Letter | Published:

Directional dominance on stature and cognition in diverse human populations

Nature volume 523, pages 459462 (23 July 2015) | Download Citation

Abstract

Homozygosity has long been associated with rare, often devastating, Mendelian disorders1, and Darwin was one of the first to recognize that inbreeding reduces evolutionary fitness2. However, the effect of the more distant parental relatedness that is common in modern human populations is less well understood. Genomic data now allow us to investigate the effects of homozygosity on traits of public health importance by observing contiguous homozygous segments (runs of homozygosity), which are inferred to be homozygous along their complete length. Given the low levels of genome-wide homozygosity prevalent in most human populations, information is required on very large numbers of people to provide sufficient power3,4. Here we use runs of homozygosity to study 16 health-related quantitative traits in 354,224 individuals from 102 cohorts, and find statistically significant associations between summed runs of homozygosity and four complex traits: height, forced expiratory lung volume in one second, general cognitive ability and educational attainment (P < 1 × 10−300, 2.1 × 10−6, 2.5 × 10−10 and 1.8 × 10−10, respectively). In each case, increased homozygosity was associated with decreased trait value, equivalent to the offspring of first cousins being 1.2 cm shorter and having 10 months’ less education. Similar effect sizes were found across four continental groups and populations with different degrees of genome-wide homozygosity, providing evidence that homozygosity, rather than confounding, directly contributes to phenotypic variance. Contrary to earlier reports in substantially smaller samples5,6, no evidence was seen of an influence of genome-wide homozygosity on blood pressure and low density lipoprotein cholesterol, or ten other cardio-metabolic traits. Since directional dominance is predicted for traits under directional evolutionary selection7, this study provides evidence that increased stature and cognitive function have been positively selected in human evolution, whereas many important risk factors for late-onset complex diseases may not have been.

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Acknowledgements

This paper is the work of the ROHgen consortium. We thank the participants in all ROHgen studies; cohort-specific acknowledgements are detailed in Supplementary Table 6. This work was funded by a UK Medical Research Council (MRC) PhD studentship to P.K.J.; and J.F.W. and O.P. acknowledge support from the MRC Human Genetics Unit “QTL in Health and Disease” programme. We thank W. G. Hill for discussions and comments on the manuscript and K. Lindsay for administrative assistance.

Author information

Author notes

    • Peter K. Joshi
    • , Tonu Esko
    • , Ozren Polašek
    •  & James F. Wilson

    These authors contributed equally to this work.

Affiliations

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

    • Peter K. Joshi
    • , Igor Rudan
    • , Harry Campbell
    • , Ozren Polašek
    •  & James F. Wilson
  2. Estonian Genome Center, University of Tartu, Riia 23b, 51010, Tartu, Estonia.

    • Tonu Esko
    • , Reedik Mägi
    • , Evelin Mihailov
    • , Lili Milani
    • , Mari Nelis
    • , Natalia Tšernikova
    • , Andrew P. Morris
    • , Andres Metspalu
    •  & Markus Perola
  3. Division of Endocrinology and Center for Basic and Translational Obesity Research, Boston Children's Hospital, Cambridge, 02141 Massachusetts, USA.

    • Tonu Esko
    • , Yingleong Chan
    • , Sailaja Vedantam
    • , Rany M. Salem
    •  & Joel N. Hirschhorn
  4. Program in Medical and Population Genetics, Broad Institute, Cambridge Center 7, Cambridge, Massachusetts 02242, USA.

    • Tonu Esko
    • , Yingleong Chan
    • , Sailaja Vedantam
    • , Rany M. Salem
    • , Cecilia M. Lindgren
    •  & Joel N. Hirschhorn
  5. Department of Genetics, Harvard Medical School, 25 Shattuck St, Boston, Massachusetts 02115, USA.

    • Tonu Esko
    • , Yingleong Chan
    • , Sailaja Vedantam
    • , Rany M. Salem
    •  & Joel N. Hirschhorn
  6. Unit of Public Health Genomics, National Institute for Health and Welfare, P.O. Box 104, Helsinki, FI-00251, Finland.

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    • , Niina Eklund
    • , Anni Joensuu
    • , Antti-Pekka Sarin
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  28. Institute for Translational Genomics and Population Sciences, Los Angeles Biomedical Research Institute, 1124 W. Carson Street, Torrance, California 90502, USA.

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  45. The GeneSTAR Research Program, Division of General Internal Medicine, Department of Medicine, The Johns Hopkins University School of Medicine, Baltimore, Maryland 21287, USA.

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  54. University of Groningen, University Medical Center Groningen, Department of Cardiology, Hanzeplein 1, Groningen, 9700 RB, The Netherlands.

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    • Helen R. Warren
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  60. NIHR Barts Cardiovascular Biomedical Research Unit, Queen Mary University of London, Charterhouse Square, London EC1M 6BQ, UK.

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  61. Department of Medicine, University of Mississippi Medical Center, 2500 N. State St., Jackson, Mississippi 39216, USA.

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  62. Pennington Biomedical Research Center, 6400 Perkins Rd, Baton Rouge, Louisiana 70808, USA.

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  63. MRC Epidemiology Unit, University of Cambridge School of Clinical Medicine, Cambridge Biomedical Campus, Cambridge CB2 0QQ, UK.

    • John R. B. Perry
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  64. Institute for Maternal and Child Health - IRCCS “Burlo Garofolo”, via dell'Istria 65, 34137 Trieste, Italy.

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    • , Jessica van Setten
    •  & Gerard Pasterkamp
  84. Center for Applied Genomics, Children's Hospital of Philadelphia, 3615 Civic Center Boulevard, Philadelphia, Pennsylvania 19104, USA.

    • Jonathan P. Bradfield
    • , Frank D. Mentch
    • , Struan F.A. Grant
    •  & Hakon Hakonarson
  85. Genetics of Complex Traits, University of Exeter Medical School, University of Exeter, Royal Devon and Exeter Hospital, Barrack Road, Exeter EX2 5DW, UK.

    • Andrew R. Wood
    •  & Timothy M. Frayling
  86. COPSAC, Copenhagen Prospective Studies on Asthma in Childhood, Herlev and Gentofte Hospital, University of Copenhagen, Ledreborg Allé 34, DK-2820 Copenhagen, Denmark.

    • Tarunveer S. Ahluwalia
    •  & Hans Bisgaard
  87. Novo Nordisk Centre for Basic Metabolic Research, Section of Metabolic Genetics, Faculty of Health and Medical Sciences, University of Copenhagen, Universitetsparken 1, Copenhagen, 2100, Denmark.

    • Tarunveer S. Ahluwalia
    •  & Thorkild I. A. Sørensen
  88. Department of Cardiovascular Sciences, University of Leicester, BHF Cardiovascular Research Centre, Glenfield Hospital, Groby Road, Leicester LE3 9QP, UK.

    • Leanne M. Hall
    • , Maciej Tomaszewski
    •  & Nilesh J. Samani
  89. Department of Health Sciences, University of Milan, via A. di Rudinì 8, 20142 Milan, Italy.

    • Erika Salvi
    •  & Daniele Cusi
  90. Centre for Ophthalmology and Visual Science, University of Western Australia, Lions Eye Institute, 2 Verdun Street, Perth, Western Australia 6009, Australia.

    • Seyhan Yazar
    •  & David A. Mackey
  91. Department of Epidemiology Research, Statens Serum Institut, Artillerivej 5, Copenhagen, 2300, Denmark.

    • Lisbeth Carstensen
    • , Bjarke Feenstra
    • , Frank Geller
    •  & Mads Melbye
  92. Clinical Epidemiology, Leiden University Medical Center, PO Box 9600, Leiden, 2300 RC, The Netherlands.

    • Hugoline G. de Haan
    •  & Astrid van Hylckama Vlieg
  93. Department of Human Genetics, University of Chicago, 920 E. 58th Street, Chicago, Illinois 60637, USA.

    • Mark Abney
    • , William Wentworth-Shields
    •  & Carole Ober
  94. Department of Family and Preventive Medicine, University of California San Diego, 9500 Gilman Drive, La Jolla, California 92093, USA.

    • Matthew A. Allison
  95. Department of Cardiology, Division Heart and Lungs, University Medical Center Utrecht, Heidelberglaan 100, Utrecht, 3584 CX, The Netherlands.

    • Folkert W. Asselbergs
  96. Durrer Center for Cardiogenetic Research, ICIN-Netherlands Heart Institute, Catharijnesingel 52, Utrecht, 3501 DG, The Netherlands.

    • Folkert W. Asselbergs
    •  & Pim van der Harst
  97. Institute of Cardiovascular Science, faculty of Population Health Sciences, University College London, Gower Street, London WC1E 6BT, UK.

    • Folkert W. Asselbergs
  98. University of Groningen, University Medical Center Groningen, Department of Internal Medicine, Hanzeplein 1, Groningen, 9700 RB, The Netherlands.

    • Stephan J. L. Bakker
  99. Department of Medicine, Columbia University, 622 W. 168th Street, New York, New York 10032, USA.

    • R. Graham Barr
    •  & Walter R. Palmas
  100. Institute for Community Medicine, University Medicine Greifswald, W.-Rathenau-Str. 48, Greifswald 17475, Germany.

    • Sebastian E. Baumeister
    •  & Alexander Teumer
  101. Department of Economics, Cornell University, 480 Uris Hall, Ithaca, New York 14853, USA.

    • Daniel J. Benjamin
  102. Department of Economics and Center for Economic and Social Research, University of Southern California, 314C Dauterive Hall, 635 Downey Way, Los Angeles, California 90089, USA.

    • Daniel J. Benjamin
  103. Human Genetics Center, School of Public Health, University of Texas Health Science Center at Houston, 1200 Pressler Street, Suite 453E, Houston, Texas 77030, USA.

    • Eric Boerwinkle
    •  & Alanna Morrison
  104. Centre for Genomic and Experimental Medicine, University of Edinburgh, Western General Hospital, Edinburgh EH4 2XU, UK.

    • Archie Campbell
    • , Generation Scotland
    • , Sarah E. Harris
    •  & David J. Porteous
  105. McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University School of Medicine, Baltimore, Maryland 21205, USA.

    • Aravinda Chakravarti
    •  & Georg Ehret
  106. Program in Genetic Epidemiology and Statistical Genetics, Harvard School of Public Health, 665 Huntington Ave, Boston, Massachusetts 02115, USA.

    • Constance Chen
    •  & Peter Kraft
  107. Genome Technology Branch, National Human Genome Research Institute, NIH, Bethesda, Maryland 20892, USA.

    • Francis S. Collins
  108. College of Medicine, Dentistry and Nursing, Ninewells Hospital and Medical School, College Office, Level 10, Dundee DD1 9SY, UK.

    • John Connell
  109. National Heart, Lung, and Blood Institute's Framingham Heart Study, 73 Mt. Wayte Ave, Framingham, Massachusetts 01702, USA.

    • L. Adrienne Cupples
    • , Caroline S. Fox
    • , Nancy L. Heard-Costa
    • , George T. O'Connor
    • , Sudha Seshadri
    •  & Josée Dupuis
  110. Psychology, University of Edinburgh, 7 George Square, Edinburgh EH8 9JZ, UK.

    • Gail Davies
    • , Paul Redmond
    •  & Ian J. Deary
  111. Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, 7 George Square, Edinburgh EH8 9JZ, UK.

    • Gail Davies
    • , Sarah E. Harris
    • , David C.M. Liewald
    • , John M. Starr
    • , Ian J. Deary
    •  & David J. Porteous
  112. Department of Internal Medicine B, University Medicine Greifswald, Ferdinand-Sauerbruch-Str. NK, Greifswald 17475, Germany.

    • Marcus Dörr
  113. Cardiology, Geneva University Hospitals, Rue Gabrielle-Perret-Gentil, 4, Genève 14, 1211, Switzerland.

    • Georg Ehret
  114. Robertson Centre, University of Glasgow, Boyd Orr Building, Glasgow G12 8QQ, Scotland.

    • Ian Ford
  115. Division of Endocrinology, Brigham and Women's Hospital and Harvard Medical School, 75 Francis St, Boston, Massachusetts 02115, USA.

    • Caroline S. Fox
  116. Institute of Clinical Chemistry and Laboratory Medicine, University Medicine Greifswald, Ferdinand-Sauerbruch-Str. NK, 17475 Greifswald, Germany.

    • Nele Friedrich
  117. Division of Biostatistics, Washington University, 660 S Euclid, St Louis, Missouri 63110, USA.

    • Charles Gu
  118. Roslin Institute, University of Edinburgh, Easter Bush, Midlothian, Edinburgh EH25 9RG, UK.

    • Chris Haley
  119. National Institutes on Aging, National Institutes of Health, Bethesda, Maryland 20892, USA.

    • Tamara B. Harris
    •  & Lenore J. Launer
  120. Department of Neurology, Boston University School of Medicine, 72 E Concord St, Boston, Massachusetts 02118, USA.

    • Nancy L. Heard-Costa
    •  & Sudha Seshadri
  121. Department of Public Health, University of Helsinki, Hjelt Institute, P.O.Box 41, Mannerheimintie 172, Helsinki, FI-00014, Finland.

    • Kauko Heikkilä
    • , Anu Loukola
    • , Eero Vuoksimaa
    •  & Jaakko Kaprio
  122. Musculoskeletal Research Programme, Division of Applied Medicine, University of Aberdeen, Foresterhill, Aberdeen AB25 2ZD, UK.

    • Lynne J. Hocking
  123. State Key Laboratory of Medical Genomics, Shanghai Institute of Hematology, Rui Jin Hospital Affiliated with Shanghai Jiao Tong University School of Medicine, 197 Rui Jin Er Road, Shanghai, 200025, China.

    • Jinyan Huang
  124. Department of Radiology, Erasmus Medical Center, PO Box 2040, Rotterdam, 3000 CA, The Netherlands.

    • M. Arfan Ikram
  125. Department of Medical Sciences, Molecular Epidemiology and Science for Life Laboratory, Uppsala University, Uppsala, SE-17121, Sweden.

    • Erik Ingelsson
  126. Uppsala Clinical Research Center, Uppsala University, Uppsala, SE-75237, Sweden.

    • Åsa Johansson
  127. Department of Chronic Disease Prevention, National Institute for Health and Welfare, P.O. Box 30, Helsinki, FI-00271, Finland.

    • Pekka Jousilahti
    • , Satu Männistö
    • , Veikko Salomaa
    • , Johan G. Eriksson
    •  & Erkki Vartiainen
  128. Department of Cardiology C5-P, Leiden University Medical Center, PO Box 9600, Leiden, 2300 RC, The Netherlands.

    • J. Wouter Jukema
  129. Department of Clinical Physiology, University of Tampere and Tampere University Hospital, P.O. Box 2000, Tampere, FI-33521, Finland.

    • Mika Kähönen
  130. Diabetes Prevention Unit, National Institute for Health and Welfare, P.O. Box 30, FI-00271 Helsinki, Finland.

    • Heikki A. Koistinen
    •  & Jaakko Tuomilehto
  131. Department of Medicine, Division of Endocrinology, Helsinki University Central Hospital, P.O.Box 340, Haartmaninkatu 4, Helsinki, FI-00029, Finland.

    • Heikki A. Koistinen
    •  & Kirsi Pietilainen
  132. Minerva Foundation Institute for Medical Research, Biomedicum 2U, Tukholmankatu 8, Helsinki, FI-00290, Finland.

    • Heikki A. Koistinen
  133. Laboratory for Genotyping Development RCfIMS, 1-7-22 Suehiro-cho, Tsurumi-ku, Yokohama, Kanagawa, 230-0045, Japan.

    • Michiaki Kubo
  134. Department of Medicine, University of Eastern Finland and Kuopio University Hospital, Kuopio, FI-70210, Finland.

    • Johanna Kuusisto
    •  & Markku Laakso
  135. Institute of Behavioural Sciences, University of Helsinki, P.O. Box 9, University of Helsinki, Helsinki, FI-00014, Finland.

    • Jari Lahti
  136. Folkhälsan Reasearch Centre, PB 63, Helsinki, FI-00014 University of Helsinki, Finland.

    • Jari Lahti
  137. Department of Clinical Chemistry, Fimlab Laboratories and School of Medicine University of Tampere, Tampere, FI-33520, Finland.

    • Terho Lehtimäki
  138. Department of Medical Sciences, University Hospital, Uppsala, 75185, Sweden.

    • Lars Lind
  139. Transplantation laboratory, Haartman Institute, University of Helsinki, P.O. Box 21, Helsinki, FI-00014, Finland.

    • Marja-Liisa Lokki
    •  & Efthymia Vlachopoulou
  140. National Institute of Environmental Health Sciences, National Institutes of Health, Department of Health and Human Services, Research Triangle Park, North Carolina 27709, USA.

    • Stephanie J. London
  141. Ophthalmology, Massachusetts Eye and Ear, 243 Charles Street, Boston, Massachusetts 02114, USA.

    • Stephanie J. Loomis
  142. Department of Statistics, University of Auckland, 303.325 Science Centre, Private Bag 92019, Auckland, 1142, New Zealand.

    • Thomas Lumley
  143. Department of Health, Functional Capacity and Welfare, National Institute for Health and Welfare, P.O. Box 30, Helsinki, FI-00271, Finland.

    • Annamari Lundqvist
    • , Harri Rissanen
    •  & Paul Knekt
  144. Department of Internal Medicine, University Hospital, Rue du Bugnon 44, Lausanne, 1011, Switzerland.

    • Pedro Marques-Vidal
    •  & Peter Vollenweider
  145. Human Genetics, Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.

    • Angela Matchan
    • , Nicole Soranzo
    • , Lorraine Southam
    •  & Eleftheria Zeggini
  146. Division of Allergy and Clinical Immunology, Department of Medicine, The Johns Hopkins University School of Medicine, Baltimore, Maryland 21224, USA.

    • Rasika A. Mathias
  147. Laboratory of Molecular Medicine, Human Genome Center, Institute of Medical Science, The University of Tokyo, 4-6-1 Shirokanedai, Minato-ku, Tokyo, 108-8639, Japan.

    • Koichi Matsuda
  148. Division of General Internal Medicine, Massachusetts General Hospital, 50 Staniford St, Boston, Massachusetts 02114, USA.

    • James B. Meigs
  149. Institute of Human Genetics, Helmholtz Zentrum München, German Research Center for Environmental Health, Ingolstädter Landstr. 1, Neuherberg 85764, Germany.

    • Thomas Meitinger
  150. Institute of Human Genetics, Klinikum rechts der Isar, Technische Universität München, Ismaninger Str. 22, München 81675, Germany.

    • Thomas Meitinger
  151. Molecular Epidemiology, QIMR Berghofer Medical Research Institute, 300 Herston Rd, Herston, Brisbane, Queensland 4006, Australia.

    • Grant W. Montgomery
  152. Genome Science Institute, Boston University School of Medicine, 72 East Concord Street, E-304, Boston, Massachusetts 02118, USA.

    • Richard H. Myers
  153. HUCH Heart and Lung center, Helsinki University Central Hospital, P.O. Box 340, Helsinki, FI-00029, Finland.

    • Markku S. Nieminen
    •  & Juha Sinisalo
  154. Pulmonary Center and Department of Medicine, Boston University School of Medicine, 72 E Concord St, Boston, Massachusetts 02118, USA.

    • George T. O'Connor
  155. Department of Medicine, University of Ibadan, Ibadan, Nigeria.

    • Adesola Ogunniyi
    •  & Babatunde Salako
  156. ICAMS, University of Glasgow, 126 University Way, Glasgow G12 8TA, UK.

    • Sandosh Padmanabhan
  157. Division of Epidemiology and Community Health, University of Minnesota, 1300 S 2nd Street, Minneapolis, Minnesota 55454, USA.

    • James S. Pankow
  158. Centre for Global Health and Department of Public Health, School of Medicine, University of Split, Soltanska 2, 21000 Split, Croatia.

    • Inga Patarcic
    •  & Ozren Polašek
  159. Obesity Research Unit, Research Programs Unit, Diabetes and Obesity, University of Helsinki, P.O.Box 63, Haartmaninkatu 8, FI-00014, Helsinki, Finland.

    • Kirsi Pietilainen
  160. International Centre for Circulatory Health, Imperial College London, London W2 1LA, UK.

    • Neil Poulter
    •  & Peter Sever
  161. Department of Genomics of Common Disease, School of Public Health, Imperial College London, London SW7 2AZ, UK.

    • Inga Prokopenko
    •  & Philippe Froguel
  162. Department of Cardiology and Cardio thoracic Surgery Hero DMC Heart Institute, Civil Lines, 141001, Ludhiana, India.

    • Sarju Ralhan
  163. Department Public Health Sciences, University of Virginia School of Medicine, 3232 West Complex, Charlottesville, Virginia 22908, USA.

    • Stephen S. Rich
  164. Department of Psychological & Brain Sciences, Indiana University Bloomington, 1101 E. 10th Street, Bloomington, Indiana 47405, USA.

    • Richard Rose
  165. Institute of Molecular Biology and Biochemistry, Medical University Graz, Harrachgasse 21, Graz, A-8010, Austria.

    • Helena Schmidt
    •  & Anna Maria Töglhofer
  166. Science for Life Laboratory, Karolinska Institutet, Stockholm, SE-17121, Sweden.

    • Bengt Sennblad
  167. University of Dundee, Kirsty Semple Way, Dundee DD2 4DB, UK.

    • Blair H. Smith
  168. Cardiovascular Health Research Unit, Division of Cardiology, University of Washington, 1730 Minor Ave, Suite 1360, Seattle, Washington 98101, USA.

    • Nona Sotoodehnia
  169. Molecular and Cellular Therapeutics, Royal College of Surgeons in Ireland, St. Stephen's Green, Dublin 2, Ireland.

    • Alice V. Stanton
  170. UMR INSERM U1122; IGE-PCV “Interactions Gène-Environnement en Physiopathologie Cardio-Vasculaire”, INSERM, University of Lorraine, 30 Rue Lionnois, 54000 Nancy, France.

    • Maria G. Stathopoulou
  171. Institute of Medical Informatics, Biometry and Epidemiology, Chair of Genetic Epidemiology, Ludwig-Maximilians-Universität, Munich 81377, Germany.

    • Konstantin Strauch
  172. Department of Endocrinology, All India Institute of Medical Sciences, Ansari Nagar East, New Delhi, 110029, India.

    • Nikhil Tandon
  173. National Heart and Lung Institute, Imperial College London, Du Cane Road, London W12 0NN, UK.

    • Sian-Tsun Tang
    •  & Jaspal S. Kooner
  174. Department of Public Health Sciences, Stritch School of Medicine, Loyola University Chicago, Maywood, Illinois 60153, USA.

    • Bamidele O. Tayo
    •  & Richard S. Cooper
  175. NIHR Leicester Cardiovascular Biomedical Research Unit, University of Leicester, Glenfield Hospital, Groby Road, Leicester LE3 9QP, UK.

    • Maciej Tomaszewski
    •  & Nilesh J. Samani
  176. Institute of Molecular and Cell Biology, University of Tartu, Riia 23, Tartu, 51010, Estonia.

    • Natalia Tšernikova
    •  & Andres Metspalu
  177. Centre for Vascular Prevention, Danube-University Krems, 3500 Krems, Austria.

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

    • Jaakko Tuomilehto
  179. Department of Internal Medicine, Erasmus Medical Center, PO Box 2040, Rotterdam, 3000 CA, The Netherlands.

    • Andre G. Uitterlinden
  180. Department of Epidemiology, Johns Hopkins Bloomberg School of Public Health, Baltimore, Maryland 21205, USA.

    • Dhananjay Vaidya
  181. Finnish Lung Health Association, Sibeliuksenkatu 11 A 1, Helsinki, FI-00250, Finland.

    • Tuula Vasankari
  182. Genetic Epidemiology, QIMR Berghofer Medical Research Institute, 300 Herston Rd, Herston, Brisbane, Queensland 4006, Australia.

    • John B. Whitfield
    •  & Nicholas G. Martin
  183. Diabetes Unit, KEM Hospital and Research Centre, Rasta Peth, Pune, 411011, India.

    • Chittaranjan S. Yajnik
  184. Renal Unit, Department of Medicine, University of Verona, Piazzale A. Stefani 1, 37124 Verona, Italy.

    • Gianluigi Zaza
  185. Department of Epidemiology and Biostatistics, Case Western Reserve University, Cleveland, Ohio 44106, USA.

    • Xiaofeng Zhu
  186. Department of Medicine, Stanford University, 300 Pasteur Drive, Stanford, California 94305, USA.

    • Mads Melbye
  187. Department of Pediatrics, Perelman School of Medicine, The University of Pennsylvania, 3615 Civic Center Boulevard, Philadelphia, Pennsylvania 19104, USA.

    • Struan F.A. Grant
    •  & Hakon Hakonarson
  188. Translational Gerontology Branch, National institute on Aging, Baltimore, Maryland 21225, USA.

    • Luigi Ferrucci
  189. Usher Institute for Population Health Sciences and Informatics, University of Edinburgh, No. 9 Edinburgh Bioquarter, 9 Little France Road, Edinburgh EH16 4UX, UK.

    • Andrew D. Morris
  190. Centre for Pharmacogenetics and Pharmacogenomics, Medical Research Institute, University of Dundee, Ninewells Hospital and School of Medicine, Dundee DD1 9SY, UK.

    • Colin N. A. Palmer
  191. Princess Al-Jawhara Al-Brahim Centre of Excellence in Research of Hereditary Disorders (PACER-HD), King Abdulaziz University, Jeddah, 21589, Saudi Arabia.

    • Panos Deloukas
  192. Faculty of Medicine, Imperial College London, Charing Cross Campus, St Dunstan's Road, London W6 8RP, UK.

    • Lars Bertram
  193. Department of Medicine, University of Leipzig, Leipzig 04103, Germany.

    • Anke Tönjes
  194. Institute of Preventive Medicine, Bispebjerg and Frederiksberg Hospital, The Capital Region, Copenhagen, 2000, Denmark.

    • Thorkild I. A. Sørensen
  195. Department of Epidemiology, University of Alabama at Birmingham, 1665 University Boulevard, Birmingham, Alabama 35294, USA.

    • Donna K. Arnett
  196. Department of Psychiatry, University Medical Center Groningen, University of Groningen, P.O. Box 30.001, Groningen, 9700 RB, The Netherlands.

    • Albertine J. Oldehinkel
  197. Epidemiology of diabetes, obesity and chronic kidney disease over the lifecourse, Inserm, CESP Center for Research in Epidemiology and Population Health U1018, 16 Avenue Paul Vaillant Couturier, 94807 Villejuif, France.

    • Beverley Balkau
  198. Dipartimento di Scienze Mediche, Catholic University of the Sacred Heart, Via G. Moscati 31/34, 00168 Roma, Italy.

    • Giovanni Gambaro
  199. Department of Biostatistics, University of Liverpool, Duncan Building, Daulby Stree, Liverpool L69 3GA, UK.

    • Andrew P. Morris
  200. Department of General Practice and Primary Health Care, University of Helsinki, P.O. Box 20, University of Helsinki, Helsinki, FI-00014, Finland.

    • Johan G. Eriksson
  201. Vasa Central Hospital, Sandviksgatan 2-4, Vasa, FI-65130, Finland.

    • Johan G. Eriksson
  202. Folkhälsan Reasearch Centre, PB 63, University of Helsinki, Helsinki, FI-00014, Finland.

    • Johan G. Eriksson
  203. Unit of General Practice, Helsinki University Central Hospital, Haartmaninkatu 4, Helsinki, FI-00290, Finland.

    • Johan G. Eriksson
  204. Neuro-Imaging Genetics, QIMR Berghofer Medical Research Institute, 300 Herston Rd, Herston, Brisbane, Queensland 4006, Australia.

    • Margie J. Wright
  205. Cardiovascular Genetics Division, University of Utah, 420 Chipeta Way, Room 1160, Salt Lake City, Utah 84117, USA.

    • Steven C. Hunt
  206. Alzheimer Scotland Research Centre, University of Edinburgh, 7 George Square, Edinburgh EH8 9JZ, UK.

    • John M. Starr
  207. Department of Psychiatry, Erasmus Medical Center, PO Box 2040, Rotterdam, 3000 CA, The Netherlands.

    • Henning Tiemeier
  208. National Institute for Health and Welfare (THL), P.O.Box 30, Mannerheimintie 166, Helsinki, FI-00271, Finland.

    • Jaakko Kaprio
  209. Department of Kinesiology, Laval University, 2300 rue de la Terrasse, Quebec G1V 0A6, Canada.

    • Louis Pérusse
  210. Department of Physiology and Biophysics, University of Mississippi Medical Center, 2500 N. State Street, Jackson, Mississippi 39216, USA.

    • James G. Wilson
  211. Department of Clinical Physiology and Nuclear Medicine, University of Turku and Turku University Hospital, Turku, FI-20521, Finland.

    • Olli Raitakari
  212. Research Center of Applied and Preventive Cardiovascular medicine, University of Turku, Turku, FI-20521, Finland.

    • Olli Raitakari
  213. University of Groningen, University Medical Center Groningen, Department of Genetics, Hanzeplein 1, Groningen, 9700 RB, The Netherlands.

    • Pim van der Harst
  214. Department of Economics, Stockholm School of Economics, Box 6501, Stockholm, SE-113 83, Sweden.

    • Magnus Johannesson
  215. Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Box 281, Stockholm, SE-171 77, Sweden.

    • Patrik K. E. Magnusson
  216. Department of Genetics and Biostatistics, Washington University School of Medicine, 4444 Forest Park Boulevard, Saint Louis, Missouri 63108, USA.

    • Ingrid B. Borecki
  217. Department of Health Policy and Management, Johns Hopkins Bloomberg School of Public Health, Baltimore, Maryland 21205, USA.

    • Diane M. Becker
  218. Department of Pediatrics, University of Oklahoma Health Sciences Center, 940 Stanton Young Boulevard, Oklahoma City, Oklahoma 73104, USA.

    • Dharambir K. Sanghera
  219. Department of Pharmaceutical Sciences, University of Oklahoma Health Sciences Center, Oklahoma City, Oklahoma 73104, USA.

    • Dharambir K. Sanghera
  220. Sidra Medical and Research Centre, Doha, Qatar.

    • Paolo Gasparini
  221. The Mindich Child Health and Development Institute, Icahn School of Medicine at Mount Sinai, One Gustave L. Levy Place, New York, New York 10029, USA.

    • Ruth J. F. Loos
  222. Genome Institute of Singapore, 60 Biopolis Street, #02-01 Genome, Singapore, 138672, Singapore.

    • Giriraj R. Chandak
  223. Program for Personalised and Genomic Medicine, Department of Medicine, University of Maryland School of Medicine, 685 Baltimore St. MSTF, Baltimore, Maryland 21201, USA.

    • Alan R. Shuldiner
  224. Geriatric Research and Education Clinical Center, Veterans Administration Medical Center, 685 W Baltimore MSTF, Baltimore, Maryland 21201, USA.

    • Alan R. Shuldiner
  225. BHF Glasgow Cardiovascular Research Centre, University of Glasgow, 126 University Place, Glasgow G12 8TA, UK.

    • Naveed Sattar
  226. Carolina Center for Genome Sciences, University of North Carolina at Chapel Hill, 137 E. Franklin Street, Suite 306, Chapel Hill, North Carolina 27599, USA.

    • Kari E. North
  227. Cardiovascular Health Research Unit, Departments of Medicine, Epidemiology and Health Services, University of Washington, 1730 Minor Ave, Suite 1360, Seattle, Washington 98101, USA.

    • Bruce M. Psaty
  228. Group Health Research Institute, Group Health Cooperative, 1730 Minor Ave, Suite 1360, Seattle, Washington 98101, USA.

    • Bruce M. Psaty
  229. Imperial College Healthcare NHS Trust, Imperial College London, Praed Street, London W2 1NY, UK.

    • John C. Chambers
    •  & Jaspal S. Kooner
  230. Population Health Research Institute, St George's, University of London, Cranmer Terrace, London SW17 0RE, UK.

    • David P. Strachan
  231. Steno Diabetes Centre, Niels Steensens Vej 2, Gentofte, 2820, Denmark.

    • Tarunveer S. Ahluwalia
  232. Translational Laboratory in Genetic Medicine (TLGM), Agency for Science, Technology and Research (A*STAR), 8A Biomedical Grove, 138648, Singapore.

    • Marie Loh
  233. Centre for Population Health Sciences, Usher Institute for Population Health Sciences and Informatics, University of Edinburgh, Teviot Place, Edinburgh EH8 9AG, UK.

    • Sarah Wild
  234. Institutes for Neurogenetics and Integrative & Experimental Genomics, University of Lübeck, Lübeck 23562, Germany.

    • Lars Bertram
  235. School of Biotechnology, Jawaharlal Nehru University, New Delhi 110067, India.

    • Dwaipayan Bharadwaj

Consortia

  1. The BioBank Japan Project

    Laboratory for Statistical Analysis, RIKEN Center for Integrative Medical Sciences, 1-7-22 Suehiro-cho, Tsurumi-ku, Yokohama, Kanagawa, 230-0045, Japan.

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Contributions

C.Hal., P.N., M.Me., H.B., N.J.S., D.C., D.A.M., R.S.C., P.F., G.P., S.F.G., H.H., L.F., R.A.S., A.D.M., C.N.P., G.De., P.D., L.B., U.L., S.I.B., C.M.L., N.J.T., A.Ton., P.B.M., T.I.S., C.N.R., D.K.A., A.J.O., S.L.K., B.B., G.Ga., A.P.M., J.G.E., M.J.W., N.G.M., S.C.H., J.M.S., I.J.D., L.R.G., H.T., N.Pi., J.Ka., N.J.W., L.P., J.G.W., G.Gi., M.J.C., O.R., D.D.B., C.Gi., P.v.d.H., A.A.H., P.Kr., J.S., P.Kn., M.J., P.K.M., A.H., R.Sc., I.B.B., E.Va., D.M.B., D.B., K.L.M., M.B., C.M.v.D., D.K.S., A.Te., E.Z., A.Me., P.G., S.U., C.O., D.T., G.D.S., I.R., D.J.P., M.C., T.D.S., C.Hay., J.D., R.J.L., A.F.W., G.R.C., P.V., A.Sh., P.M.R., J.I.R., N.S., U.G., K.E.N., M.P., B.M.P., D.R.W., M.La., V.G., A.Ta., J.C.C., J.S.K., D.P.S., H.C., J.N.H., M.P., O.P. and J.F.W. designed individual studies. T.N., J.D.F., S.E., V.V., S.Tr., D.I.C., S.S.N., M.Ma., D.R., A.F., L.R.Y., E.H., C.Bo., J.R.P., S.C., U.B., G.M., T.Li., I.D., J.Z., J.P.B., E.S., S.Y., M.A.A., S.J.B., G.R.B., E.P.B., A.Ca., Y. Chan, S.J.C., Y.D.I.C., F.S.C., J.C., A.Co., L.Cu., G.Da., M.D., S.B.E., B.F., M.F.F., I.F., C.S.F., T.M.F., N.Fri., F.Ge., I.Gi., O.G., F.Gr., C.Gu., C.J.H., S.E.H., N.D.H., N.L.H., K.H., L.J.H., G.Ho., P.G.H., E.I., Å.J., P.J., J.J., M.Ka., S.K., S.M.K., N.M.K., H.K.K., M.Ku., J.Ku., J.L., R.A.L., T.Le., D.C.L., L.Li., M.L.L., A.Lo., T.Lu., A.Lu., S.M., K.M., J.B.M., C.Mei., T.M., C.Men., F.D.M., L.M., G.W.M., R.H.M., R.N., M.N., M.S.N., G.T.O., A.O., S.P., W.R.P., J.S.P., I.Pa., K.P., N.Po., S.Ra., P.R., S.S.R., H.R., A.R., L.M.R., R.R., B.Sa., R.M.S., V.S., A.Sa., L.J.S., S.Se., P.S., B.H.S., N.Sor., A.V.St., M.G.S., K.S., N.Ta., K.D.T., B.O.T., A.Tog., M.To., J.T., A.G.U., A.v.H.V., T.V., S.V., E.Vl., E.Vu., M.W., J.B.W., S.W., G.W., C.S.Y., G.Z., X.Z., M.Me., H.B., N.J.S., D.C., D.A.M., R.S.C., G.P., S.F.G., H.H., L.F., R.A.S., G.De., P.D., L.B., U.L., S.I.B., G.D.S., N.J.T., A.Ton., P.B.M., T.I.S., C.N.R., D.K.A., A.J.O., S.L.K., B.B., M.K.K., G.Ga., J.G.E., M.J.W., N.G.M., S.C.H., J.M.S., I.J.D., L.R.G., J.Ka., N.J.W., L.P., J.G.W., G.Gi., M.J.C., O.R., D.D.B., C.Gi., P.v.d.H., A.A.H., P.Kr., J.S., P.Kn., M.J., P.K.M., A.H., R.Sc., I.B.B., E.Va., D.M.B., D.B., K.L.M., M.B., C.M.v.D., D.K.S., E.Z., A.Me., P.G., C.O., D.T., D.J.P., M.C., T.D.S., C.Hay., R.J.L., A.F.W., G.R.C., P.V., A.Sh., P.M.R., J.I.R., N.S., U.G., M.P., B.M.P., D.R.W., M.La., J.C.C., J.S.K., D.P.S., J.N.H., M.P., O.P. and J.F.W. collected the data. S.Tr., D.I.C., M.C.C., C.Bo., U.B., I.D., M.A., F.W.A., S.J.B., D.J.B., E.B., E.P.B., A.Cc., S.J.C., J.C., I.F., T.M.F., C.Gu., C.J.H., T.B.H., N.D.H., M.I., E.I., J.J., P.Ko., M.Ku., L.J.L., R.A.L., L.Li., R.A.M., K.M., J.B.M., G.W.M., R.H.M., P.A.P., K.P., S.S.R., R.R., H.S., P.S., B.H.S., N.Sor., N.Sot., D.Va., J.B.W., C.S.Y., M.Me., N.J.S., D.C., D.A.M., R.S.C., P.F., G.P., S.F.G., H.H., L.F., G.De., P.D., L.B., U.L., S.I.B., C.M.L., A.Ton., P.B.M.,.C.N.R., D.K.A., A.J.O., S.L.K., B.B., G.Ga., A.P.M., M.J.W., N.G.M., S.C.H., J.M.S., I.J.D., L.R.G., J.Ka., N.J.W., L.P., M.J.C., D.D.B., P.v.d.H., P.Kr., M.J., P.K.M., A.H., R.Sc., I.B.B., D.M.B., D.B., K.L.M., M.B., C.M.v.D., D.K.S., E.Z., A.Me., P.G., S.U., C.O., I.R., D.J.P., M.C., T.D.S., C.Hay., A.F.W., G.R.C., P.V., A.Sh., P.M.R., J.I.R., N.S., U.G., K.E.N., B.M.P., D.R.W., M.La., V.G., D.P.S., H.C., O.P. and J.F.W. contributed to funding. P.K.J., T.E., H.Ma., N.E., I.Ga., T.N., A.U.J., C.Sc., A.V.Sm., W.Zhan., Y.O., A.Stc., J.D.F., W. Zhao, T.M.B., M.P.C., N.Fra., S.E., V.V., S.Tr., X.G., D.I.C., J.R.O., T.C., S.S.N., Y. Chen, M.Ma., D.R., M.Ta., A.F., T.Kac., A.Bj., A.v.d.S., Y.W., A.K.G., L.R.Y., L.W., E.H., C.A.R., O.M., M.C.C., C.P., N.V., C.Ba., A.A.A., H.R.W., D.Vu., H.Me., J.R.P., S.S.Mi., M.C.B., S.S.Me., P.A.L., G.M., A.D., L.Y., L.F.B., D.Z., P.J.v.d.M., D.S., R.M., G.He., T.Kar., Z.W., T.Li., I.D., J.Z., W.M., L.La., S.W.v.L., J.P.B., A.R.W., A.Bo., T.S.A., L.M.H., E.S., S.Y., I.M.M., L.Ca., H.G.d.H., M.A., U.A., N.A., F.W.A., S.E.B., S.B., A.Ca., Y. Chan, C.C., G.Da., G.E., B.F., M.F.F., F.Ge., M.G., S.E.H., J.J.H., J.H., J.E.H., P.G.H., A.J., Y.K., S.K., R.A.L., B.L., M.Lo., S.J.Loo., Y.L., P.M., A.Ma., C.Men., F.D.M., E.M., M.E.M., A.Mo., A.O., I.Pa., F.P., I.Pr., L.M.R., B.Sa., R.M.S., R.Sa., H.S., W.R.S., C.Sa., C.Ma., B.Se., S.Sh., S.J.Lon., J.A.S., L.S., R.J.S., M.J.S., S.Ta., B.O.T., A.Tog., M.To., N.Ts., J.v.S., S.V., D.Vo., E.B.W., W.W., J.Y., G.Z., N.J.S., R.A.S., A.D.M., C.N.P., S.I.B., N.J.T., A.P.M., S.C.H., H.T., N.Pi., L.P., P.v.d.H., P.Kr., R.Sc., I.B.B., A.Te., C.O., M.C., J.D., J.I.R., N.S., K.E.N., A.Ta., J.C.C., J.S.K. and D.P.S. analysed the data. P.K.J., T.E., H.Ma., N.E., I.Ga., T.N., A.U.J., C.Sc., A.V.Sm., M.C.B. and D.P.S. performed beta-testing of scripts. P.K.J. and T.E. performed the meta-analysis. P.K.J., T.E., O.P. and J.F.W. wrote the manuscript. All authors approved the final manuscript.

Competing interests

G.P. is a co-founder of CAVADIS B.V. S.W.v.L. is a former employee of CAVADIS B.V. B.M.P. serves on the Data and Safety Monitoring Board of a clinical trial funded by the LifeVest manufacturer (Zoll Lifecor) and on the Yale Open Data Access Project funded by Johnson & Johnson. N.Po. has received financial support and consultancy fees from several pharmaceutical companies that manufacture either blood-pressure-lowering or lipid-lowering agents or both. P.S. has received research awards from Pfizer. No other authors declared a conflict of interest.

Corresponding author

Correspondence to James F. Wilson.

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https://doi.org/10.1038/nature14618

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