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New genetic loci link adipose and insulin biology to body fat distribution

Nature volume 518, pages 187–196 (2015)Cite this article

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Abstract

Body fat distribution is a heritable trait and a well-established predictor of adverse metabolic outcomes, independent of overall adiposity. To increase our understanding of the genetic basis of body fat distribution and its molecular links to cardiometabolic traits, here we conduct genome-wide association meta-analyses of traits related to waist and hip circumferences in up to 224,459 individuals. We identify 49 loci (33 new) associated with waist-to-hip ratio adjusted for body mass index (BMI), and an additional 19 loci newly associated with related waist and hip circumference measures (P < 5 × 10−8). In total, 20 of the 49 waist-to-hip ratio adjusted for BMI loci show significant sexual dimorphism, 19 of which display a stronger effect in women. The identified loci were enriched for genes expressed in adipose tissue and for putative regulatory elements in adipocytes. Pathway analyses implicated adipogenesis, angiogenesis, transcriptional regulation and insulin resistance as processes affecting fat distribution, providing insight into potential pathophysiological mechanisms.

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Figure 1: Regional SNP association plots illustrating the complex genetic architecture at two WHRadjBMI loci.
Figure 2: Gene set enrichment and tissue expression of genes at WHRadjBMI-associated loci (GWAS-only P < 10−5).

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Acknowledgements

We thank the more than 224,000 volunteers who participated in this study. Detailed acknowledgment of funding sources is provided in the Supplementary Note.

Author information

Author notes

  1. Dmitry Shungin, Thomas W. Winkler, Damien C. Croteau-Chonka, Teresa Ferreira, Adam E. Locke and Reedik Mägi: These authors contributed equally to this work.

  2. Iris M. Heid, Ruth J. F. Loos, L. Adrienne Cupples, Andrew P. Morris, Cecilia M. Lindgren and Karen L Mohlke: These authors jointly supervised this work.

Authors and Affiliations

  1. Department of Public Health and Clinical Medicine, Unit of Medicine, Umeå University, 901 87 Umeå, Sweden.,

    Dmitry Shungin & Paul W. Franks

  2. Department of Clinical Sciences, Genetic & Molecular Epidemiology Unit, Lund University Diabetes Center, Skåne University Hosptial, 205 02 Malmö, Sweden.,

    Dmitry Shungin, Frida Renstrom & Paul W. Franks

  3. Department of Odontology, Umeå University, 901 85 Umeå, Sweden.,

    Dmitry Shungin & Ingegerd Johansson

  4. Department of Genetic Epidemiology, Institute of Epidemiology and Preventive Medicine, University of Regensburg, D-93053 Regensburg, Germany.,

    Thomas W. Winkler, Matthias Olden & Iris M. Heid

  5. Department of Genetics, University of North Carolina, Chapel Hill, 27599, North Carolina, USA

    Damien C. Croteau-Chonka, Martin L. Buchkovich, Tamara S. Roman & Karen L Mohlke

  6. Channing Division of Network Medicine, Department of Medicine, Brigham and Women's Hospital and Harvard Medical School, Boston, 02115, Massachusetts, USA

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

  7. Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford OX3 7BN, UK.,

    Teresa Ferreira, Reedik Mägi, Alexander W. Drong, Joshua C. Randall, Anuj Goel, Inga Prokopenko, Thorhildur Juliusdottir, Anubha Mahajan, Nigel W. Rayner, Neil R. Robertson, Åsa K. Hedman, Sarah Keildson, John R. B. Perry, Krina T. Zondervan, Martin Farrall, Hugh Watkins, Mark I. McCarthy, Erik Ingelsson, Andrew P. Morris & Cecilia M. Lindgren

  8. Department of Biostatistics, Center for Statistical Genetics, University of Michigan, Ann Arbor, 48109, Michigan, USA

    Adam E. Locke, Jennifer L. Bragg-Gresham, Anne U. Jackson, Heather M. Stringham, Goncalo R. Abecasis & Michael Boehnke

  9. Estonian Genome Center, University of Tartu, Tartu 51010, Estonia.,

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

  10. Department of Medicine, Atherosclerosis Research Unit, Center for Molecular Medicine, Karolinska Institutet, Stockholm 17176, Sweden.,

    Rona J. Strawbridge, Bengt Sennblad, Per Eriksson, Lasse Folkersen & Anders Hamsten

  11. Divisions of Endocrinology and Genetics and Center for Basic and Translational Obesity Research, Boston Children's Hospital, Boston, 02115, Massachusetts, USA

    Tune H. Pers, Tonu Esko, Sailaja Vedantam, Cameron D. Palmer, Rany M. Salem & Joel N. Hirschhorn

  12. Broad Institute of the Massachusetts Institute of Technology and Harvard University, Cambridge, 02142, Massachusetts, USA

    Tune H. Pers, Tonu Esko, Sailaja Vedantam, Cameron D. Palmer, Karol Estrada, Rany M. Salem, Joel N. Hirschhorn & Cecilia M. Lindgren

  13. Department of Genetics, Harvard Medical School, Boston, 02115, Massachusetts, USA

    Tune H. Pers, Tonu Esko, Rany M. Salem & Joel N. Hirschhorn

  14. Department of Systems Biology, Center for Biological Sequence Analysis, Technical University of Denmark, Lyngby 2800, Denmark.,

    Tune H. Pers

  15. Department of Epidemiology, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina 27599, USA.,

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

  16. Department of Nutrition, Harvard School of Public Health, Boston, 02115, Massachusetts, USA

    Tsegaselassie Workalemahu & David J. Hunter

  17. Department of Biostatistics, Boston University School of Public Health, Boston, 02118, Massachusetts, USA

    Joseph M. W. Wu, Ching-Ti Liu & L. Adrienne Cupples

  18. National Heart, Lung, and Blood Institute, the Framingham Heart Study, Framingham, 01702, Massachusetts, USA

    Nancy L. Heard-Costa, Joanne M. Murabito, Caroline S. Fox & L. Adrienne Cupples

  19. Department of Neurology, Boston University School of Medicine, Boston, 02118, Massachusetts, USA

    Nancy L. Heard-Costa

  20. Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm 17177, Sweden.,

    Ci Song, Tove Fall, Patrik K. E. Magnusson & Nancy L. Pedersen

  21. Science for Life Laboratory, Uppsala University, Uppsala 75185, Sweden.,

    Ci Song, Stefan Gustafsson, Tove Fall, Johan Ärnlöv, Ann-Christine Syvänen, Åsa K. Hedman & Erik Ingelsson

  22. Department of Medical Sciences, Molecular Epidemiology, Uppsala University, Uppsala 75185, Sweden.,

    Ci Song, Stefan Gustafsson, Tove Fall, Johan Ärnlöv, Åsa K. Hedman & Erik Ingelsson

  23. MRC Epidemiology Unit, University of Cambridge School of Clinical Medicine, Institute of Metabolic Science, Cambridge Biomedical Campus, Cambridge CB2 0QQ, UK.,

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

  24. Institute of Social and Preventive Medicine (IUMSP), Centre Hospitalier Universitaire Vaudois (CHUV), Lausanne 1010, Switzerland.,

    Zoltán Kutalik

  25. Swiss Institute of Bioinformatics, Lausanne 1015, Switzerland.,

    Zoltán Kutalik & Toby Johnson

  26. Department of Medical Genetics, University of Lausanne, Lausanne 1005, Switzerland.,

    Zoltán Kutalik & Toby Johnson

  27. Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1SA, UK.,

    Joshua C. Randall, Nigel W. Rayner, Kathleen Stirrups, Panos Deloukas & Inês Barroso

  28. Institute for Medical Informatics, Biometry and Epidemiology (IMIBE), University Hospital Essen, Essen, 45147 Germany.,

    André Scherag, Sonali Pechlivanis & Karl-Heinz Jöckel

  29. Clinical Epidemiology, Integrated Research and Treatment Center, Center for Sepsis Control and Care (CSCC), Jena University Hospital, Jena 07743, Germany.,

    André Scherag

  30. Genetics of Complex Traits, University of Exeter Medical School, University of Exeter, Exeter EX1 2LU, UK.,

    Andrew R. Wood, Dorota Pasko, John R. B. Perry & Timothy M. Frayling

  31. Department of Internal Medicine, Division of Cardiovascular Medicine, University of Michigan, Ann Arbor, 48109, Michigan, USA

    Jin Chen & Cristen J. Willer

  32. Department of Genetics, University Medical Center Groningen, University of Groningen, 9700 RB Groningen, The Netherlands.,

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

  33. Department of Internal Medicine, Division of Gastroenterology, and Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, 48109, Michigan, USA

    Bratati Kahali & Elizabeth K. Speliotes

  34. Department of Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, 48109, Michigan, USA

    Ellen M. Schmidt & Cristen J. Willer

  35. HudsonAlpha Institute for Biotechnology, Huntsville, 35806, Alabama, USA

    Devin Absher & Lindsay L. Waite

  36. Department of Epidemiology, Genetic Epidemiology Unit, Erasmus MC University Medical Center, 3015 GE Rotterdam, The Netherlands.,

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

  37. Telethon Institute for Child Health Research, Centre for Child Health Research, The University of Western Australia, Perth, 6008, Western Australia, Australia

    Denise Anderson

  38. Netherlands Consortium for Healthy Aging (NCHA), Leiden University Medical Center, Leiden 2300 RC, The Netherlands.,

    Marian Beekman, Stefan Böhringer, Joris Deelen, Quinta Helmer, Simon P. Mooijaart, Lisette Stolk & P. Eline Slagboom

  39. Department of Molecular Epidemiology, Leiden University Medical Center, 2300 RC Leiden, The Netherlands.,

    Marian Beekman, Joris Deelen & P. Eline Slagboom

  40. Kidney Epidemiology and Cost Center, University of Michigan, Ann Arbor, 48109, Michigan, USA

    Jennifer L. Bragg-Gresham

  41. Department of Statistics & Biostatistics, Rutgers University, Piscataway, 08854, New Jersey, USA

    Steven Buyske

  42. Department of Genetics, Rutgers University, Piscataway, 08854, New Jersey, USA

    Steven Buyske & Tara C. Matise

  43. Department of Human Genetics, Leiden University Medical Center, 2333 ZC Leiden, The Netherlands.,

    Ayse Demirkan

  44. Center for Complex Disease Genomics, McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University School of Medicine, Baltimore, 21205, Maryland, USA

    Georg B. Ehret & Aravinda Chakravarti

  45. Department of Specialties of Internal Medicine, Cardiology, Geneva University Hospital, Geneva 1211, Switzerland.,

    Georg B. Ehret & François Mach

  46. Department of Genetics, Washington University School of Medicine, St Louis, 63110, Missouri, USA

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

  47. Division of Cardiovacular Medicine, Radcliffe Department of Medicine, University of Oxford, Oxford OX3 9DU, UK.,

    Anuj Goel, Martin Farrall & Hugh Watkins

  48. University Institute for Social and Preventative Medicine, Centre Hospitalier Universitaire Vaudois (CHUV), University of Lausanne, Lausanne 1005, Switzerland.,

    Toby Johnson

  49. Vth Department of Medicine (Nephrology, Hypertensiology, Endocrinology, Diabetology, Rheumatology), Medical Faculty of Mannheim, University of Heidelberg, D-68187 Mannheim, Germany.,

    Marcus E. Kleber & Winfried März

  50. Department of Internal Medicine II, Ulm University Medical Centre, D-89081 Ulm, Germany.,

    Marcus E. Kleber & Wolfgang Koenig

  51. National Institute for Health and Welfare, FI-00271 Helsinki, Finland.,

    Kati Kristiansson, Niina Eklund, Leena Kinnunen, Jaana Lindström, Johan G. Eriksson, Markku Heliövaara, Pekka Jousilahti, Antti M. Jula, Jaakko Kaprio, Kari Kuulasmaa, Satu Männistö, Veikko Salomaa, Markus Perola & Jaakko Tuomilehto

  52. Department of Twin Research and Genetic Epidemiology, King’s College London, London SE1 7EH, UK.,

    Massimo Mangino, Cristina Menni, Alireza Moayyeri, John R. B. Perry & Tim D. Spector

  53. Department of Cardiology, University Medical Center Groningen, University of Groningen, 9700RB Groningen, The Netherlands.,

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

  54. Netherlands Consortium for Healthy Aging (NCHA), 3015GE Rotterdam, The Netherlands.,

    Carolina Medina-Gomez, Marjolein J. Peters, Oscar H. Franco, Albert Hofman, Fernando Rivadeneira, André G. Uitterlinden, Cornelia M. van Duijn & M. Carola Zillikens

  55. Department of Epidemiology, Erasmus MC University Medical Center, 3015GE Rotterdam, The Netherlands.,

    Carolina Medina-Gomez, Oscar H. Franco, Albert Hofman, Jacqueline C. Witteman, Fernando Rivadeneira, André G. Uitterlinden & Cornelia M. van Duijn

  56. Department of Internal Medicine, Erasmus MC University Medical Center, 3015GE Rotterdam, The Netherlands.,

    Carolina Medina-Gomez, Marjolein J. Peters, Karol Estrada, Lisette Stolk, Fernando Rivadeneira, André G. Uitterlinden & M. Carola Zillikens

  57. Oxford Centre for Diabetes, Endocrinology and Metabolism, University of Oxford, Oxford OX3 7LJ, UK.,

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

  58. Department of Genomics of Common Disease, School of Public Health, Imperial College London, Hammersmith Hospital, London W12 0NN, UK.,

    Inga Prokopenko & Philippe Froguel

  59. University of Eastern Finland, FI-70210 Kuopio, Finland.,

    Alena Stančáková

  60. Division of Biostatistics, Washington University School of Medicine, St Louis, 63110, Missouri, USA

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

  61. Translational Gerontology Branch, National Institute on Aging, Baltimore, 21225, Maryland, USA

    Toshiko Tanaka & Luigi Ferrucci

  62. Interfaculty Institute for Genetics and Functional Genomics, University Medicine Greifswald, D-17475 Greifswald, Germany.,

    Alexander Teumer, Georg Homuth & Uwe Völker

  63. Department of Endocrinology, University of Groningen, University Medical Center Groningen, Groningen, 9700 RB, The Netherlands.,

    Jana V. Van Vliet-Ostaptchouk

  64. CNRS UMR 8199, F-59019 Lille, France.,

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

  65. European Genomic Institute for Diabetes, F-59000 Lille, France.,

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

  66. Université de Lille 2, F-59000 Lille, France.,

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

  67. Ealing Hospital NHS Trust, Middlesex UB1 3HW, UK.,

    Weihua Zhang, Jagvir Grewal, Jaspal S. Kooner & John C. Chambers

  68. Department of Epidemiology and Biostatistics, Imperial College London, London W2 1PG, UK.,

    Weihua Zhang, Jagvir Grewal, Marjo-Riitta Jarvelin & John C. Chambers

  69. Institute of Genetic Epidemiology, Helmholtz Zentrum München - German Research Center for Environmental Health, D-85764 Neuherberg, Germany.,

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

  70. School of Health and Social Studies, Dalarna University, SE-791 88 Falun, Sweden.,

    Johan Ärnlöv

  71. PathWest Laboratory Medicine of Western Australia, Nedlands, 6009, Western Australia, Australia

    Gillian M. Arscott, John Beilby & Jennie Hui

  72. Geriatric Unit, Azienda Sanitaria Firenze (ASF), 50125 Florence, Italy.,

    Stefania Bandinelli

  73. Department of Genetics, Texas Biomedical Research Institute, San Antonio, 78227, Texas, USA

    Claire Bellis & John Blangero

  74. Genomics Research Centre, Institute of Health and Biomedical Innovation, Queensland University of Technology, Brisbane, 4001, Queensland, Australia

    Claire Bellis

  75. Department of Medical Sciences, Endocrinology, Diabetes and Metabolism, Uppsala University, Uppsala 75185, Sweden.,

    Christian Berne

  76. Integrated Research and Treatment Center (IFB) Adiposity Diseases, University of Leipzig, D-04103 Leipzig, Germany.,

    Matthias Blüher, Yvonne Böttcher, Peter Kovacs, Anke Tönjes & Michael Stumvoll

  77. Department of Medicine, University of Leipzig, D-04103 Leipzig, Germany.,

    Matthias Blüher, Peter Kovacs, Anke Tönjes & Michael Stumvoll

  78. Department of Medical Statistics and Bioinformatics, Leiden University Medical Center, 2300 RC Leiden, The Netherlands.,

    Stefan Böhringer & Quinta Helmer

  79. Department of Endocrinology, Inserm UMR991, University of Rennes, F-35000 Rennes, France.,

    Fabrice Bonnet

  80. LifeLines Cohort Study, University Medical Center Groningen, University of Groningen, 9700 RB Groningen, The Netherlands.,

    Marcel Bruinenberg & Salome Scholtens

  81. USC-Office of Population Studies Foundation, Inc., University of San Carlos, Cebu City 6000, Philippines.,

    Delia B. Carba & Nanette R. Lee

  82. Department of Biology, Norwegian University of Science and Technology, 7491 Trondheim, Norway.,

    Ida H. Caspersen & Berit Johansen

  83. Nuffield Department of Population Health, Clinical Trial Service Unit and Epidemiological Studies Unit, University of Oxford, Oxford OX3 7LF, UK.,

    Robert Clarke

  84. Information Sciences Institute, University of Southern California, Marina del Rey, 90292, California, USA

    Ewa Deelman

  85. Medical Research Institute, University of Dundee, Ninewells Hospital and Medical School, Dundee DD1 9SY, UK.,

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

  86. Institute for Molecular Medicine, University of Helsinki, FI-00014 Helsinki, Finland.,

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

  87. Medical Genomics and Metabolic Genetics Branch, National Human Genome Research Institute, NIH, Bethesda, Maryland 20892, USA.,

    Michael R. Erdos, Narisu Narisu, Amy J. Swift, Peter S. Chines & Francis S. Collins

  88. Analytic and Translational Genetics Unit, Massachusetts General Hospital and Harvard Medical School, Boston, 02114, Massachusetts, USA

    Karol Estrada

  89. Institute of Clinical Chemistry and Laboratory Medicine, University Medicine Greifswald, D-17475 Greifswald, Germany.,

    Nele Friedrich

  90. Laboratory of Epidemiology and Population Sciences, National Institute on Aging, NIH, Bethesda, Maryland 20892, USA.,

    Melissa E. Garcia & Tamara B. Harris

  91. Department of Public Health and Caring Sciences, Geriatrics, Uppsala University, Uppsala 75185, Sweden.,

    Vilmantas Giedraitis

  92. Division of Cardiovascular Epidemiology, Institute of Environmental Medicine, Karolinska Institutet, Stockholm, Sweden, Stockholm 17177, Sweden.,

    Bruna Gigante, Karin Leander & Ulf de Faire

  93. Division of Research, Kaiser Permanente, Oakland, 94612, California, USA

    Alan S. Go

  94. Service of Therapeutic Education for Diabetes, Obesity and Chronic Diseases, Geneva University Hospital, Geneva CH-1211, Switzerland.,

    Alain Golay

  95. Research Unit of Molecular Epidemiology, Helmholtz Zentrum München - German Research Center for Environmental Health, D-85764 Neuherberg, Germany.,

    Harald Grallert, Thomas Illig & Annette Peters

  96. German Center for Diabetes Research (DZD), D-85764 Neuherberg, Germany.,

    Harald Grallert & Barbara Thorand

  97. Department of Medicine III, University Hospital Carl Gustav Carus, Technische Universität Dresden, D-01307 Dresden, Germany.,

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

  98. Department of Public Health and Clinical Medicine, Unit of Nutritional Research, Umeå University, Umeå 90187, Sweden.,

    Goran Hallmans

  99. Department of Psychiatry, University of Groningen, University Medical Center Groningen, 9700RB Groningen, The Netherlands.,

    Catharina A. Hartman & Albertine J. Oldehinkel

  100. Kuopio Research Institute of Exercise Medicine, FI-70100 Kuopio, Finland.,

    Maija Hassinen, Timo A. Lakka & Rainer Rauramaa

  101. MRC Human Genetics Unit, Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh EH4 2XU, UK.,

    Caroline Hayward

  102. Hjelt Institute Department of Public Health, University of Helsinki, FI-00014 Helsinki, Finland.,

    Kauko Heikkilä & Jaakko Kaprio

  103. Institute of Biomedicine, University of Oulu, FI-90014 Oulu, Finland.,

    Karl-Heinz Herzig

  104. Medical Research Center Oulu and Oulu University Hospital, FI-90014 Oulu, Finland.,

    Karl-Heinz Herzig

  105. Biocenter Oulu, University of Oulu, FI-90014 Oulu, Finland.,

    Karl-Heinz Herzig & Marjo-Riitta Jarvelin

  106. Faculty of Psychology and Education, VU University Amsterdam, 1081BT Amsterdam, The Netherlands.,

    Quinta Helmer

  107. Department of Epidemiology, University Medical Center Groningen, University of Groningen, 9700 RB Groningen, The Netherlands.,

    Hans L. Hillege, Ilja M. Nolte, Judith M. Vonk, Ronald P. Stolk & Harold Snieder

  108. Department of Public Health and General Practice, Norwegian University of Science and Technology, Trondheim 7489, Norway.,

    Oddgeir Holmen & Kristian Hveem

  109. Cardiovascular Genetics Division, Department of Internal Medicine, University of Utah, Salt Lake City, 84108, Utah, USA

    Steven C. Hunt

  110. Center for Medical Sytems Biology, 2300 RC Leiden, The Netherlands.,

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

  111. Institute for Community Medicine, University Medicine Greifswald, D-17475 Greifswald, Germany.,

    Till Ittermann & Wolfgang Hoffmann

  112. Department of Pulmonary Physiology and Sleep Medicine, Nedlands, 6009, Western Australia, Australia

    Alan L. James

  113. School of Medicine and Pharmacology, University of Western Australia, Crawley 6009, Australia.,

    Alan L. James

  114. Department of Dietetics-Nutrition, Harokopio University, 17671 Athens, Greece.,

    Ioanna-Panagiota Kalafati & George Dedoussis

  115. Department of Internal Medicine I, Ulm University Medical Centre, D-89081 Ulm, Germany.,

    Wolfgang Kratzer & Thomas Seufferlein

  116. Division of Genetic Epidemiology, Department of Medical Genetics, Molecular and Clinical Pharmacology, Innsbruck Medical University, 6020 Innsbruck, Austria.,

    Claudia Lamina

  117. Institute of Human Genetics, Helmholtz Zentrum München - German Research Center for Environmental Health, D-85764 Neuherberg, Germany.,

    Peter Lichtner

  118. Department of Medical Sciences, Cardiovascular Epidemiology, Uppsala University, Uppsala 75185, Sweden.,

    Lars Lind & Johan Sundström

  119. Department of Internal Medicine and Clinical Nutrition, Centre for Bone and Arthritis Research, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, Gothenburg 413 45, Sweden.,

    Mattias Lorentzon & Claes Ohlsson

  120. School of Social and Community Medicine, University of Bristol, Bristol BS8 2BN, UK.,

    Wendy L. McArdle & Josine L. Min

  121. Division of Endocrinology, Diabetes and Metabolism, Ulm University Medical Centre, D-89081 Ulm, Germany.,

    Sigrun Merger & Simone Claudi-Boehm

  122. Institute of Molecular and Cell Biology, University of Tartu, Tartu 51010, Estonia.,

    Evelin Mihailov & Andres Metspalu

  123. Farr Institute of Health Informatics Research, University College London, London NW1 2DA, UK,

    Alireza Moayyeri

  124. The Center for Observational Research, Amgen, Inc., Thousand Oaks, 91320, California, USA

    Keri L. Monda

  125. Department of Gerontology and Geriatrics, Leiden University Medical Center, 2300 RC Leiden, The Netherlands.,

    Simon P. Mooijaart

  126. Department of Genomics, Life & Brain Center, University of Bonn, 53127 Bonn, Germany.,

    Thomas W. Mühleisen

  127. Institute of Human Genetics, University of Bonn, 53127 Bonn, Germany.,

    Thomas W. Mühleisen

  128. Istituto di Ricerca Genetica e Biomedica (IRGB), Consiglio Nazionale delle Ricerche, Cagliari, 09042, Sardinia, Italy

    Antonella Mulas & Serena Sanna

  129. Center for Evidence-based Healthcare, University Hospital Carl Gustav Carus, Technische Universität Dresden, D-01307 Dresden, Germany.,

    Gabriele Müller

  130. Department of Medicine I, University Hospital Grosshadern, Ludwig-Maximilians-Universität, D-81377 Munich, Germany.,

    Martina Müller-Nurasyid

  131. Institute of Medical Informatics, Biometry and Epidemiology, Chair of Genetic Epidemiology, Ludwig-Maximilians-Universität, D-81377 Munich, Germany.,

    Martina Müller-Nurasyid & Konstantin Strauch

  132. Deutsches Forschungszentrum für Herz-Kreislauferkrankungen (DZHK) (German Research Centre for Cardiovascular Research), Munich Heart Alliance, D-80636 Munich, Germany.,

    Martina Müller-Nurasyid & Annette Peters

  133. Laboratory of Genetics, National Institute on Aging, Baltimore, 21224, Maryland, USA

    Ramaiah Nagaraja & David Schlessinger

  134. Laboratory of Neurogenetics, National Institute on Aging, National Institutes of Health, Bethesda, 20892, Maryland, USA

    Michael A. Nalls

  135. Hypertension and Related Diseases Centre - AOU, University of Sassari Medical School, Sassari 07100, Italy.,

    Nicola Glorioso & Chiara Troffa

  136. Division of Preventive Medicine, Brigham and Women's Hospital, Boston, 02215, Massachusetts, USA

    Lynda M. Rose, Paul M. Ridker & Daniel I. Chasman

  137. Clinical Institute of Medical and Chemical Laboratory Diagnostics, Medical University of Graz, Graz 8036, Austria.,

    Hubert Scharnagl & Winfried März

  138. Science for Life Laboratory, Karolinska Institutet, Stockholm 171 65, Sweden.,

    Bengt Sennblad

  139. Department of Medicine, University of Washington, Seattle, 98101, Washington, USA

    Colleen M. Sitlani

  140. Icelandic Heart Association, Kopavogur 201, Iceland.,

    Albert Vernon Smith & Vilmundur Gudnason

  141. University of Iceland, Reykjavik 101, Iceland.,

    Albert Vernon Smith & Vilmundur Gudnason

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

    Kathleen Stirrups, Stavroula Kanoni & Panos Deloukas

  143. Department of Medical Sciences, Molecular Medicine, Uppsala University, Uppsala 75144, Sweden.,

    Ann-Christine Syvänen

  144. Department of Public Health Sciences, Stritch School of Medicine, Loyola University of Chicago, Maywood, 61053, Illinois, USA

    Bamidele O. Tayo & Richard S. Cooper

  145. Institute of Epidemiology II, Helmholtz Zentrum München - German Research Center for Environmental Health, Neuherberg, Germany, D-85764 Neuherberg, Germany.,

    Barbara Thorand & Annette Peters

  146. deCODE Genetics, Amgen Inc., Reykjavik 101, Iceland.,

    Gudmar Thorleifsson, Valgerdur Steinthorsdottir, Unnur Thorsteinsdottir & Kari Stefansson

  147. Department of Cardiology, Medical University of Graz, Graz 8036, Austria.,

    Andreas Tomaschitz

  148. Department of Child and Adolescent Psychiatry, Psychology, Erasmus MC University Medical Centre, 3000 CB Rotterdam, The Netherlands.,

    Floor V. A. van Oort

  149. Department of Clinical Chemistry, Ulm University Medical Centre, D-89081 Ulm, Germany.,

    Roman Wennauer

  150. Department of Community Medicine, Faculty of Health Sciences, UiT The Arctic University of Norway, 9037 Tromsø, Norway.,

    Tom Wilsgaard & Inger Njølstad

  151. MRC Unit for Lifelong Health and Ageing at University College London, London WC1B 5JU, UK.,

    Andrew Wong, Ken K. Ong & Diana Kuh

  152. Diabetes Complications Research Centre, Conway Institute, School of Medicine and Medical Sciences, University College Dublin, Dublin 4, Ireland.,

    Eoin P. Brennan

  153. Department of Biomedical Sciences, Seoul National University College of Medicine, Seoul 110-799, Korea.,

    Murim Choi

  154. Department of Molecular Medicine and Surgery, Cardiothoracic Surgery Unit, Karolinska Institutet, Stockholm 17176, Sweden.,

    Anders Franco-Cereceda & Christian Olsson

  155. Department of Medicine, Columbia University College of Physicians and Surgeons, 10032, New York, USA

    Ali G. Gharavi & Krzysztof Kiryluk

  156. Department of Population Medicine, Harvard Pilgrim Health Care Institute, Harvard Medical School, Boston, 02215, Massachusetts, USA

    Marie-France Hivert

  157. Massachusetts General Hospital, Boston, 02114, Massachusetts, USA

    Marie-France Hivert

  158. State Key Laboratory of Medical Genomics, Shanghai Institute of Hematology, Rui Jin Hospital Affiliated with Shanghai Jiao Tong University School of Medicine, Shanghai 200025, China.,

    Jinyan Huang

  159. Department of Epidemiology, Harvard School of Public Health, Boston, 02115, Massachusetts, USA

    Jinyan Huang, Liming Liang, Baoshan Ma & David J. Hunter

  160. NIHR Oxford Biomedical Research Centre, OUH Trust, Oxford OX3 7LE, UK.,

    Fredrik Karpe

  161. Department of Biostatistics, Harvard School of Public Health, Harvard University, Boston, 02115, Massachusetts, USA

    Liming Liang

  162. Department of Genetics, Howard Hughes Medical Institute, Yale University School of Medicine, New Haven, New Haven, Connecticut 06520, USA.,

    Richard P. Lifton

  163. College of Information Science and Technology, Dalian Maritime University, Dalian, 116026, Liaoning, China

    Baoshan Ma

  164. Nephrology Research, Centre for Public Health, Queen’s University of Belfast, Belfast, County Down BT9 7AB, UK.,

    Amy J. McKnight

  165. University of Ottawa Heart Institute, Ottawa K1Y 4W7, Canada.,

    Ruth McPherson

  166. National Heart and Lung Institute, Imperial College London, London SW3 6LY, UK.,

    Miriam F. Moffatt & Jaspal S. Kooner

  167. QIMR Berghofer Medical Research Institute, Brisbane, 4006, Queensland, Australia

    Grant W. Montgomery & Dale R. Nyholt

  168. Section of General Internal Medicine, Boston University School of Medicine, Boston, 02118, Massachusetts, USA

    Joanne M. Murabito

  169. Department of Statistics, University of Oxford, 1 South Parks Road, Oxford OX1 3TG, UK.,

    George Nicholson

  170. MRC Harwell, Harwell Science and Innovation Campus, Harwell OX11 0QG, UK.,

    George Nicholson

  171. Institute of Health and Biomedical Innovation, Queensland University of Technology, Brisbane, 4059, Queensland, Australia

    Dale R. Nyholt

  172. Department of Biomedical Engineering and Computational Science, Aalto University School of Science, FI-00076 Helsinki, Finland.,

    Niina Sandholm

  173. Department of Medicine, Division of Nephrology, Helsinki University Central Hospital, FI-00290 Helsinki, Finland.,

    Niina Sandholm

  174. Folkhälsan Institute of Genetics, Folkhälsan Research Center, FI-00290 Helsinki, Finland.,

    Niina Sandholm

  175. Icahn Institute for Genomics and Multiscale Biology, Icahn School of Medicine at Mount Sinai, New York, 10580, New York, USA

    Eric E. Schadt

  176. Computer Science Department, Tecnológico de Monterrey, Atizapán de Zaragoza, 52926, Mexico.,

    Edgar E. Vallejo

  177. Nuffield Department of Obstetrics & Gynaecology, University of Oxford, Oxford OX3 7BN, UK.,

    Krina T. Zondervan

  178. Institut Pasteur de Lille; INSERM, U744; Université de Lille 2; F-59000 Lille, France.,

    Philippe Amouyel

  179. INSERM, U744,

    Philippe Amouyel

  180. Department of Epidemiology and Public Health, EA3430, University of Strasbourg, Faculty of Medicine, Strasbourg, France

    Dominique Arveiler

  181. Department of Internal Medicine, University Medical Center Groningen, University of Groningen, 9700RB Groningen, The Netherlands.,

    Stephan J. L. Bakker & Ron T. Gansevoort

  182. Pathology and Laboratory Medicine, The University of Western Australia, Perth, 6009, Western Australia, Australia

    John Beilby & Jennie Hui

  183. Cedars-Sinai Diabetes and Obesity Research Institute, Los Angeles, 90048, California, USA

    Richard N. Bergman

  184. Clinical Pharmacology Unit, University of Cambridge, Addenbrooke’s Hospital, Hills Road, Cambridge CB2 2QQ, UK.,

    Morris J. Brown

  185. Department of Medicine, Service of Nephrology, Lausanne University Hospital (CHUV), Lausanne 1005, Switzerland.,

    Michel Burnier

  186. Centre for Population Health Sciences, University of Edinburgh, Teviot Place, Edinburgh EH8 9AG, UK.,

    Harry Campbell, Igor Rudan & James F. Wilson

  187. Center for Human Genetics Research, Vanderbilt University Medical Center, Nashville, 37203, Tennessee, USA

    Dana C. Crawford

  188. Department of Molecular Physiology and Biophysics, Vanderbilt University, Nashville, 37232, Tennessee, USA

    Dana C. Crawford

  189. Department of Public Health and Primary Care, University of Cambridge, Cambridge CB1 8RN, UK.,

    John Danesh & Danish Saleheen

  190. Biological Psychology, VU University Amsterdam, 1081BT Amsterdam, The Netherlands.,

    Eco J. C. de Geus

  191. Institute for Research in Extramural Medicine, Institute for Health and Care Research, VU University, 1081BT Amsterdam, The Netherlands.,

    Eco J. C. de Geus & Brenda W. Penninx

  192. Department of Internal Medicine B, University Medicine Greifswald, D-17475 Greifswald, Germany.,

    Marcus Dörr

  193. DZHK (Deutsches Zentrum für Herz-Kreislaufforschung – German Centre for Cardiovascular Research), partner site Greifswald, D-17475 Greifswald, Germany.,

    Marcus Dörr, Wolfgang Hoffmann & Uwe Völker

  194. Clinic of Cardiology, West-German Heart Centre, University Hospital Essen, 45122 Essen, Germany.,

    Raimund Erbel & Stefan Möhlenkamp

  195. Department of General Practice and Primary Health Care, University of Helsinki, FI-00290 Helsinki, Finland.,

    Johan G. Eriksson

  196. Unit of General Practice, Helsinki University Central Hospital, Helsinki FI-00290, Finland.,

    Johan G. Eriksson

  197. Department of Internal Medicine, University of Pisa, Pisa 56100, Italy.,

    Ele Ferrannini

  198. National Research Council Institute of Clinical Physiology, University of Pisa, Pisa 56124, Italy.,

    Ele Ferrannini

  199. Department of Cardiology, Toulouse University School of Medicine, Rangueil Hospital, 31400 Toulouse, France.,

    Jean Ferrières

  200. UWI Solutions for Developing Countries, The University of the West Indies, Mona, Kingston 7, Jamaica.,

    Terrence Forrester & Colin A. McKenzie

  201. Department of Preventive Medicine, Keck School of Medicine, University of Southern California, Los Angeles, 90089, California, USA

    Christopher A. Haiman

  202. Institute of Biomedical & Clinical Science, University of Exeter, Barrack Road, Exeter EX2 5DW, UK.,

    Andrew T. Hattersley

  203. Center for Biomedicine, European Academy Bozen, Bolzano (EURAC), Bolzano 39100, Italy (affiliated Institute of the University of Lübeck, D-23562 Lübeck, Germany).,

    Andrew A. Hicks & Peter P. Pramstaller

  204. Institute of Cardiovascular Science, University College London, London WC1E 6BT, UK.,

    Aroon D. Hingorani

  205. Centre for Cardiovascular Genetics, Institute Cardiovascular Sciences, University College London, London WC1E 6JJ, UK.,

    Steve E. Humphries

  206. Sansom Institute for Health Research, University of South Australia, Adelaide 5000, South Australia, Australia.,

    Elina Hyppönen

  207. School of Population Health, University of South Australia, Adelaide 5000, South Australia, Australia.,

    Elina Hyppönen

  208. South Australian Health and Medical Research Institute, Adelaide 5000, South Australia, Australia.,

    Elina Hyppönen

  209. Population, Policy, and Practice, University College London Institute of Child Health, London WC1N 1EH, UK.,

    Elina Hyppönen & Chris Power

  210. Hannover Unified Biobank, Hannover Medical School, Hannover, D-30625 Hannover, Germany.,

    Thomas Illig

  211. National Institute for Health and Welfare, FI-90101 Oulu, Finland.,

    Marjo-Riitta Jarvelin

  212. MRC Health Protection Agency (HPA) Centre for Environment and Health, School of Public Health, Imperial College London, London W2 1PG, UK.,

    Marjo-Riitta Jarvelin

  213. Unit of Primary Care, Oulu University Hospital, FI-90220 Oulu, Finland.,

    Marjo-Riitta Jarvelin

  214. Institute of Health Sciences, University of Oulu, FI-90014 Oulu, Finland.,

    Marjo-Riitta Jarvelin

  215. UK Clinical Research Collaboration Centre of Excellence for Public Health (NI), Queens University of Belfast, Belfast BT7 1NN, Northern Ireland, UK.,

    Frank Kee

  216. Institute of Health Sciences, Faculty of Medicine, University of Oulu, FI-90014 Oulu, Finland.,

    Sirkka M. Keinanen-Kiukaanniemi

  217. Unit of Primary Health Care/General Practice, Oulu University Hospital, FI-90220 Oulu, Finland.,

    Sirkka M. Keinanen-Kiukaanniemi

  218. Imperial College Healthcare NHS Trust, London W12 0HS, UK.,

    Jaspal S. Kooner & John C. Chambers

  219. Division of Public Health Sciences, Fred Hutchinson Cancer Research Center, Seattle, 98109, Washington, USA

    Charles Kooperberg & Ulrike Peters

  220. Department of Epidemiology and Public Health, University College London, London WC1E 6BT, UK.,

    Meena Kumari, Claudia Langenberg & Mika Kivimaki

  221. Department of Biological and Social Epidemiology, University of Essex, Wivenhoe Park, Colchester, Essex CO4 3SQ, UK.,

    Meena Kumari

  222. Department of Medicine, Kuopio University Hospital and University of Eastern Finland, FI-70210 Kuopio, Finland.,

    Johanna Kuusisto & Markku Laakso

  223. Department of Physiology, Institute of Biomedicine, University of Eastern Finland, Kuopio Campus, FI-70211 Kuopio, Finland.,

    Timo A. Lakka

  224. Department of Clinical Physiology and Nuclear Medicine, Kuopio University Hospital and University of Eastern Finland, FI-70210 Kuopio, Finland.,

    Timo A. Lakka & Rainer Rauramaa

  225. Epidemiology Program, University of Hawaii Cancer Center, Honolulu, 96813, Hawaii, USA

    Loic Le Marchand

  226. Department of Clinical Chemistry, Fimlab Laboratories and School of Medicine University of Tampere, FI-33520 Tampere, Finland.,

    Terho Lehtimäki

  227. Steno Diabetes Center A/S, Gentofte DK-2820, Denmark.,

    Valeriya Lyssenko

  228. Lund University Diabetes Centre and Department of Clinical Science, Diabetes & Endocrinology Unit, Lund University, Malmö 221 00, Sweden.,

    Valeriya Lyssenko & Leif C. Groop

  229. Institut Universitaire de Cardiologie et de Pneumologie de Québec, Faculty of Medicine, Laval University, Quebec QC G1V 0A6, Canada.,

    André Marette

  230. Institute of Nutrition and Functional Foods, Laval University, Quebec QC G1V 0A6, Canada.,

    André Marette, Marie-Claude Vohl & Louis Pérusse

  231. Department of Biostatistics, University of Washington, Seattle, 98195, Washington, USA

    Barbara McKnight

  232. Department of Respiratory Medicine, Sir Charles Gairdner Hospital, Nedlands, 6009, Western Australia, Australia

    Arthur W. Musk

  233. Epidemiology and Obstetrics & Gynaecology, University of Toronto, Toronto, Ontario M5G 1E2, Canada.,

    Lyle J. Palmer

  234. Genetic Epidemiology & Biostatistics Platform, Ontario Institute for Cancer Research, Toronto, Ontario M5G 0A3, Canada.,

    Lyle J. Palmer

  235. Department of Psychiatry, Neuroscience Campus, VU University Amsterdam, 1081 BT Amsterdam, The Netherlands.,

    Brenda W. Penninx

  236. Department of Neurology, General Central Hospital, Bolzano 39100, Italy.,

    Peter P. Pramstaller

  237. Department of Clinical Physiology and Nuclear Medicine, Turku University Hospital, FI-20521 Turku, Finland.,

    Olli T. Raitakari

  238. Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, FI-20521 Turku, Finland.,

    Olli T. Raitakari

  239. Human Genomics Laboratory, Pennington Biomedical Research Center, Baton Rouge, 70808, Louisiana, USA

    Tuomo Rankinen, Mark A. Sarzynski & Claude Bouchard

  240. Department of Psychiatry, Washington University School of Medicine, St Louis, 63110, Missouri, USA

    D. C. Rao & Treva K. Rice

  241. Harvard Medical School, Boston, 02115, Massachusetts, USA

    Paul M. Ridker & Daniel I. Chasman

  242. Center for Systems Genomics, The Pennsylvania State University, University Park, Pennsylvania 16802, USA.,

    Marylyn D. Ritchie

  243. Croatian Centre for Global Health, Faculty of Medicine, University of Split, 21000 Split, Croatia.,

    Igor Rudan

  244. Department of Cardiovascular Sciences, University of Leicester, Glenfield Hospital, Leicester LE3 9QP, UK.,

    Nilesh J. Samani

  245. National Institute for Health Research (NIHR) Leicester Cardiovascular Biomedical Research Unit, Glenfield Hospital, Leicester LE3 9QP, UK.,

    Nilesh J. Samani

  246. South Carelia Central Hospital, 53130 Lappeenranta, Finland.,

    Jouko Saramies

  247. Paul Langerhans Institute Dresden, German Center for Diabetes Research (DZD), 01307 Dresden, Germany.,

    Peter E. H. Schwarz

  248. Division of Endocrinology, Diabetes and Nutrition, University of Maryland School of Medicine, Baltimore, 21201, Maryland, USA

    Alan R. Shuldiner & Jeffrey R. O’Connell

  249. Program for Personalized and Genomic Medicine, University of Maryland School of Medicine, Baltimore, 21201, Maryland, USA

    Alan R. Shuldiner & Jeffrey R. O’Connell

  250. Geriatric Research and Education Clinical Center, Veterans Administration Medical Center, Baltimore, 21201, Maryland, USA

    Alan R. Shuldiner

  251. Department of Epidemiology, Maastricht University, 6229 HA Maastricht, The Netherlands.,

    Jan A. Staessen

  252. KU Leuven Department of Cardiovascular Sciences, Research Unit Hypertension and Cardiovascular Epidemiology, University of Leuven, B-3000 Leuven, Belgium.,

    Jan A. Staessen

  253. Department of Kinesiology, Laval University, Quebec, QC G1V 0A6, Canada.,

    Angelo Tremblay & Louis Pérusse

  254. Dipartimento di Scienze Farmacologiche e Biomolecolari, Università di Milano & Centro Cardiologico Monzino, Instituto di Ricovero e Cura a Carattere Scientifico, Milan 20133, Italy.,

    Elena Tremoli

  255. Department of Food Science and Nutrition, Laval University, Quebec, QC G1V 0A6, Canada.,

    Marie-Claude Vohl

  256. Department of Internal Medicine, University Hospital (CHUV) and University of Lausanne, 1011, Switzerland.,

    Peter Vollenweider

  257. Department of Nutrition, University of North Carolina, Chapel Hill, 27599, North Carolina, USA

    Linda S. Adair

  258. Institute of Social and Preventive Medicine (IUMSP), Centre Hospitalier Universitaire Vaudois and University of Lausanne, 1010 Lausanne, Switzerland.,

    Murielle Bochud

  259. Ministry of Health, Victoria, Republic of Seychelles

    Murielle Bochud

  260. Lee Kong Chian School of Medicine, Imperial College London and Nanyang Technological University, Singapore, 637553 Singapore, Singapore.,

    Bernhard O. Boehm

  261. Department of Internal Medicine I, Ulm University Medical Centre, D-89081 Ulm, Germany.,

    Bernhard O. Boehm

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

    Mark J. Caulfield & Patricia B. Munroe

  263. Department of Psychiatry and Psychotherapy, University Medicine Greifswald, HELIOS-Hospital Stralsund, D-17475 Greifswald, Germany.,

    Hans-Jörgen Grabe

  264. German Center for Neurodegenerative Diseases (DZNE), Rostock, Greifswald, D-17475 Greifswald, Germany.,

    Hans-Jörgen Grabe

  265. School of Population Health, The University of Western Australia, Nedlands, 6009, Western Australia, Australia

    Jennie Hui

  266. Division of Public Health Sciences, Center for Human Genetics, Wake Forest School of Medicine, Winston-Salem, 27157, North Carolina, USA

    Yongmei Liu

  267. Synlab Academy, Synlab Services GmbH, 68163 Mannheim, Germany.,

    Winfried März

  268. Department of Clinical Genetics, Erasmus MC University Medical Center, 3000 CA Rotterdam, The Netherlands.,

    Ben A. Oostra

  269. Department of Medicine, Stanford University School of Medicine, Palo Alto, 94304, California, USA

    Thomas Quertermous & Themistocles L. Assimes

  270. Finnish Diabetes Association, Kirjoniementie 15, FI-33680 Tampere, Finland.,

    Timo E. Saaristo

  271. Pirkanmaa Hospital District, FI-33521 Tampere, Finland.,

    Timo E. Saaristo

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

    Danish Saleheen

  273. Department of Medicine, University of Pennsylvania, Philadelphia, 19104, Pennsylvania, USA

    Danish Saleheen

  274. Department of Medicine, Helsinki University Central Hospital Heart and Lung Center, Helsinki University Central Hospital, FI-00290 Helsinki, Finland.,

    Juha Sinisalo

  275. Faculty of Medicine, University of Iceland, Reykjavik 101, Iceland.,

    Unnur Thorsteinsdottir & Kari Stefansson

  276. Instituto de Investigacion Sanitaria del Hospital Universario LaPaz (IdiPAZ), 28046 Madrid, Spain.,

    Jaakko Tuomilehto

  277. Diabetes Research Group, King Abdulaziz University, 21589 Jeddah, Saudi Arabia.,

    Jaakko Tuomilehto

  278. Centre for Vascular Prevention, Danube-University Krems, 3500 Krems, Austria.,

    Jaakko Tuomilehto

  279. Department of Public Health and Clinical Nutrition, University of Eastern Finland, FI-70211 Kuopio, Finland.,

    Matti Uusitupa

  280. Research Unit, Kuopio University Hospital, FI-70210 Kuopio, Finland.,

    Matti Uusitupa

  281. Durrer Center for Cardiogenetic Research, Interuniversity Cardiology Institute Netherlands-Netherlands Heart Institute, 3501 DG Utrecht, The Netherlands.,

    Pim van der Harst

  282. Department of Clinical and Experimental Medicine, EPIMED Research Center, University of Insubria, Varese I-21100, Italy.,

    Giovanni Veronesi

  283. Institute of Cellular Medicine, Newcastle University, Newcastle NE1 7RU, UK.,

    Mark Walker

  284. Institute of Medical Informatics, Biometry and Epidemiology, Chair of Epidemiology, Ludwig-Maximilians-Universität, D-85764 Munich, Germany.,

    H-Erich Wichmann

  285. Klinikum Grosshadern, D-81377 Munich, Germany.,

    H-Erich Wichmann

  286. Institute of Epidemiology I, Helmholtz Zentrum München - German Research Center for Environmental Health, Neuherberg, Germany, D-85764 Neuherberg, Germany.,

    H-Erich Wichmann

  287. Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, 20892, Maryland, USA

    Sonja I. Berndt

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

    Panos Deloukas

  289. Department of Epidemiology and Population Health, Albert Einstein College of Medicine, Belfer 1306, New York 10461, USA.,

    Robert C. Kaplan

  290. Division of Population Health Sciences & Education, St George's, University of London, London SW17 0RE, UK.,

    David P. Strachan

  291. Department of Human Genetics, University of Michigan, Ann Arbor, 48109, Michigan, USA

    Cristen J. Willer

  292. Queensland Brain Institute, The University of Queensland, Brisbane 4072, Australia.,

    Peter M. Visscher & Jian Yang

  293. The University of Queensland Diamantina Institute, The Translation Research Institute, Brisbane 4012, Australia.,

    Peter M. Visscher & Jian Yang

  294. Oxford NIHR Biomedical Research Centre, Oxford University Hospitals NHS Trust, Oxford OX3 7LJ, UK.,

    Mark I. McCarthy

  295. Carolina Center for Genome Sciences, University of North Carolina at Chapel Hill, Chapel Hill, 27599, North Carolina, USA

    Kari E. North

  296. University of Cambridge Metabolic Research Laboratories, Institute of Metabolic Science, Addenbrooke’s Hospital, Cambridge CB2 OQQ, UK.,

    Inês Barroso

  297. NIHR Cambridge Biomedical Research Centre, Institute of Metabolic Science, Addenbrooke’s Hospital, Cambridge CB2 OQQ, UK.,

    Inês Barroso

  298. The Charles Bronfman Institute for Personalized Medicine, Icahn School of Medicine at Mount Sinai, New York, 10029, New York, USA

    Ruth J. F. Loos

  299. The Genetics of Obesity and Related Metabolic Traits Program, The Icahn School of Medicine at Mount Sinai, New York, 10029, New York, USA

    Ruth J. F. Loos

  300. The Mindich Child Health and Development Institute, Icahn School of Medicine at Mount Sinai, New York, 10029, New York, USA

    Ruth J. F. Loos

  301. Department of Biostatistics, University of Liverpool, Liverpool L69 3GA, UK.,

    Andrew P. Morris

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