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Rare and low-frequency coding variants alter human adult height

Nature volume 542, pages 186–190 (2017)Cite this article

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Abstract

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

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Figure 1: Variants with a larger effect size on height variation tend to be rarer.
Figure 2: Heat map showing subset of DEPICT gene set enrichment results.
Figure 3: STC2 mutants p.Arg44Leu (R44L) and p.Met86Ile (M86I) show compromised proteolytic inhibition of PAPP-A.

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Acknowledgements

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

Author information

Author notes

  1. Eirini Marouli, Mariaelisa Graff, Carolina Medina-Gomez, Ken Sin Lo, Andrew R. Wood, Troels R. Kjaer, Rebecca S. Fine and Yingchang Lu: These authors contributed equally to this work.

  2. Claus Oxvig, Zoltán Kutalik, Fernando Rivadeneira, Ruth J. F. Loos, Timothy M. Frayling, Joel N. Hirschhorn, Panos Deloukas and Guillaume Lettre: These authors jointly supervised this work.

Authors and Affiliations

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

    Eirini Marouli, Kathleen E. Stirrups, Mark J. Caulfield, Stavroula Kanoni, Patricia B. Munroe, Ioanna Ntalla, Helen R. Warren & Panos Deloukas

  2. Department of Epidemiology, University of North Carolina, Chapel Hill, 27514, North Carolina, USA

    Mariaelisa Graff, Heather M. Highland, Anne E. Justice, Kristin L. Young & Angela L. Mazul

  3. Department of Epidemiology, Erasmus Medical Center, Rotterdam, 3015 GE, The Netherlands

    Carolina Medina-Gomez, M. Arfan Ikram, André G. Uitterlinden, Cornelia M. van Duijn & Fernando Rivadeneira

  4. Department of Internal Medicine, Erasmus Medical Center, Rotterdam, 3015 GE, The Netherlands

    Carolina Medina-Gomez, Linda Broer, André G. Uitterlinden & Fernando Rivadeneira

  5. Montreal Heart Institute, Montreal, H1T 1C8, Quebec, Canada

    Ken Sin Lo, Valérie Turcot, Simon de Denus, John D. Rioux, Jean-Claude Tardif & Guillaume Lettre

  6. Genetics of Complex Traits, University of Exeter Medical School, University of Exeter, Exeter, EX2 5DW, UK

    Andrew R. Wood, Katherine S. Ruth, Hanieh Yaghootkar & Timothy M. Frayling

  7. Department of Molecular Biology and Genetics, Aarhus University, Aarhus, 8000, Denmark

    Troels R. Kjaer & Claus Oxvig

  8. Broad Institute of MIT and Harvard, Cambridge, 02142, Massachusetts, USA

    Rebecca S. Fine, Tõnu Esko, Manuel A. Rivas, Sailaja Vedantam, Daniel I. Chasman, Aniruddh P. Patel, Sekar Kathiresan & Joel N. Hirschhorn

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

    Rebecca S. Fine & Sailaja Vedantam

  10. Division of Endocrinology and Center for Basic and Translational Obesity Research, Boston Children’s Hospital, Boston, 02115, Massachusetts, USA

    Rebecca S. Fine, Tõnu Esko, Sailaja Vedantam & Joel N. Hirschhorn

  11. Division of Epidemiology, Department of Medicine, Vanderbilt-Ingram Cancer Center, Vanderbilt Epidemiology Center, Vanderbilt University School of Medicine, Nashville, 37203, Tennessee, USA

    Yingchang Lu

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

    Yingchang Lu, Claudia Schurmann, Erwin P. Bottinger & Ruth J. F. Loos

  13. The Genetics of Obesity and Related Metabolic Traits Program, Ichan School of Medicine at Mount Sinai, New York, 10069, New York, USA

    Yingchang Lu, Claudia Schurmann & Ruth J. F. Loos

  14. Human Genetics Center, The University of Texas School of Public Health, The University of Texas Graduate School of Biomedical Sciences at Houston, The University of Texas Health Science Center at Houston, Houston, 77030, Texas, USA

    Heather M. Highland

  15. Institute of Social and Preventive Medicine, Lausanne University Hospital, Lausanne, 1010, Switzerland

    Sina Rüeger & Zoltán Kutalik

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

    Sina Rüeger, David Lamparter, Sven Bergmann & Zoltán Kutalik

  17. deCODE Genetics/Amgen inc., Reykjavik, 101, Iceland

    Gudmar Thorleifsson, Valgerdur Steinthorsdottir, Unnur Thorsteinsdottir & Kari Stefansson

  18. Department of Computational Biology, University of Lausanne, Lausanne, 1011, Switzerland

    David Lamparter & Sven Bergmann

  19. Department of Haematology, University of Cambridge, Cambridge, CB2 0PT, UK

    Kathleen E. Stirrups

  20. Department of Genetic Epidemiology, University of Regensburg, Regensburg, D-93051, Germany

    Thomas W. Winkler, Mathias Gorski & Iris M. Heid

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

    Tõnu Esko & Andres Metspalu

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

    Tugce Karaderi, Anubha Mahajan, Wei Gan, Hidetoshi Kitajima, Andrew P. Morris, Neil Robertson, Lorraine Southam, Mark I. McCarthy & Cecilia M. Lindgren

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

    Adam E. Locke, Goncalo Abecasis, Shuang Feng, Anne U. Jackson, Xueling Sim, Heather M. Stringham & Michael Boehnke

  24. McDonnell Genome Institute, Washington University School of Medicine, Saint Louis, 63108, Missouri, USA

    Adam E. Locke

  25. Department of Cardiovascular Sciences, Univeristy of Leicester, Glenfield Hospital, Leicester, LE3 9QP, UK

    Nicholas G. D. Masca, Christopher P. Nelson & Nilesh J. Samani

  26. NIHR Leicester Cardiovascular Biomedical Research Unit, Glenfield Hospital, Leicester, LE3 9QP, UK

    Nicholas G. D. Masca, Christopher P. Nelson & Nilesh J. Samani

  27. Center for Diabetes Research, Wake Forest School of Medicine, Winston-Salem, 27157, North Carolina, USA

    Maggie C. Y. Ng, Poorva Mudgal, Donald W. Bowden & Amanda J. Cox

  28. Center for Genomics and Personalized Medicine Research, Wake Forest School of Medicine, Winston-Salem, 27157, North Carolina, USA

    Maggie C. Y. Ng, Donald W. Bowden & Amanda J. Cox

  29. Nuffield Department of Clinical Medicine, Oxford, OX3 7BN, UK

    Manuel A. Rivas

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

    Xiuqing Guo, Yii-Der Ida Chen, Yucheng Jia, Yeheng Liu, Kevin Sandow, Kent D. Taylor, Jie Yao & Jerome I Rotter

  31. Netherlands Comprehensive Cancer Organisation, Utrecht, 3501 DB, The Netherlands

    Katja K. Aben

  32. Radboud University Medical Center, Nijmegen, 6500 HB, The Netherlands

    Katja K. Aben, Tessel E. Galesloot, Lambertus A. Kiemeney, Kirsten B. Kluivers & Sita H. Vermeulen

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

    Linda S. Adair

  34. Centre for Control of Chronic Diseases (CCCD), Dhaka, 1212, Bangladesh

    Dewan S. Alam

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

    Eva Albrecht, Iris M. Heid & Martina Müller-Nurasyid

  36. The Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, 2100, Denmark

    Kristine H. Allin, Emil V. Appel, Jette Bork-Jensen, Niels Grarup, Torben Hansen, Mette Hollensted, Oluf Pedersen, Henrik Vestergaard & Tune H. Pers

  37. Department of Family Medicine & Public Health, University of California, San Diego, La Jolla, 92093, California, USA

    Matthew Allison

  38. INSERM U1167, Lille, F-59019, France

    Philippe Amouyel

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

    Philippe Amouyel

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

    Philippe Amouyel

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

    Dominique Arveiler

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

    Dominique Arveiler

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

    Folkert W. Asselbergs & Jessica van Setten

  44. Durrer Center for Cardiogenetic Research, ICIN-Netherlands Heart Institute, Utrecht, The Netherlands

    Folkert W. Asselbergs

  45. Institute of Cardiovascular Science, Faculty of Population Health Sciences, University College London, London, UK

    Folkert W. Asselbergs

  46. Zilber School of Public Health, University of Wisconsin-Milwaukee, Milwaukee, 53201, Wisconsin, USA

    Paul L. Auer

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

    Beverley Balkau

  48. Department of Nephrology, University Hospital Regensburg, Regensburg, 93042, Germany

    Bernhard Banas, Carsten A. Böger & Mathias Gorski

  49. Department of Cardiology, Rigshospitalet, Copenhagen University Hospital, Copenhagen, 2100, Denmark

    Lia E. Bang

  50. Department of Clinical Biochemistry, Herlev and Gentofte Hospital, Copenhagen University Hospital, Herlev, 2730, Denmark

    Marianne Benn, Pia R. Kamstrup, Sune F. Nielsen, Børge G. Nordestgaard & Anette Varbo

  51. Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, 2200, Denmark

    Marianne Benn, Ruth Frikke-Schmidt, Torben Jørgensen, Allan Linneberg, Sune F. Nielsen, Børge G. Nordestgaard, Anne Tybjaerg-Hansen & Anette Varbo

  52. Department of Epidemiology, School of Public Health, University of Michigan, Ann Arbor, 48109, Michigan, USA

    Lawrence F. Bielak, Min A. Jhun, Sharon L. R. Kardia, Jennifer A. Smith & Wei Zhao

  53. IFB Adiposity Diseases, University of Leipzig, Leipzig, 04103, Germany

    Matthias Blüher, Peter Kovacs & Michael Stumvoll

  54. Department of Medicine, University of Leipzig, Leipzig, 04103, Germany

    Matthias Blüher & Michael Stumvoll

  55. Department of Epidemiology, German Institute of Human Nutrition Potsdam-Rehbruecke (DIfE), Nuthetal, 14558, Germany

    Heiner Boeing

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

    Eric Boerwinkle & Megan L. Grove

  57. Human Genome Sequencing Center, Baylor College of Medicine, Houston, 77030, Texas, USA

    Eric Boerwinkle

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

    Lori L. Bonnycastle & Francis S. Collins

  59. Julius Center for Health Sciences and Primary Care, University Medical Center Utrecht, Utrecht, The Netherlands

    Michiel L. Bots & Paul I. W. de Bakker

  60. Department of Biochemistry, Wake Forest School of Medicine, Winston-Salem, 27157, North Carolina, USA

    Donald W. Bowden

  61. Department of Clinical Biochemistry, Lillebaelt Hospital, Vejle, 7100, Denmark

    Ivan Brandslund

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

    Ivan Brandslund

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

    Gerome Breen

  64. Marshfield Clinic Research Foundation, Marshfield, 54449, Wisconsin, USA

    Murray H. Brilliant, Peggy L. Peissig & Thomas N. Person

  65. Department of Medicine, University of Washington, Seattle, 98195, Washington, USA

    Amber A. Burt, Gail P. Jarvik & Eric B. Larson

  66. Department of Public Health and Primary Care, MRC/BHF Cardiovascular Epidemiology Unit, University of Cambridge, Cambridge, CB1 8RN, UK

    Adam S. Butterworth, Rajiv Chowdhury, John Danesh, Emanuele Di Angelantonio, Joanna M. M. Howson, Praveen Surendran & Robin Young

  67. NIHR Blood and Transplant Research Unit in Donor Health and Genomics, University of Cambridge, Cambridge, CB1 8RN, UK

    Adam S. Butterworth, John Danesh & Emanuele Di Angelantonio

  68. The Sigfried and Janet Weis Center for Research, Danville, 17822, Pennsylvania, USA

    David J. Carey, Helena Kuivaniemi & Gerard Tromp

  69. NIHR Barts Cardiovascular Research Unit, Barts and The London School of Medicine & Dentistry, Queen Mary University, London, EC1M 6BQ, UK

    Mark J. Caulfield, Patricia B. Munroe & Helen R. Warren

  70. Department of Cardiology, London North West Healthcare NHS Trust, Ealing Hospital, Middlesex, UB1 3HW, UK

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

  71. Department of Epidemiology and Biostatistics, School of Public Health, Imperial College London, London, W2 1PG, UK

    John C. Chambers, Evangelos Evangelou, Marcelo P. Segura-Lepe & Weihua Zhang

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

    John C. Chambers & Jaspal S. Kooner

  73. Division of Genetics, Brigham and Women's Hospital and Harvard Medical School, Boston, 02115, Massachusetts, USA

    Daniel I. Chasman

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

    Daniel I. Chasman, Audrey Y. Chu & Paul M. Ridker

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

    Daniel I. Chasman, Aniruddh P. Patel, Paul M. Ridker & Sekar Kathiresan

  76. Medical Department, Lillebaelt Hospital, Vejle, 7100, Denmark

    Cramer Christensen

  77. NHLBI Framingham Heart Study, Framingham, 01702, Massachusetts, USA

    Audrey Y. Chu & Nancy L. Heard-Costa

  78. Department of Medical, Surgical and Health Sciences, University of Trieste, Trieste, 34100, Italy

    Massimiliano Cocca

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

    James P. Cook & Andrew P. Morris

  80. Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, EH8 9JZ, UK

    Janie Corley, Gail Davies, Ian J. Deary & John M. Starr

  81. Department of Psychology, University of Edinburgh, Edinburgh, EH8 9JZ, UK

    Janie Corley, Gail Davies, Ian J. Deary & Alison Pattie

  82. Department of Human Genetics, Radboud University Medical Center, Nijmegen, 6500 HB, The Netherlands

    Jordi Corominas Galbany

  83. Menzies Health Institute Queensland, Griffith University, Southport, Queensland, Australia

    Amanda J. Cox

  84. Diamantina Institute, University of Qeensland, Brisbane, 4072, Queensland, Australia

    Gabriel Cuellar-Partida

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

    Gabriel Cuellar-Partida, Scott D. Gordon, Nicholas G. Martin & Yadav Sapkota

  86. Wellcome Trust Sanger Institute, Wellcome Genome Campus, Hinxton, CB10 1SA, Cambridge, UK

    John Danesh, Gaëlle Marenne, Lorraine Southam & Eleftheria Zeggini

  87. Department of Medicine, British Heart Foundation, Cambridge Centre of Excellence, University of Cambridge, Cambridge, CB2 0QQ, UK

    John Danesh

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

    Paul I. W. de Bakker

  89. Department of Vascular Surgery, Division of Surgical Specialties, University Medical Center Utrecht, Utrecht, 3584 CX, The Netherlands

    Gert J. de Borst

  90. Faculty of Pharmacy, Université de Montréal, Montreal, H3T 1J4, Quebec, Canada

    Simon de Denus

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

    Mark C. H. de Groot

  92. Division Pharmacoepidemiology & Clinical Pharmacology, Utrecht Institute for Pharmaceutical Sciences, Utrecht University, Utrecht, 3508 TB, The Netherlands

    Mark C. H. de Groot

  93. Department of Clinical Epidemiology, Leiden University Medical Center, Leiden, 2300 RC, The Netherlands

    Renée de Mutsert, Ruifang Li-Gao & Dennis O. Mook-Kanamori

  94. Department of Nutrition and Dietetics, School of Health Science and Education, Harokopio University, Athens, 17671, Greece

    George Dedoussis & Aliki-Eleni Farmaki

  95. Division of Epidemiology & Community Health, School of Public Health, University of Minnesota, Minneapolis, 55454, Minnesota, USA

    Ellen W. Demerath

  96. Department of Ophthalmology, Radboud University Medical Center, Nijmegen, 6500 HB, The Netherlands

    Anneke I. den Hollander & Carel B. Hoyng

  97. Department of Public Health and Primary Care, Centre for Cancer Genetic Epidemiology, University of Cambridge, Cambridge, CB1 8RN, UK

    Joe G. Dennis, Douglas F. Easton & Deborah J. Thompson

  98. Institute of Cardiovascular Science, University College London, London, WC1E 6JF, UK

    Fotios Drenos

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

    Fotios Drenos

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

    Mengmeng Du

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

    Mengmeng Du

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

    Alison M. Dunning, Douglas F. Easton, Ailith Pirie & Jonathan P. Tyrer

  103. Department of Medicine, Oulu University Hospital, Oulu, 90029, Finland

    Tapani Ebeling

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

    Tapani Ebeling

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

    Todd L. Edwards & Ayush Giri

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

    Patrick T. Ellinor, Jose C. Florez, Paul L. Huang, Steven A. Lubitz, Alisa K. Manning, Aniruddh P. Patel, Gina M. Peloso, Suthesh Sivapalaratnam & Sekar Kathiresan

  107. Medical and Population Genetics Program, Broad Institute, Cambridge, 02141, Massachusetts, USA

    Patrick T. Ellinor, Jose C. Florez, Steven A. Lubitz, Alisa K. Manning & Gina M. Peloso

  108. Department of Epidemiology and Biostatistics, MRC-PHE Centre for Environment and Health, School of Public Health, Imperial College London, London, W2 1PG, UK

    Paul Elliott

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

    Evangelos Evangelou

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

    Jessica D. Faul

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

    Mary F. Feitosa & Ingrid B. Borecki

  112. CNR Institute of Clinical Physiology, Pisa, Italy

    Ele Ferrannini

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

    Ele Ferrannini

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

    Marco M. Ferrario

  115. Toulouse University School of Medicine, Toulouse, 50032 31059, TSA, France

    Jean Ferrieres

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

    Jose C. Florez & Alisa K. Manning

  117. University of Glasgow, Glasgow, G12 8QQ, UK

    Ian Ford, Chris J. Packard, Sandosh Padmanabhan, Naveed Sattar & Robin Young

  118. Institute of Molecular Medicine, The University of Texas Health Science Center at Houston, Houston, 77030, Texas, USA

    Myriam Fornage & Li-An Lin

  119. Department of Clinical Sciences, Genetic and Molecular Epidemiology Unit, Lund University, Malmö, SE-20502, Sweden

    Paul W. Franks, Frida Renström & Tibor V. Varga

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

    Paul W. Franks

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

    Paul W. Franks

  122. Department of Clinical Biochemistry, Rigshospitalet, Copenhagen University Hospital, Copenhagen, 2100, Denmark

    Ruth Frikke-Schmidt & Anne Tybjaerg-Hansen

  123. Department of Medical Sciences, University of Trieste, Trieste, 34137, Italy

    Ilaria Gandin, Paolo Gasparini, Giorgia Girotto & Anna Morgan

  124. Division of Experimental Genetics, Sidra Medical and Research Center, Doha, 26999, Qatar

    Paolo Gasparini, Giorgia Girotto & Diego Vozzi

  125. Department of Public Health, Geriatrics, Uppsala University, Uppsala, 751 85, Sweden

    Vilmantas Giedraitis

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

    Penny Gordon-Larsen & Kathleen Mullan Harris

  127. Department of Nutrition, Gillings School of Global Public Health, University of North Carolina, Chapel Hill, 27514, North Carolina, USA

    Penny Gordon-Larsen

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

    Vilmundur Gudnason, Albert Vernon Smith, Unnur Thorsteinsdottir & Kari Stefansson

  129. Icelandic Heart Association, Kopavogur, 201, Iceland

    Vilmundur Gudnason & Albert Vernon Smith

  130. Department of Medical Sciences, Molecular Epidemiology and Science for Life Laboratory, Uppsala University, Uppsala, 751 41, Sweden

    Stefan Gustafsson & Erik Ingelsson

  131. Department of Sociology, University of North Carolina, Chapel Hill, 27514, North Carolina, USA

    Kathleen Mullan Harris

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

    Tamara B. Harris

  133. University of Exeter Medical School, University of Exeter, Exeter, EX2 5DW, UK

    Andrew T. Hattersley

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

    Caroline Hayward, Jonathan Marten & Veronique Vitart

  135. Biodemography of Aging Research Unit, Social Science Research Institute, Duke University, Durham, 27708, North Carolina, USA

    Liang He

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

    Liang He, Kauko Heikkilä & Jaakko Kaprio

  137. Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, FI-00014, Finland

    Kauko Heikkilä & Jaakko Kaprio

  138. Department of Pediatrics, Haukeland University Hospital, Bergen, 5021, Norway

    Øyvind Helgeland & Pål R. Njølstad

  139. Department of Clinical Science, KG Jebsen Center for Diabetes Research, University of Bergen, Bergen, 5020, Norway

    Øyvind Helgeland, Stefan Johansson & Pål R. Njølstad

  140. Department of Cardiology, Heart Center, Tampere University Hospital, Tampere, 33521, Finland

    Jussi Hernesniemi

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

    Jussi Hernesniemi, Terho Lehtimäki & Leo-Pekka Lyytikäinen

  142. Department of Clinical Chemistry, University of Tampere School of Medicine, Tampere, 33014, Finland

    Jussi Hernesniemi, Terho Lehtimäki & Leo-Pekka Lyytikäinen

  143. Centre for Eye Research Australia, Royal Victorian Eye and Ear Hospital, University of Melbourne, Melbourne, 3002, Victoria, Australia

    Alex W. Hewitt

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

    Alex W. Hewitt & David A. Mackey

  145. Menzies Research Institute Tasmania, University of Tasmania, Hobart, 7000, Tasmania, Australia

    Alex W. Hewitt

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

    Lynne J. Hocking

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

    Lynne J. Hocking

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

    Oddgeir L. Holmen

  149. Department of Vascular Medicine, AMC, Amsterdam, 1105 AZ, The Netherlands

    G. Kees Hovingh

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

    Kristian Hveem

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

    M. Arfan Ikram

  152. Department of Radiology, Erasmus Medical Center, Rotterdam, 3015 GE, The Netherlands

    M. Arfan Ikram

  153. Department of Medicine, Division of Cardiovascular Medicine, Stanford University School of Medicine, Stanford, 94305, California, USA

    Erik Ingelsson

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

    Jan-Håkan Jansson & Olov Rolandsson

  155. Research Unit Skellefteå, Skellefteå, SE-93141, Sweden

    Jan-Håkan Jansson

  156. Department of Genome Sciences, University of Washington, Seattle, 98195, Washington, USA

    Gail P. Jarvik

  157. The Copenhagen City Heart Study, Frederiksberg Hospital, Frederiksberg, 2000, Denmark

    Gorm B. Jensen

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

    Xuejuan Jiang

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

    Xuejuan Jiang & Rohit Varma

  160. Center for Medical Genetics and Molecular Medicine, Haukeland University Hospital, Bergen, 5021, Norway

    Stefan Johansson

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

    Marit E. Jørgensen

  162. Steno Diabetes Center, Gentofte, 2820, Denmark

    Marit E. Jørgensen

  163. Aalborg University, Aalborg, DK-9000, Denmark

    Torben Jørgensen

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

    Torben Jørgensen, Allan Linneberg & Betina H. Thuesen

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

    Pekka Jousilahti, Jaakko Kaprio, Jukka Kontto, Jaana Lindström, Satu Männistö, Markus Perola, Veikko Salomaa & Jaakko Tuomilehto

  166. Department of Cardiology, Leiden University Medical Center, Leiden, 2333, The Netherlands

    J. Wouter Jukema & Stella Trompet

  167. The Interuniversity Cardiology Institute of the Netherlands, Utrecht, 2333, The Netherlands

    J. Wouter Jukema

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

    Bratati Kahali, Elizabeth K. Speliotes, Wei Zhou & Cristen J. Willer

  169. Department of Internal Medicine, University of Michigan, Ann Arbor, 48109, Michigan, USA

    Bratati Kahali, Elizabeth K. Speliotes, Wei Zhou & Cristen J. Willer

  170. Division of Gastroenterology, University of Michigan, Ann Arbor, 48109, Michigan, USA

    Bratati Kahali & Elizabeth K. Speliotes

  171. Department of Psychiatry, Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht, 3584 CG, The Netherlands

    René S. Kahn & Roel A. Ophoff

  172. Department of Clinical Physiology, University of Tampere School of Medicine, Tampere, 33014, Finland

    Mika Kähönen

  173. Echinos Medical Centre, Echinos, Greece

    Maria Karaleftheri

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

    Fredrik Karpe, Matt Neville, Katharine R. Owen, Neil Robertson & Mark I. McCarthy

  175. Oxford NIHR Biomedical Research Centre, Oxford University Hospitals Trust, Oxford, OX3 7LE, UK

    Fredrik Karpe, Matt Neville, Katharine R. Owen & Mark I. McCarthy

  176. UKCRC Centre of Excellence for Public Health Research, Queens University Belfast, Belfast, BT12 6BJ, UK

    Frank Kee

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

    Renske Keeman & Marjanka K. Schmidt

  178. Department of Restorative Dentistry, Periodontology and Endodontology, University Medicine Greifswald, Greifswald, 17475, Germany

    Thomas Kocher

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

    Pirjo Komulainen, Timo A. Lakka & Rainer Rauramaa

  180. National Heart and Lung Institute, Imperial College London, Hammersmith Hospital Campus, London, W12 0NN, UK

    Jaspal S. Kooner

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

    Charles Kooperberg & Alex P. Reiner

  182. German Center for Diabetes Research, München-Neuherberg, 85764, Germany

    Jennifer Kriebel, Karina Meidtner & Matthias B. Schulze

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

    Jennifer Kriebel

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

    Jennifer Kriebel

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

    Helena Kuivaniemi

  186. CHU Nantes, Service de Génétique Médicale, Nantes, 44093, France

    Sébastien Küry

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

    Johanna Kuusisto & Markku Laakso

  188. Institute for Maternal and Child Health - IRCCS ‘Burlo Garofolo’, Trieste, 34137, Italy

    Martina La Bianca, Antonietta Robino & Sheila Ulivi

  189. Institute of Biomedicine & Physiology, University of Eastern Finland, Kuopio, 70210, Finland

    Timo A. Lakka

  190. Department of Genetics, University of North Carolina, Chapel Hill, North, 27514, Carolina, USA

    Ethan M. Lange, Leslie A. Lange, Karen L. Mohlke & Ying Wu

  191. Department of Biostatistical Sciences and Center for Public Health Genomics, Wake Forest School of Medicine, Winston-Salem, North, 27157, Carolina, USA

    Carl D. Langefeld

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

    Claudia Langenberg, Jian'an Luan, John R. B. Perry, Robert A. Scott, Nicholas J. Wareham, Sara M. Willems & Jing Hua Zhao

  193. Group Health Research Institute, Seattle, 98101, Washington, USA

    Eric B. Larson

  194. Department of Health Services, University of Washington, Seattle, 98101, Washington, USA

    Eric B. Larson

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

    I-Te Lee

  196. School of Medicine, National Yang-Ming University, Taipei, 112, Taiwan

    I-Te Lee

  197. School of Medicine, Chung Shan Medical University, Taichung, 402, Taiwan

    I-Te Lee

  198. Division of Preventive Medicine University of Alabama at Birmingham, Birmingham, 35205, Alabama, USA

    Cora E. Lewis

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

    Huaixing Li, Xu Lin, Feijie Wang, Yiqin Wang, Pang Yao & He Zheng

  200. Department of Medicine, Division of Cardiovascular Medicine, Stanford University School of Medicine, Palo Alto, 94304, California, USA

    Jin Li

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

    Honghuang Lin

  202. Uppsala University, Uppsala, 75185, Sweden

    Lars Lind

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

    Allan Linneberg

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

    Yongmei Liu

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

    Artitaya Lophatananon

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

    Pamela A. F. Madden

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

    Karina Meidtner & Matthias B. Schulze

  208. Centre for Vision Research and Department of Ophthalmology, Westmead Millennium Institute of Medical Research, University of Sydney, Sydney, 2022, New South Wales, Australia

    Paul Mitchell

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

    Dennis O. Mook-Kanamori

  210. Centre for Global Health Research, Usher Institute of Population Health Sciences and Informatics, University of Edinburgh, Edinburgh, EH8 9AG, UK

    Andrew D. Morris, Ozren Polasek & Igor Rudan

  211. Department of Medicine I, Ludwig-Maximilians-Universität, Munich, 81377, Germany

    Martina Müller-Nurasyid

  212. DZHK (German Centre for Cardiovascular Research), partner site Munich Heart Alliance, Munich, 80802, Germany

    Martina Müller-Nurasyid

  213. Laboratory of Neurogenetics, National Institute on Aging, NIH, Bethesda, 20892, Maryland, USA

    Mike A. Nalls

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

    Matthias Nauck

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

    Matthias Nauck

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

    Kjell Nikus

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

    Jeffrey R. O'Connel, James A. Perry & Laura M. Yerges-Armstrong

  218. Department of Medicine, Tampere University Hospital, Tampere, 33521, Finland

    Heikki Oksa

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

    Loes M. Olde Loohuis & Roel A. Ophoff

  220. Pat Macpherson Centre for Pharmacogenetics and Pharmacogenomics, Medical Research Institute, Ninewells Hospital and Medical School, Dundee, DD1 9SY, UK

    Colin N. A. Palmer

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

    Gerard Pasterkamp

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

    Gerard Pasterkamp & Sander W. van der Laan

  223. School of Women’s and Infants’ Health, The University of Western Australia, Perth, 6009, Western Australia, Australia

    Craig E. Pennell & Carol A. Wang

  224. University of Helsinki, Institute for Molecular Medicine (FIMM) and Diabetes and Obesity Research Program, Helsinki, FI00014, Finland

    Markus Perola

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

    Markus Perola

  226. School of Medicine, University of Split, Split, 21000, Croatia

    Ozren Polasek

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

    Danielle Posthuma

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

    Danielle Posthuma

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

    Olli T. Raitakari

  230. Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku, 20520, Finland

    Olli T. Raitakari

  231. Centre for Non-Communicable Diseases, Karachi, Pakistan

    Asif Rasheed & Danish Saleheen

  232. Department of Clinical Physiology and Nuclear Medicine, Kuopio University Hospital, Kuopio, 70210, Finland

    Rainer Rauramaa

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

    Dermot F. Reilly

  234. Department of Epidemiology, University of Washington, Seattle, 98195, Washington, USA

    Alex P. Reiner

  235. Department of Biobank Research, Umeå University, Umeå, SE-90187, Sweden

    Frida Renström

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

    Paul M. Ridker

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

    John D. Rioux, Jean-Claude Tardif & Guillaume Lettre

  238. Department of Public Health and Clinical Medicine, Unit of Family Medicine, Umeå University, Umeå, 90185, Sweden

    Olov Rolandsson

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

    Danish Saleheen & Wei Zhao

  240. Division of Epidemiology & Community Health University of Minnesota, Minneapolis, 55454, Minnesota, USA

    Pamela J. Schreiner

  241. Duke University, Durham, 27703, North Carolina, USA

    Svati Shah

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

    Xueling Sim

  243. Departement of Haematology, University of Cambridge, Cambridge, CB2 OPT, UK

    Suthesh Sivapalaratnam

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

    Suthesh Sivapalaratnam

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

    Kerrin S. Small & Timothy D. Spector

  246. Alzheimer Scotland Dementia Research Centre, University of Edinburgh, Edinburgh, EH8 9JZ, UK

    John M. Starr

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

    Leen M. ‘t Hart & Natasja M. van Schoor

  248. Department of Molecular Cell Biology, Leiden University Medical Center, Leiden, 2333ZC, The Netherlands

    Leen M. ‘t Hart

  249. Department of Molecular Epidemiology, Leiden University Medical Center, Leiden, 2333ZC, The Netherlands

    Leen M. ‘t Hart

  250. College of Biomedical and Life Sciences, Cardiff University, Cardiff, CF14 4EP, UK

    Katherine E. Tansey

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

    Katherine E. Tansey

  252. Institute for Community Medicine, University Medicine Greifswald, Greifswald, 17475, Germany

    Alexander Teumer

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

    Anke Tönjes

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

    Gerard Tromp

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

    Stella Trompet

  256. Anogia Medical Centre, Anogia, Greece

    Emmanouil Tsafantakis

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

    Jaakko Tuomilehto

  258. Dasman Diabetes Institute, Dasman, 15462, Kuwait

    Jaakko Tuomilehto

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

    Jaakko Tuomilehto

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

    Rudolf Uher

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

    Andries R. Van Der Leij

  262. Department of Obstetrics and Gynecology, Institute for Medicine and Public Health, Vanderbilt Genetics Institute, Vanderbilt University, Nashville, 37203, Tennessee, USA

    Digna R. Velez Edwards

  263. MRL, Merck & Co., Inc., Cardiometabolic Disease, Kenilworth, 07033, New Jersey, USA

    Thomas F. Vogt

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

    Mark Walker

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

    Shuai Wang

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

    Jennifer Wessel

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

    James G. Wilson

  268. Danish Diabetes Academy, Odense, 5000, Denmark

    Daniel R. Witte

  269. Department of Public Health, Aarhus University, Aarhus, 8000, Denmark

    Daniel R. Witte

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

    Michael O. Woods

  271. GlaxoSmithKlein, King of Prussia, Pennsylvania, 19406, USA

    Laura M. Yerges-Armstrong

  272. Department of Clinical Sciences, Quantitative Biomedical Research Center, Center for the Genetics of Host Defense, University of Texas Southwestern Medical Center, Dallas, 75390, Texas, USA

    Xiaowei Zhan

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

    Cristen J. Willer

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

    Dajiang J. Liu

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

    Kari E. North

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

    Nancy L. Heard-Costa

  277. Department of Epidemiology Research, Statens Serum Institut, Copenhagen, 2200, Denmark

    Tune H. Pers

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

    Cecilia M. Lindgren

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

    Ruth J. F. Loos

  280. Departments of Pediatrics and Genetics, Harvard Medical School, Boston, 02115, Massachusetts, USA

    Joel N. Hirschhorn

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

    Panos Deloukas

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