richard M Watanabe
    • , Thomas A Buchanan
    •  & Richard N Bergman
  87. Department of Preventive Medicine, Keck School of Medicine, University of Southern California, Los Angeles, California, USA.

    • Richard M Watanabe
  88. Division of Preventive Medicine, Brigham and Women's Hospital, Boston, Massachusetts, USA.

    • Daniel I Chasman
    •  & Paul M Ridker
  89. Centre for Genetic Epidemiology and Biostatistics, University of Western Australia, Crawley, Western Australia, Australia.

    • Matthew N Cooper
    • , Robert W Lawrence
    • , Jennie Hui
    •  & Lyle J Palmer
  90. Department of Physiology, Institute of Neuroscience and Physiology, Sahlgrenska Academy, University of Gothenburg, Gothenburg, Sweden.

    • John-Olov Jansson
  91. Department of Internal Medicine, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, Gothenburg, Sweden.

    • Liesbeth Vandenput
    •  & Claes Ohlsson
  92. Lund University Diabetes Centre, Department of Clinical Sciences, Lund University, Malmö, Sweden.

    • Peter Almgren
    •  & Leif C Groop
  93. Zentrum für Zahn-, Mund- und Kieferheilkunde, Greifswald, Germany.

    • Reiner Biffar
  94. Department of Medicine III, University of Dresden, Dresden, Germany.

    • Stefan R Bornstein
    •  & Barbara Ludwig
  95. Division of Endocrinology, Keck School of Medicine, University of Southern California, Los Angeles, California, USA.

    • Thomas A Buchanan
  96. Centre for Population Health Sciences, University of Edinburgh, Teviot Place, Edinburgh, Scotland, UK.

    • Harry Campbell
    • , Sarah H Wild
    • , Igor Rudan
    •  & James F Wilson
  97. Department of Internal Medicine B, Ernst-Moritz-Arndt University, Greifswald, Germany.

    • Marcus Dörr
  98. MRC-Health Protection Agency (HPA) Centre for Environment and Health, London, UK.

    • Paul Elliott
  99. Department of General Practice and Primary Health Care, University of Helsinki, Helsinki, Finland.

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

    • Johan G Eriksson
    • , Eero Kajantie
    •  & Leena Peltonen
  101. Helsinki University Central Hospital, Unit of General Practice, Helsinki, Finland.

    • Johan G Eriksson
  102. Folkhalsan Research Centre, Helsinki, Finland.

    • Johan G Eriksson
    • , Bo Isomaa
    •  & Tiinamaija Tuomi
  103. Vasa Central Hospital, Vasa, Finland.

    • Johan G Eriksson
  104. Center for Neurobehavioral Genetics, University of California, Los Angeles, California, USA.

    • Nelson B Freimer
  105. Department of Medicine, University of Maryland School of Medicine, Baltimore, Maryland, USA.

    • Mao Fu
    • , Alan R Shuldiner
    •  & Jeffrey R O'Connell
  106. Department of Medicine III, Pathobiochemistry, University of Dresden, Dresden, Germany.

    • Jürgen Gräßler
  107. Department of Clinical Sciences/Obstetrics and Gynecology, University of Oulu, Oulu, Finland.

    • Anna-Liisa Hartikainen
    •  & Anneli Pouta
  108. National Institute for Health and Welfare, Department of Chronic Disease Prevention, Chronic Disease Epidemiology and Prevention Unit, Helsinki, Finland.

    • Aki S Havulinna
    • , Pekka Jousilahti
    • , Seppo Koskinen
    •  & Veikko Salomaa
  109. Institute of Biomedicine, Department of Physiology, University of Oulu, Oulu, Finland.

    • Karl-Heinz Herzig
  110. Department of Psychiatry, Kuopio University Hospital and University of Kuopio, Kuopio, Finland.

    • Karl-Heinz Herzig
  111. Institute of Genetic Medicine, European Academy Bozen-Bolzano (EURAC), Bolzano-Bozen, Italy (affiliated Institute of the University of Lübeck, Lübeck, Germany).

    • Andrew A Hicks
    • , Irene Pichler
    • , Claudia B Volpato
    •  & Peter P Pramstaller
  112. PathWest Laboratory of Western Australia, Department of Molecular Genetics, J Block, QEII Medical Centre, Nedlands, Western Australia, Australia.

    • Jennie Hui
    •  & John P Beilby
  113. Busselton Population Medical Research Foundation Inc., Sir Charles Gairdner Hospital, Nedlands, Western Australia, Australia.

    • Jennie Hui
    • , John P Beilby
    • , Alan L James
    •  & Lyle J Palmer
  114. National Institute for Health and Welfare, Department of Chronic Disease Prevention, Population Studies Unit, Turku, Finland.

    • Antti Jula
  115. Hospital for Children and Adolescents, Helsinki University Central Hospital and University of Helsinki, Helsinki, Finland.

    • Eero Kajantie
  116. National Institute for Health and Welfare, Diabetes Prevention Unit, Helsinki, Finland.

    • Leena Kinnunen
    • , Jaakko Tuomilehto
    •  & Timo T Valle
  117. Interdisciplinary Centre for Clinical Research, University of Leipzig, Leipzig, Germany.

    • Peter Kovacs
  118. Institut für Pharmakologie, Universität Greifswald, Greifswald, Germany.

    • Heyo K Kroemer
  119. Croatian Centre for Global Health, School of Medicine, University of Split, Split, Croatia.

    • Vjekoslav Krzelj
    •  & Igor Rudan
  120. Department of Medicine, University of Kuopio and Kuopio University Hospital, Kuopio, Finland.

    • Johanna Kuusisto
    •  & Markku Laakso
  121. Nord-Trøndelag Health Study (HUNT) Research Centre, Department of Public Health and General Practice, Norwegian University of Science and Technology, Levanger, Norway.

    • Kirsti Kvaloy
    • , Carl G P Platou
    • , Kristian Hveem
    •  & Kristian Midthjell
  122. Finnish Institute of Occupational Health, Oulu, Finland.

    • Jaana Laitinen
  123. Institut inter-regional pour la sante (IRSA), La Riche, France.

    • Olivier Lantieri
  124. Centre National de Genotypage, Evry, Paris, France.

    • G Mark Lathrop
  125. Transplantation Laboratory, Haartman Institute, University of Helsinki, Helsinki, Finland.

    • Marja-Liisa Lokki
  126. Department of Public Health and Primary Care, Institute of Public Health, University of Cambridge, Cambridge, UK.

    • Robert N Luben
    •  & Kay-Tee Khaw
  127. Avon Longitudinal Study of Parents and Children (ALSPAC) Laboratory, Department of Social Medicine, University of Bristol, Bristol, UK.

    • Wendy L McArdle
  128. Division of Health, Research Board, An Bord Taighde Sláinte, Dublin, Ireland.

    • Anne McCarthy
  129. Department of Pathology and Molecular Medicine, McMaster University, Hamilton, Ontario, Canada.

    • Guillaume Paré
  130. Amgen, Cambridge, Massachusetts, USA.

    • Alex N Parker
  131. Department of Cardiovascular Medicine, University of Oxford, John Radcliffe Hospital, Headington, Oxford, UK.

    • John F Peden
  132. Finnish Twin Cohort Study, Department of Public Health, University of Helsinki, Helsinki, Finland.

    • Kirsi H Pietiläinen
    •  & Jaakko Kaprio
  133. Obesity Research Unit, Department of Psychiatry, Helsinki University Central Hospital, Helsinki, Finland.

    • Kirsi H Pietiläinen
    •  & Aila Rissanen
  134. Department of Medicine, Levanger Hospital, The Nord-Trøndelag Health Trust, Levanger, Norway.

    • Carl G P Platou
  135. National Institute for Health and Welfare, Oulu, Finland.

    • Anneli Pouta
    •  & Marjo-Riitta Jarvelin
  136. Department of Clinical Sciences, Lund University, Malmö, Sweden.

    • Martin Ridderstråle
  137. Department of Cardiovascular Sciences, University of Leicester, Glenfield Hospital, Leicester, UK.

    • Nilesh J Samani
  138. Leicester National Institute for Health Research (NIHR) Biomedical Research Unit in Cardiovascular Disease, Glenfield Hospital, Leicester, UK.

    • Nilesh J Samani
  139. South Karelia Central Hospital, Lappeenranta, Finland.

    • Jouko Saramies
  140. Division of Cardiology, Cardiovascular Laboratory, Helsinki University Central Hospital, Helsinki, Finland.

    • Juha Sinisalo
    •  & Markku S Nieminen
  141. Department of Psychiatry/Instituut voor Extramuraal Geneeskundig Onderzoek (EMGO) Institute, VU University Medical Center, Amsterdam, The Netherlands.

    • Jan H Smit
    •  & Brenda W Penninx
  142. Atherosclerosis Research Unit, Department of Medicine, Solna, Karolinska Institutet, Karolinska University Hospital, Stockholm, Sweden.

    • Rona J Strawbridge
    •  & Anders Hamsten
  143. Department of Human Genetics, Leiden University Medical Center, Leiden, The Netherlands.

    • Gert-Jan van Ommen
  144. Center of Medical Systems Biology, Leiden University Medical Center, Leiden, The Netherlands.

    • Gert-Jan van Ommen
  145. Institut für Klinische Chemie und Laboratoriumsmedizin, Universität Greifswald, Greifswald, Germany.

    • Henri Wallaschofski
  146. Steno Diabetes Center, Gentofte, Denmark.

    • Daniel R Witte
  147. Department of Physiatrics, Lapland Central Hospital, Rovaniemi, Finland.

    • Paavo Zitting
  148. School of Pathology and Laboratory Medicine, University of Western Australia, Nedlands, Western Australia, Australia.

    • John P Beilby
  149. School of Medicine and Pharmacology, University of Western Australia, Perth, Western Australia, Australia.

    • Alan L James
  150. Department of Clinical Physiology, University of Tampere and Tampere University Hospital, Tampere, Finland.

    • Mika Kähönen
  151. Department of Clinical Chemistry, University of Tampere and Tampere University Hospital, Tampere, Finland.

    • Terho Lehtimäki
  152. Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku, Finland.

    • Olli Raitakari
  153. The Department of Clinical Physiology, Turku University Hospital, Turku, Finland.

    • Olli Raitakari
  154. Department of Medicine, University of Leipzig, Leipzig, Germany.

    • Michael Stumvoll
    •  & Anke Tönjes
  155. Leipziger Interdisziplin?r Forschungskomplex zu molekularen Ursachen umwelt- und lebensstilassoziierter Erkrankungen (LIFE) Study Centre, University of Leipzig, Leipzig, Germany.

    • Michael Stumvoll
  156. Coordination Centre for Clinical Trials, University of Leipzig, Leipzig, Germany.

    • Anke Tönjes
  157. Department of Medicine, University of Turku and Turku University Hospital, Turku, Finland.

    • Jorma Viikari
  158. INSERM Centre de Recherche en Epidémiologie et Santé des Populations (CESP) U1018, Villejuif, France.

    • Beverley Balkau
  159. University Paris Sud 11, Unité Mixte de Recherche en Santé (UMRS) 1018, Villejuif, France.

    • Beverley Balkau
  160. Department of Social Medicine, University of Bristol, Bristol, UK.

    • Yoav Ben-Shlomo
  161. The London School of Hygiene and Tropical Medicine, London, UK.

    • Shah Ebrahim
  162. South Asia Network for Chronic Disease, New Delhi, India.

    • Shah Ebrahim
  163. Department of Genomics of Common Disease, School of Public Health, Imperial College London, London, UK.

    • Philippe Froguel
  164. Faculty of Health Science, University of Southern Denmark, Odense, Denmark.

    • Torben Hansen
  165. Klinik und Poliklinik für Innere Medizin II, Universität Regensburg, Regensburg, Germany.

    • Christian Hengstenberg
  166. Regensburg University Medical Center, Innere Medizin II, Regensburg, Germany.

    • Christian Hengstenberg
  167. Department of Social Services and Health Care, Jakobstad, Finland.

    • Bo Isomaa
  168. Research Centre for Prevention and Health, Glostrup University Hospital, Glostrup, Denmark.

    • Torben Jørgensen
  169. Faculty of Health Science, University of Copenhagen, Copenhagen, Denmark.

    • Torben Jørgensen
  170. NIHR Oxford Biomedical Research Centre, Churchill Hospital, Oxford, UK.

    • Fredrik Karpe
    •  & Mark I McCarthy
  171. Department of Endocrinology, Diabetology and Nutrition, Bichat-Claude Bernard University Hospital, Assistance Publique des Hôpitaux de Paris, Paris, France.

    • Michel Marre
  172. Cardiovascular Genetics Research Unit, Université Henri Poincaré-Nancy 1, Nancy, France.

    • Michel Marre
  173. Institute of Human Genetics, Klinikum rechts der Isar der Technischen Universität München, Munich, Germany.

    • Thomas Meitinger
  174. Institute of Human Genetics, Helmholtz Zentrum München-German Research Center for Environmental Health, Neuherberg, Germany.

    • Thomas Meitinger
  175. Institute of Biomedical Sciences, University of Copenhagen, Copenhagen, Denmark.

    • Oluf Pedersen
  176. Faculty of Health Science, University of Aarhus, Aarhus, Denmark.

    • Oluf Pedersen
  177. Department of Medicine III, Prevention and Care of Diabetes, University of Dresden, Dresden, Germany.

    • Peter E H Schwarz
  178. Department of Medicine, Helsinki University Central Hospital, Helsinki, Finland.

    • Tiinamaija Tuomi
  179. Research Program of Molecular Medicine, University of Helsinki, Helsinki, Finland.

    • Tiinamaija Tuomi
  180. Hjelt Institute, Department of Public Health, University of Helsinki, Helsinki, Finland.

    • Jaakko Tuomilehto
  181. South Ostrobothnia Central Hospital, Seinajoki, Finland.

    • Jaakko Tuomilehto
  182. Collaborative Health Studies Coordinating Center, Seattle, Washington, USA.

    • Alice M Arnold
  183. Service of Medical Genetics, Centre Hospitalier Universitaire Vaudois (CHUV) University Hospital, Lausanne, Switzerland.

    • Jacques S Beckmann
  184. Human Genetics Center and Institute of Molecular Medicine, University of Texas Health Science Center, Houston, Texas, USA.

    • Eric Boerwinkle
  185. Division of Research, Kaiser Permanente Northern California, Oakland, California, USA.

    • Carlos Iribarren
  186. Department of Epidemiology and Biostatistics, University of California, San Francisco, San Francisco, California, USA.

    • Carlos Iribarren
  187. Department of Epidemiology and Medicine, Johns Hopkins Bloomberg School of Public Health, Baltimore, Maryland, USA.

    • W H Linda Kao
  188. National Institute for Health and Welfare, Department of Mental Health and Substance Abuse Services, Unit for Child and Adolescent Mental Health, Helsinki, Finland.

    • Jaakko Kaprio
  189. Department of Clinical Genetics, Erasmus MC, Rotterdam, The Netherlands.

    • Ben Oostra
  190. Department of Psychiatry, Leiden University Medical Centre, Leiden, The Netherlands.

    • Brenda W Penninx
  191. Department of Psychiatry, University Medical Centre Groningen, Groningen, The Netherlands.

    • Brenda W Penninx
  192. Department of Neurology, General Central Hospital, Bolzano, Italy.

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

    • Peter P Pramstaller
  194. Departments of Epidemiology, Medicine and Health Services, University of Washington, Seattle, Washington, USA.

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

    • Bruce M Psaty
  196. Geriatrics Research and Education Clinical Center, Baltimore Veterans Administration Medical Center, Baltimore, Maryland, USA.

    • Alan R Shuldiner
  197. Uppsala University, Department of Medical Sciences, Molecular Medicine, Uppsala, Sweden.

    • Ann-Christine Syvanen
  198. Institut für Community Medicine, Greifswald, Germany.

    • Henry Völzke
  199. Department of Internal Medicine, Centre Hospitalier Universitaire Vaudois (CHUV) University Hospital, Lausanne, Switzerland.

    • Peter Vollenweider
  200. Division of Biostatistics, Washington University School of Medicine, St. Louis, Missouri, USA.

    • Ingrid B Borecki
  201. Medical Genetics Institute, Cedars-Sinai Medical Center, Los Angeles, California, USA.

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

    • Robert C Kaplan
  203. Carolina Center for Genome Sciences, School of Public Health, University of North Carolina Chapel Hill, Chapel Hill, North Carolina, USA.

    • Kari E North
  204. Department of Medical Genetics, University of Helsinki, Helsinki, Finland.

    • Leena Peltonen
  205. Laboratory of Genetics, National Institute on Aging, Baltimore, Maryland, USA.

    • David Schlessinger
  206. Division of Community Health Sciences, St George's, University of London, London, UK.

    • David P Strachan
  207. Department of Genetics, Harvard Medical School, Boston, Massachusetts, USA.

    • Joel N Hirschhorn
  208. Klinikum Grosshadern, Munich, Germany.

    • H-Erich Wichmann
  209. Ludwig-Maximilians-Universität, Institute of Medical Informatics, Biometry and Epidemiology, Munich, Germany.

    • H-Erich Wichmann
  210. Faculty of Medicine, University of Iceland, Reykjavík, Iceland.

    • Unnur Thorsteinsdottir
    •  & Kari Stefansson
  211. University of Cambridge Metabolic Research Labs, Institute of Metabolic Science Addenbrooke's Hospital, Cambridge, UK.

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

    • Caroline S Fox

Consortia

  1. MAGIC

    On behalf of the MAGIC (Meta-Analyses of Glucose and Insulin-related traits Consortium) investigators.

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Contributions

Writing group: I.B., C.S.F., I.M.H. (lead), C.M.L. (lead), M.I.M., K.L. Mohlke, L.Q., V. Steinthorsdottir, G.T., M.C.Z.

Waist phenotype working group: T.L.A., N.B., I.B., L.A.C., C.M.D., C.S.F., T.B.H., I.M.H., A.U.J., C.M.L. (lead), R.J.F.L., R.M., M.I.M., K.L. Mohlke, L.Q., J.C.R., E.K.S., V. Steinthorsdottir, K. Stefansson, G.T., U.T., C.C.W., T.W., T.W.W., H.E.W., M.C.Z.

Data cleaning and analysis: S.I.B., I.M.H. (lead), E.I., A.U.J., H.L., C.M.L. (lead), R.J.F.L. (lead), J.L., R.M., L.Q., J.C.R., E.K.S., G.T., S.V., M.N.W., E.W., C.J.W., T.W.W., T.W.

Sex-specific analyses: S.I.B., T.E., I.M.H., A.U.J., T.O.K., Z.K., S.L., C.M.L., R.J.F.L., R.M., K.L. Monda, K.E.N., L.Q., J.C.R. (lead), V. Steinthorsdottir, G.T., T.W.W. (lead).

eQTL and expression analyses: S.I.B., A.L.D., C.C.H., J.N.H., F.K., L.M.K., C.M.L., L.L., R.J.F.L., J.L., M.F.M., J.L.M., C.M., G.N., E.E.S., E.K.S., V. Steinthorsdottir, G.T., K.T.Z.

Pathway and CNV analyses: C.M.L., S.A.M., M.I.M., J.N., V. Steinthorsdottir, G.T., B.F.V.

Secondary analyses: S.I.B., I.B.B., N.C., K.E., T.M.F., M.F.F., T.F., M.E.G., J.N.H., E.I., G.L., C.M.L., H.L., R.M., M. Mangino, M.I.M., K.L. Mohlke, D.R.N., J.R.O., S.P., J.R.B.P., J.C.R., A.V.S., E.K.S., P.M.V., M.N.W., C.J.W., R.J.W., E.W., A.R.W., J.Y.

Study-specific analyses: G.R.A., D.A., N.A., T.A., T.L.A., N.B., C.C., P.S.C., L.C., L.A.C., D.I.C., M.N.C., C.M.D., T.E., K.E., E.F., M.F.F., T.F., A.P.G., N.L.G., M.E.G., C. Hayward, N.L.H., I.M.H., J.J.H., A.U.J., Å.J., T. Johnson, J.O.J., J.R.K., M. Kaakinen, K. Kapur, S. Ketkar, J.W.K., P. Kraft, A.T.K., Z.K., J. Kettunen, C. Lamina, R.J.F.L., C. Lecoeur, H.L., M.F.L., C.M.L., J.L., R.W.L., R.M., M. Mangino, B.M., K.L. Monda, A.P.M., N.N., K.E.N., D.R.N., J.R.O., K.K.O., C.O., M.J.P., O. Polasek, I. Prokopenko, N.P., M.P., L.Q., J.C.R., N.W.R., S.R., F.R., N.R.R., C.S., L.J.S., K. Silander, E.K.S., K. Stark, S.S., A.V.S., N.S., U.S., V. Steinthorsdottir, D.P.S., I.S., M.L.T., T.M.T., N.J.T., A.T., G.T., A.U., S.V., V. Vitart, L.V., P.M.V., R.M.W., R.W., R.J.W., S.W., M.N.W., C.C.W., C.J.W., T.W.W., A.R.W., J.Y., J.H.Z., M.C.Z.

Study-specific genotyping: D.A., T.L.A., L.D.A., N.B., I.B., A.J.B., E.B., L.L.B., I.B.B., H.C., D.I.C., I.N.M.D., M. Dei, M.R.E., P.E., K.E., N.B.F., M.F., A.P.G., H.G., C.G., E.J.C.G., C.J.G., T. Hansen, A.L.H., N.H., C. Hayward, A.A.H., J.J.H., F.B.H., D.J.H., J.H., W.I., M.R.J., Å.J., J.O.J., J.W.K., P. Kovacs, A.T.K., H.K.K., J. Kettunen, P. Kraft, R.N.L., C.M.L., R.J.F.L., J.L., M.L.L., M.A.M., M. Mangino, W.L.M., M.I.M., J.B.J.M., M.J.N., M.N., D.R.N., K.K.O., C.O., O. Pedersen, L.P., M.J.P., G.P., A.N.P., N.P., L.Q., N.W.R., F.R., N.R.R., C.S., A.J.S., N.S., A.C.S., M.T., B.T., A.U., G.U., V. Vatin, P.M.V., H.W., P.Z.

Study-specific phenotyping: H.A., P.A., D.A., A.M.A., T.L.A., B.B., S.R.B., R.B., E.B., I.B.B., J.P.B., M. Dörr, C.M.D., P.E., M.F.F., C.S.F., T.M.F., M.F., S.G., J.G., L.C.G., T. Hansen, A.S.H., C. Hengstenberg, A.L.H., A.T.H., K.H.H., A. Hofman, F.B.H., D.J.H., B.I., T.I., T. Jørgensen, P.J., M.R.J., Å.J., A.J., A.L.J., J.O.J., F.K., L.K., J. Kuusisto, K. Kvaloy, R.K., S. Ketkar, J.W.K., I.K., S. Koskinen, V.K., M. Kähönen, P. Kovacs, O.L., R.N.L., B.L., J.L., G.M.L., R.J.F.L., T.L., M. Mangino, M.I.M., C.O., B.M.P., O. Pedersen, C.G.P.P., J.F.P., I. Pichler, K.P., O. Polasek, A.P., L.Q., M.R., I.R., O.R., V. Salomaa, J. Saramies, P.E.H.S., K. Silander, N.J.S., J.H.S., T.D.S., D.P.S., R.S., H.M.S., J. Sinisalo, T.T., A.T., M.U., P.V., C.B.V., L.V., J.V., D.R.W., G.B.W., S.H.W., G.W., J.C.W., A.F.W., L.Z., P.Z.

Study-specific management: G.R.A., A.M.A., B.B., Y.B.S., R.N.B., H.B., J.S.B., S.B., M.B., E.B., D.I.B., I.B.B., J.P.B., M.J.C., F.S.C., L.A.C., G.D., C.M.D., S.E., G.E., P.F., C.S.F., T.M.F., L.C.G., V.G., U.G., M.E.G., T. Hansen, C. Hengstenberg, K.H., A. Hamsten, T.B.H., A.T.H., A. Hofman, F.B.H., D.J.H., B.I., T.I., C.I., T. Jørgensen, M.R.J., A.L.J., F.K., K.T.K., W.H.L.K., R.K., J. Kaprio, M. Kähönen, M.L., D.A.L., L.J.L., C.M.L., R.J.F.L., T.L., M. Marre, T.M., A.M.E.T., K.M., M.I.M., K.L. Mohlke, P.B.M., K.E.N., M.S.N., D.R.N., B.O., C.O., O. Pedersen, L.P., B.W.P., P.P.P., B.M.P., L.J.P., T.Q., A.R., I.R., O.R., P.M.R., V. Salomaa, P.S., D.S., A.R.S., N.S., T.D.S., K. Stefansson, D.P.S., A.C.S., M.S., T.T., J.T., U.T., A.T., M.U., A.U., T.T.V., P.V., H.V., J.V., P.M.V., N.J.W., H.E.W., J.F.W., J.C.W., A.F.W.

Steering committee: G.R.A., T.L.A., I.B., S.I.B., M.B., I.B.B., P.D., C.M.D., C.S.F., T.M.F., L.C.G., T. Haritunians, J.N.H. (chair), D.J.H., E.I., R.K., R.J.F.L., M.I.M., K.L. Mohlke, K.E.N., J.R.O., L.P., D.S., D.P.S., U.T., H.E.W.

Competing interests

I.B. and spouse own stock in Incyte Ltd and GlaxoSmithKline. J.H. is a member of the Scientific Advisory Board, Correlagen, Inc. A.P. is employed by Amgen. K.S., V.S., G.T., U.T. and G.B.W. are employed by deCODE Genetics.

Corresponding authors

Correspondence to Iris M Heid or Mark I McCarthy or Caroline S Fox or Karen L Mohlke or Cecilia M Lindgren.

Supplementary information

PDF files

  1. 1.

    Supplementary Text and Figures

    Supplementary Tables 1–11, Supplementary Figure 1 and Supplementary Note.

Excel files

  1. 1.

    Supplementary Table 11

    3,113 SNPs tagging the 856 CNVs in the HapMap 3 catalog across all HapMap3 populations