Genetic evidence of assortative mating in humans

ABSTRACT In human populations, assortative mating is almost univer­sally positive, with similarities between partners for quantit­ative phenotypes1–6, common disease risk1,3,7–10,

beha­vi­our6,11, social factors12–14 and personality4,5,11. The causes and genetic consequences of assortative mating remain un­re­solved because partner similarity can arise from different

mechanisms: phenotypic assortment based on mate choice15,16, partner interaction and convergence in phenotype over time14,17, or social homogamy where individuals pair according to social or

environmental background. Here, we present theory and an analytical approach to test for genetic evidence of assortative mating and find a correlation in genetic value among partners for a

range of phenotypes. Across three independent samples of 24,662 spousal pairs in total, we infer a correlation at trait-associated loci between partners for height (0.200, 0.004 standard

error, SE) that matched the phenotypic correlation (0.201, 0.004 SE), and a correlation at trait-associated loci for BMI (0.143, 0.007 SE) that was significantly lower than the phenotypic

value (0.228, 0.004 SE). We extend our analysis to the UK Biobank study (7,780 pairs), finding evidence of a correlation at trait-associated loci for waist-to-hip ratio (0.101, 0.041 SE),

systolic blood pressure (0.138, 0.064 SE) and educational attainment (0.654, 0.014 SE). Our results imply that mate choice, combined with widespread pleiotropy among traits, affects the

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Contact customer support SIMILAR CONTENT BEING VIEWED BY OTHERS GENETIC FOOTPRINTS OF ASSORTATIVE MATING IN THE JAPANESE POPULATION Article Open access 22 September 2022 ASSORTATIVE MATING

BIASES MARKER-BASED HERITABILITY ESTIMATORS Article Open access 03 February 2022 GENETIC SIMILARITY BETWEEN RELATIVES PROVIDES EVIDENCE ON THE PRESENCE AND HISTORY OF ASSORTATIVE MATING

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ACKNOWLEDGEMENTS The University of Queensland group is supported by the Australian National Health and Medical Research Council (NHMRC grants 1078037, 1048853 and 1050218), the Australian

Research Council (Discovery Project 160103860) and the National Institute of Health (NIH grants R21ESO25052-01, R01AG042568 and PO1GMO99568). J.Y. is supported by a Charles and Sylvia

Viertel Senior Medical Research Fellowship. We thank the participants of the cohort studies, as well as C. Haley and our colleagues at the Program in Complex Trait Genomics, for comments and

suggestions. 23andMe cohort thank the research participants and employees of 23andMe. This work was supported by the National Human Genome Research Institute of the NIH (grant number

R44HG006981). The UK Biobank research was conducted using the UK Biobank Resource under project 12514. TWINGENE was supported by the Swedish Research Council (M-2005-1112), GenomEUtwin

(EU/QLRT-2001-01254; QLG2-CT-2002-01254), NIH DK U01-066134, the Swedish Foundation for Strategic Research, and the Heart and Lung Foundation grant no. 20070481. The Atherosclerosis Risk in

Communities Study (ARIC) is a collaborative study supported by National Heart, Lung, and Blood Institute contracts (HHSN268201100005C, HHSN268201100006C, HHSN268201100007C,

HHSN268201100008C, HHSN268201100009C, HHSN268201100010C, HHSN268201100011C, and HHSN268201100012C). K.E.N and M.G. are supported by NIDDK R01 DK089256. We thank the staff and participants of

the ARIC study for their contributions. Generation and management of GWAS genotype data for the LifeLines Cohort Study is supported by the Netherlands Organization of Scientific Research

(grant 175.010.2007.006), the Economic Structure Enhancing Fund of the Dutch government, the Ministry of Economic Affairs, the Ministry of Education, Culture and Science, the Ministry for

Health, Welfare and Sports, the Northern Netherlands Collaboration of Provinces, the Province of Groningen, University Medical Center Groningen, the University of Groningen, Dutch Kidney

Foundation and Dutch Diabetes Research Foundation. The authors acknowledge the services of the LifeLines Cohort Study, the contributing research centres delivering data to LifeLines, and all

the study participants. Minnesota Center for Twin and Family Research (MCTFR) is funded by US Public Health Service grants from the National Institute on Alcohol Abuse and Alcoholism

(AA09367, AA11886), National Institute on Drug Abuse (DA05147, DA13240, DA024417, DA036216) and National Institute of Mental Health (MH066140). The funders had no role in study design, data

collection and analysis, decision to publish, or preparation of the manuscript. AUTHOR INFORMATION AUTHORS AND AFFILIATIONS * Institute of Molecular Bioscience, The University of Queensland,

Brisbane,, 4072, Queensland, Australia Matthew R. Robinson, Anna A. E. Vinkhuyzen, Jian Yang & Peter M. Visscher * Department of Research, 23andMe Inc., Mountain View, California,

94041, USA Aaron Kleinman * Department of Epidemiology, Gillings School of Global Public Health, University of North Carolina, Chapel Hill, 27514, North Carolina, USA Mariaelisa Graff & 

Kari E. North * Department of Biostatistics, Gillings School of Global Public Health, University of North Carolina, Chapel Hill, 27514, North Carolina, USA David Couper * Department of

Psychology, University of Minnesota, Minneapolis, Minnesota, USA Michael B. Miller, William G. Iacono & Matt McGue * Department of Psychiatry, VU University Medical Centre & GGZ

inGeest, Amsterdam, The Netherlands Wouter J. Peyrot * Department of Biological Psychology, VU University Amsterdam, Amsterdam, The Netherlands Abdel Abdellaoui * School of Psychology, The

University of Queensland, Brisbane, , 4072, Queensland, Australia Brendan P. Zietsch * Department of Epidemiology, University of Groningen, University Medical Center Groningen, Groningen,

9700 RB, The Netherlands Ilja M. Nolte, Jana V. van Vliet-Ostaptchouk & Harold Snieder * Department of Endocrinology, University of Groningen, University Medical Center Groningen,

Groningen, 9700 RB, The Netherlands Jana V. van Vliet-Ostaptchouk * QIMR Berghofer Medical Research Institute, 300 Herston Road, Herston, 4006, Queensland, Australia Sarah E. Medland & 

Nicholas G. Martin * Karolinska Institutet, Stockholm, SE-171 77, Sweden Patrik K. E. Magnusson * Carolina Centre for Genome Sciences, Gillings School of Global Public Health, University of

North Carolina, Chapel Hill, 27514, North Carolina, USA Kari E. North * The Queensland Brain Institute, The University of Queensland, Brisbane, 4072, Queensland, Australia Jian Yang & 

Peter M. Visscher * Department of Epidemiology, University of Groningen, University Medical Center Groningen, The Netherlands Harold Snieder, Behrooz Z. Alizadeh & H. Marike Boezen *

Department of Genetics, University of Groningen, University Medical Center Groningen, The Netherlands Lude Franke, Morris Swertz & Cisca Wijmenga * Department of Cardiology, University

of Groningen, University Medical Center Groningen, The Netherlands Pim van der Harst * Department of Internal Medicine, Division of Nephrology, University of Groningen, University Medical

Center Groningen, The Netherlands Gerjan Navis * Department of Medical Biology, University of Groningen, University Medical Center Groningen, The Netherlands Marianne Rots * Department of

Endocrinology, University of Groningen, University Medical Center Groningen, The Netherlands Bruce H. R. Wolffenbuttel * Department of Biostatistics, Center for Statistical Genetics,

University of Michigan, Ann Arbor, 48109, Michigan, USA Goncalo R. Abecasis, Michael Boehnke, Jennifer L. Bragg-Gresham, Anne U. Jackson, Adam E. Locke, Laura J. Scott, Nicole Soranzo, 

Heather M. Stringham, Tanya M. Teslovich, Ryan Welch, Robert J. Weyant & Cristen J. Willer * Hudson Alpha Institute for Biotechnology, Huntsville, 35806, Alabama, USA Devin Absher & 

Lindsay L. Waite * Estonian Genome Center, University of Tartu, Tartu, 50410, Estonia Helene Alavere, Tõnu Esko, Krista Fischer, Reedik Mägi, Andres Metspalu, Evelin Mihailov, Lili Milani, 

Mari Nelis, Markus Perola, Mari-Liis Tammesoo & Maris Teder-Laving * Institute of Genetic Epidemiology, Helmholtz Zentrum Mìnchen - German Research Center for Environmental Health,

Neuherberg, 85764, Germany Eva Albrecht, Christian Gieger & Martina Mìller-Nurasyid * Genetics of Complex Traits, Peninsula College of Medicine and Dentistry, University of Exeter,

Exeter, EX1 2LU, UK Hana Lango Allen, Timothy Frayling, Andrew T. Hattersley, John R. B. Perry, Michael N. Weedon & Andrew R. Wood * Department of Clinical Sciences, Lund University

Diabetes Centre, Lund University, Malmö, 20502, Sweden Peter Almgren & Leif C. Groop * Department of Epidemiology, Erasmus MC, Rotterdam, 3015GE, The Netherlands Najaf Amin, Cornelia M.

van Duijn, Karol Estrada, Albert Hofman, Carolina Medina-Gomez, Joyce B. J. van Meurs, Marjolein J. Peters, Fernando Rivadeneira, André G. Uitterlinden, Sophie van Wingerden & Jacqueline

C. M. Witteman * Institut Pasteur de Lille, INSERM U744, Université Lille Nord de France, Lille, F-59000, France Philippe Amouyel * Telethon Institute for Child Health Research, West Perth

Western Australia, 6872, Australia Denise Anderson & Aarno Palotie * Centre for Child Health Research, The University of Western Australia, Australia Denise Anderson * Departments of

Biostatistics, University of Washington, Seattle, 98195, Washington, USA Alice M. Arnold & Barbara McKnight * Collaborative Health Studies Coordinating Center, Seattle, 98115,

Washington, USA Alice M. Arnold * Department of Epidemiology and Public Health, Faculty of Medicine, Strasbourg, France Dominique Arveiler & Kaarel Krjutškov * Icelandic Heart

Association, Kopavogur, Iceland Thor Aspelund, Gudny Eiriksdottir, Vilmundur Gudnason & Albert Vernon Smith * University of Iceland, Reykjavik, Iceland Thor Aspelund, Vilmundur Gudnason 

& Albert Vernon Smith * Department of Cardiology, Division Heart & Lungs, University Medical Center Utrecht, The Netherlands Folkert W. Asselbergs * Department of Medicine, Stanford

University School of Medicine, Stanford, 94305, California, USA Themistocles L. Assimes, Joshua W. Knowles & Thomas Quertermous * Institute of Biomedicine/Physiology, University of

Eastern Finland, Kuopio Campus, Finland Mustafa Atalay & Timo A. Lakka * Wellcome Trust Sanger Institute, Hinxton, Cambridge, CB10 1SA, UK Antony P. Attwood, Inês Barroso, Panos

Deloukas, Sarah Edkins, Sarah E. Hunt, Stavroula Kanoni, Willem H. Ouwehand, Leena Peltonen, Aparna Radhakrishnan, Joshua C. Randall, Samuli Ripatti, So-Youn Shin, Kathleen Stirrups & 

Eleanor Wheeler * Department of Haematology, University of Cambridge, Cambridge, CB2 0PT, UK Antony P. Attwood, Jennifer Jolley, Willem H. Ouwehand, Aparna Radhakrishnan, Augusto Rendon, 

Jennifer G. Sambrook & Jonathan C. Stephens * NIHR Cambridge Biomedical Research Centre, Cambridge, UK Antony P. Attwood, Willem H. Ouwehand & Augusto Rendon * Department of

Neurology, Boston University School of Medicine, Boston, 02118, Massachusetts, USA Larry D. Atwood & Nancy L. Heard-Costa * Department of Internal Medicine, University Medical Center

Groningen, University of Groningen, Groningen, The Netherlands Stephan J. L. Bakker & Gerjan Navis * INSERM CESP Centre for Research in Epidemiology and Public Health U1018, Epidemiology

of diabetes, obesity and chronic kidney disease over the lifecourse, Villejuif, 94807, France Beverley Balkau * University Paris Sud 11, UMRS 1018, Villejuif, 94807, France Beverley Balkau

* Multidisciplinary Cardiovascular Research Centre (MCRC), Leeds Institute of Genetics, Health and Therapeutics (LIGHT), University of Leeds, Leeds, LS2 9JT, UK Anthony J. Balmforth *

Department of Medicine, University of Milan, Surgery and Dentistry, Milano, 20139, Italy Cristina Barlassina * University of Cambridge Metabolic Research Labs, Institute of Metabolic Science

Addenbrooke’s Hospital, Cambridge, CB2 OQQ, UK Inês Barroso * Department of Vascular Medicine, Academic Medical Center, Amsterdam, The Netherlands Hanneke Basart, Kees G. Hovingh & 

Mieke D. Trip * Regensburg University Medical Center, Innere Medizin I, Regensburg, 93053, Germany Sabrina Bauer, Christa Buechler & Kristina Eisinger * Department of Medical Genetics,

University of Lausanne, Lausanne, 1005, Switzerland Jacques S. Beckmann, Sven Bergmann, Toby Johnson, Karen Kapur, Zoltán Kutalik, Diana Marek & Carlo Rivolta * Service of Medical

Genetics, Centre Hospitalier Universitaire Vaudois (CHUV) University Hospital, Lausanne, 1011, Switzerland Jacques S. Beckmann * Department of Molecular Genetics, PathWest Laboratory of

Western Australia, J Block, QEII Medical Centre, Nedlands, 6009, Western Australia, Australia John P. Beilby & Jennie Hui * Busselton Population Medical Research Foundation Inc., Sir

Charles Gairdner Hospital, Nedlands, 6009, Western Australia, Australia John P. Beilby, Jennie Hui, Joseph Hung, Alan L. James, Arthur W. Musk & Lyle J. Palmer * School of Pathology and

Laboratory Medicine, University of Western Australia, Nedlands, 6009, Western Australia, Australia John P. Beilby & Jennie Hui * Department of Surgery and Pathology, University of

Western Australia, Nedlands, 6009, Australia John P. Beilby * Oxford Centre for Diabetes, Endocrinology and Metabolism, University of Oxford, Oxford, OX3 7LJ, UK Amanda J. Bennett, 

Christopher J. Groves, Neelam Hassanali, Fredrik Karpe, Cecilia M. Lindgren, Mark I. McCarthy, Matt J. Neville, Inga Prokopenko, Nigel W. Rayner & Neil R. Robertson * Department of

Social Medicine, University of Bristol, Bristol, BS8 2PS, UK Yoav Ben-Shlomo * Department of Physiology and Biophysics, Keck School of Medicine, University of Southern California, Los

Angeles, 90033, California, USA Richard N. Bergman, Thomas A. Buchanan & Richard M. Watanabe * Swiss Institute of Bioinformatics, Lausanne, 1015, Switzerland Sven Bergmann, Toby Johnson,

 Karen Kapur, Zoltán Kutalik & Diana Marek * Division of Cancer Epidemiology and Genetics, Department of Health and Human Services, National Cancer Institute, National Institutes of

Health, Bethesda, 20892, Maryland, USA Sonja I. Berndt, Stephen J. Chanock, Kevin B. Jacobs, Jianxin Shi & Zhaoming Wang * Zentrum fìr Zahn-, Mund- und Kieferheilkunde, Greifswald,

17489, Germany Reiner Biffar * Molecular Biology Department, Istituto Auxologico Italiano, Milano, Italy Anna Maria Di Blasio & Davide Gentilini * Division of Endocrinology and Diabetes,

Department of Medicine, University Hospital, Ulm, Germany Bernhard O. Boehm * Department of Epidemiology, German Institute of Human Nutrition Potsdam-Rehbruecke, Nuthetal, 14558, Germany

Heiner Boeing & Eva Fisher * Human Genetics Center and Institute of Molecular Medicine, University of Texas Health Science Center, Houston, 77030, Texas, USA Eric Boerwinkle * Centre for

Population Health Sciences, University of Edinburgh, Teviot Place, Edinburgh, EH8 9AG, Scotland Jennifer L. Bolton, Harry Campbell, Gerry Fowkes, Ross M. Fraser, Jackie F. Price, Igor

Rudan, Sarah H. Wild & James F. Wilson * CNRS UMR8199-IBL-Institut Pasteur de Lille, Lille, F-59000, France Amélie Bonnefond, Philippe Froguel, Cecile Lecoeur, David Meyre & Boris

Skrobek * National Human Genome Research Institute, National Institutes of Health, Bethesda, Maryland, 20892, USA Lori L. Bonnycastle, Peter S. Chines, Francis S. Collins, Michael R. Erdos, 

Mario A. Morken, Narisu Narisu & Amy J. Swift * Genome Technology Branch, National Human Genome Research Institute, NIH, Bethesda, 20892, MD, USA Lori L. Bonnycastle, Peter S. Chines, 

Francis S. Collins, Narisu Narisu & Amy J. Swift * Department of Biological Psychology, VU University Amsterdam, Amsterdam, 1081, BT, The Netherlands Dorret I. Boomsma, Eco J. C. Geus, 

Jouke-Jan Hottenga & Gonneke Willemsen * Department of Genetics, Washington University School of Medicine, St Louis, 63110, Missouri, USA Ingrid B. Borecki, Mary F. Feitosa, Shamika

Ketkar, Aldi T. Kraja, Lyle J. Palmer & Michael A. Province * Division of Biostatistics, Washington University School of Medicine, St Louis, 63110, Missouri, USA Ingrid B. Borecki *

Department of Medicine III, University of Dresden, Dresden, 01307, Germany Stefan R. Bornstein & Barbara Ludwig * Department of Medicine III, University of Dresden, Medical Faculty Carl

Gustav Carus, Fetscherstrasse 74, Dresden, 01307, Germany Stefan R. Bornstein & Peter E. H. Schwarz * CNRS UMR8199-IBL-Institut Pasteur de Lille, Lille, F-59019, France Nabila

Bouatia-Naji, Christine Cavalcanti-Proença, Philippe Froguel, Stefan Gaget, Cecile Lecoeur & Vincent Vatin * University Lille Nord de France, Lille, 59000, France Nabila Bouatia-Naji, 

Christine Cavalcanti-Proença, Philippe Froguel, Stefan Gaget, Cecile Lecoeur, David Meyre, Boris Skrobek & Vincent Vatin * Montreal Heart Institute, Montreal, H1T 1C8, Quebec, Canada

Gabrielle Boucher & Guillaume Lettre * Dipartimento di Medicina Sperimentale. Università degli Studi Milano-Bicocca, Monza, Italy Paolo Brambilla * LifeLines Cohort Study, University

Medical Center Groningen, University of Groningen, The Netherlands Harold Snieder, Marcel Bruinenberg, Lude Franke, Melanie M. Van der Klauw, Ronald P. Stolk, Jana V. Van Vliet-Ostaptchouk 

& Bruce H. R. Wolffenbuttel * Division of Endocrinology, Keck School of Medicine, University of Southern California, Los Angeles, 90033, California, USA Thomas A. Buchanan * Genetic

Epidemiology and Biostatistics Platform, Ontario Institute for Cancer Research, Toronto, M5G 1L7, Canada Gemma Cadby * Prosserman Centre for Health Research, Samuel Lunenfeld Research

Institute, Toronto, M5G 1X5, Canada Gemma Cadby & Lyle J. Palmer * Clinical Pharmacology and Barts and The London Genome Centre, William Harvey Research Institute, Barts and The London

School of Medicine and Dentistry, Queen Mary University of London, Charterhouse Square, London, EC1M 6BQ, UK Mark J. Caulfield, Toby Johnson & Patricia B. Munroe * Department of Clinical

Medicine, University of Milano-Bicocca, Monza, Italy Giancarlo Cesana * Harvard Medical School, Boston, 02115, Massachusetts, USA Daniel I. Chasman, Lee M. Kaplan & Paul M. Ridker *

Division of Preventive Medicine, Brigham and Women’s Hospital, Boston, 02215, Massachusetts, USA Daniel I. Chasman, Paul M. Ridker & Lynda M. Rose * Department of OB/GYN and Medical

Genetics Institute, Cedars-Sinai Medical Center, Los Angeles, CA, USA Yii-Der Ida Chen * Department of Medicine, David Geffen School of Medicine at University of California, Los Angeles,

California, USA Yii-Der Ida Chen * University of Texas Southwestern Medical Center, 5323 Harry Hines Blvd, Dallas, 75390-8854, Texas, USA Deborah J Clegg & Thomas Person * Department of

Epidemiology and Biostatistics, School of Public Health, Faculty of Medicine, Imperial College London, London, W2 1PG, UK Lachlan Coin, Paul Elliott, Marjo-Riitta Jarvelin & Ulla Sovio *

British Heart Foundation Glasgow Cardiovascular Research Centre, University of Glasgow, Glasgow, G12 8TA, UK John M. Connell * University of Dundee, Ninewells Hospital & Medical School,

Dundee, DD1 9SY, UK John M. Connell * National Heart and Lung Institute, Imperial College London, London, SW3 6LY, UK William Cookson & Miriam F. Moffatt * Centre for Genetic

Epidemiology and Biostatistics, University of Western Australia, Crawley, 6009, Western Australia, Australia Matthew N. Cooper, Jennie Hui, Robert W. Lawrence & Lyle J. Palmer *

Department of Genetics, University of North Carolina, Chapel Hill, 27599, North Carolina, USA Damien C. Croteau-Chonka, Jennifer R. Kulzer & Karen L. Mohlke * Department of

Biostatistics, Boston University School of Public Health, Boston, 02118, Massachusetts, USA L. Adrienne Cupples, Julius S Ngwa & Charles C. White * Department of Health Sciences,

University of Milan, Ospedale San Paolo, Milano, 20139, Italy Daniele Cusi & Francesca Frau * Fondazione Filarete, Milano, Italy Daniele Cusi * MRC Epidemiology Unit, Institute of

Metabolic Science, Addenbrooke’s Hospital, CB2 0QQ, Cambridge, UK Felix R. Day, Tuomas O. Kilpeläinen, Claudia Langenberg, Shengxu Li, Ruth J. F. Loos, Jian’an Luan, Ken K. Ong, Nicholas J.

Wareham & Jing Hua Zhao * Department of Social Medicine, MRC Centre for Causal Analyses in Translational Epidemiology, Oakfield House, Bristol, BS8 2BN, UK Ian N. M. Day, Debbie A.

Lawlor, George Davey Smith & Nicholas John Timpson * Department of Dietetics-Nutrition, Harokopio University, 70 El. Venizelou Str, Athens, Greece George V. Dedoussis, Maria Dimitriou 

& Eirini V. Theodoraki * Istituto di Neurogenetica e Neurofarmacologia del CNR, Monserrato, 09042, Cagliari, Italy Mariano Dei, Andrea Maschio, Serena Sanna, Manuela Uda & Gianluca

Usala * Istituto di Ricerca Genetica e Biomedicadel CNR, Monserrato, 09042, Cagliari, Italy Mariano Dei & Serena Sanna * Department of Genetic Medicine and Development, University of

Geneva Medical School, Geneva, 1211, Switzerland Emmanouil T. Dermitzakis & Antigone S. Dimas * Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, OX3 7BN, UK

Antigone S. Dimas, Teresa Ferreira, Jon P. Krohn, Cecilia M. Lindgren, Reedik Mägi, Iain Mathieson, Mark I. McCarthy, Josine L. Min, Andrew P. Morris, John F. Peden, Inga Prokopenko, Joshua

C. Randall, Nigel W. Rayner & Neil R. Robertson * Biomedical Sciences Research Center Al, 16672, Fleming, Vari, Greece Antigone S. Dimas * Department of Pharmacy and Pharmacology,

University of Bath, Bath, BA1 1RL, UK Anna L. Dixon * Department of Internal Medicine B, Ernst-Moritz-Arndt University, Greifswald, 17475, Germany Marcus Dörr * Netherlands Genomics

Initiative (NGI)-sponsored Netherlands Consortium for Healthy Aging (NCHA), The Netherlands Cornelia M. van Duijn, Karol Estrada, Albert Hofman, Carolina Medina-Gomez, Joyce B. J. van Meurs,

 Ben A. Oostra, Marjolein J. Peters, Fernando Rivadeneira, André G. Uitterlinden, Jacqueline C. M. Witteman & M. Carola Zillikens * Center of Medical Systems Biology, Leiden University

Medical Center, Leiden, 2333, ZC, The Netherlands Cornelia M. van Duijn & Gert-Jan van Ommen * The London School of Hygiene and Tropical Medicine, London, WC1E 7HT, UK Shah Ebrahim *

South Asia Network for Chronic Disease, Shah Ebrahim * Department of Chronic Disease Prevention, National Institute for Health and Welfare, Unit of Public Health Genomics, Helsinki, 00014,

Finland Niina Eklund, Johannes Kettunen, Kati Kristiansson, Niina Pellikka, Markus Perola, Samuli Ripatti, Kaisa Silander & Ida Surakka * Institute of Molecular and Cell Biology,

University of Tartu, Tartu, 51010, Estonia Niina Eklund, Tõnu Esko, Andres Metspalu, Evelin Mihailov, Mari Nelis & Maris Teder-Laving * MRC-HPA Centre for Environment and Health, London,

W2 1PG, UK Paul Elliott * Clinic of Cardiology, West German Heart Centre, University Hospital of Essen, University Duisburg-Essen, Germany Raimund Erbel * Nordic Center of Cardiovascular

Research (NCCR), Lìbeck, 23538, Germany Jeanette Erdmann & Heribert Schunkert * Universität zu Lìbeck, Medizinische Klinik II, Lìbeck, 23562, Germany Jeanette Erdmann, Christina Loley 

& Heribert Schunkert * Universität zu Lìbeck, Medizinische Klinik II, Lìbeck, 23538, Germany Jeanette Erdmann, Michael Preuss & Heribert Schunkert * Deutsches Zentrum fìr

Herz-Kreislaufforschung e. V. (DZHK), Universität zu Lìbeck, Lìbeck, 23538, Germany Jeanette Erdmann & Heribert Schunkert * Department of General Practice and Primary health Care,

University of Helsinki, Helsinki, Finland Johan G. Eriksson * National Institute for Health and Welfare, Helsinki, 00271, Finland Johan G. Eriksson, Eero Kajantie & Leena Peltonen *

Helsinki University Central Hospital, Unit of General Practice, Helsinki, 00280, Finland Johan G. Eriksson * Folkhalsan Research Centre, Helsinki, 00250, Finland Johan G. Eriksson, Bo Isomaa

 & Tiinamaija Tuomi * Vasa Central Hospital, Vasa, 65130, Finland Johan G. Eriksson * Estonian Biocenter, Tartu, 51010, Estonia Tõnu Esko, Andres Metspalu, Mari Nelis & Maris

Teder-Laving * Department of Internal Medicine, Erasmus MC, Rotterdam, 3015GE, The Netherlands Karol Estrada, Carolina Medina-Gomez, Joyce B. J. van Meurs, Marjolein J. Peters, Fernando

Rivadeneira, André G. Uitterlinden & M. Carola Zillikens * Department of Social Medicine, MRC Centre for Causal Analyses in Translational Epidemiology, University of Bristol, Bristol,

BS8 2BN, UK David M Evans & Lavinia Paternoster * Division of Cardiovascular Epidemiology, Institute of Environmental Medicine, Karolinska Institutet, Stockholm, Sweden Ulf de Faire 

& Bruna Gigante * Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, 171 77, Sweden Tove Fall, Andrea Ganna, Henrik Grönberg, Stefan Gustafsson, Per

Hall, Erik Ingelsson, Yudi Pawitan, Nancy L. Pedersen, Emil Rehnberg & Patrik K. E. Magnusson * Cardiovascular Medicine, University of Oxford, Wellcome Trust Centre for Human Genetics,

Oxford, OX3 7BN, UK Martin Farrall, Anuj Goel & Hugh Watkins * Department of Clinical and Experimental Medicine, Epidemiology and Preventive Medicine Research Center, University of

Insubria, Varese, Italy Marco M. Ferrario * Department of Cardiology, Toulouse University School of Medicine, Rangueil Hospital, Toulouse, France Jean Ferrières * Division of Intramural

Research, National Heart, Lung and Blood Institute, Framingham Heart Study, Framingham, 01702, Massachusetts, USA Caroline S. Fox * Department of Genetics, University Medical Center

Groningen, University of Groningen, The Netherlands Lude Franke & Pim van der Harst * Department of Clinical Sciences, Genetic and Molecular Epidemiology Unit, Skåne University Hospital

Malmö, Lund University, Malmö, Sweden Paul W. Franks & Dmitry Shungin * Department of Nutrition, Harvard School of Public Health, Boston, MA, USA Paul W. Franks * Department of Public

Health & Clinical Medicine, Umeå University, Umeå, Sweden Paul W. Franks, Göran Hallmans & Dmitry Shungin * Center for Neurobehavioral Genetics, University of California, Los

Angeles, 90095, California, USA Nelson B. Freimer * Department of Genomics of Common Disease, School of Public Health, Imperial College London, London, W12 0NN, UK Philippe Froguel *

Department of Medicine, University of Maryland School of Medicine, Baltimore, 21201, Maryland, USA Mao Fu, Jeffrey R. O’Connell & Alan R. Shuldiner * University of Chicago, Chicago, IL

Pablo V. Gejman & Alan R. Sanders * Northshore University Healthsystem, Evanston, 60201, Ilinois, USA Pablo V. Gejman & Alan R. Sanders * Hagedorn Research Institute, Gentofte, 2820,

Denmark Anette P. Gjesing, Torben Hansen, Oluf Pedersen & Camilla Sandholt * Department of Medicine, University of Washington, Seattle, 98101, Washington, USA Nicole L. Glazer *

Cardiovascular Health Research Unit, University of Washington, Seattle, 98101, Washington, USA Nicole L. Glazer, Guo Li & Bruce M. Psaty * University of Melbourne, Parkville, 3010,

Australia Michael E. Goddard * Department of Primary Industries, Melbourne, 3001, Victoria, Australia Michael E. Goddard * Institute of Epidemiology, Helmholtz Zentrum Mìnchen - German

Research Center for Environmental Health, Neuherberg, 85764, Germany Harald Grallert, Iris M. Heid, Thomas Illig, Claudia Lamina & H-Erich Wichmann * Unit for Molecular Epidemiology,

Helmholtz Zentrum Mìnchen - German Research Center for Environmental Health, Neuherberg, Germany Harald Grallert & Thomas Illig * Research Unit for Molecular Epidemiology, Helmholtz

Zentrum Mìnchen - German Research Center for Environmental Health, Neuherberg, Germany Harald Grallert, Thomas Illig & Annette Peters * Department of Medicine III, Pathobiochemistry,

University of Dresden, Dresden, 01307, Germany Jìrgen Gräßler * Department of Medicine, University of Iceland, Reykjavik, Iceland Vilmundur Gudnason & Albert Vernon Smith * Metabolism

Initiative and Program in Medical and Population Genetics, Broad Institute, Cambridge, 02142, Massachusetts, USA Candace Guiducci, Joel N. Hirschhorn, Elizabeth K. Speliotes, Brian Thomson 

& Sailaja Vedantam * Department of Genetics and Pathology, Rudbeck Laboratory, University of Uppsala, Uppsala, SE-75185, Sweden Ulf Gyllensten, Wilmar Igl & Åsa Johansson *

Department of Immunology, Genetics and Pathology, Uppsala University, Sweden Ulf Gyllensten & Åsa Johansson * Division of Cardiovascular and Neuronal Remodelling, Multidisciplinary

Cardiovascular Research Centre, Leeds Institute of Genetics, Health and Therapeutics, University of Leeds, UK Alistair S. Hall * Department of Medicine, Atherosclerosis Research Unit,

Solna,Karolinska Institutet, Karolinska University Hospital, Stockholm, 171 76, Sweden Anders Hamsten & Rona J Strawbridge * Faculty of Health Science, University of Southern Denmark,

Odense, 5000, Denmark Torben Hansen * Medical Genetics Institute, Cedars-Sinai Medical Center, Los Angeles, 90048, California, USA Talin Haritunians * Laboratory of Epidemiology, Demography,

Biometry, National Institute on Aging, National Institutes of Health, Bethesda, 20892, Maryland, USA Tamara B. Harris & Lenore J. Launer * Department of Cardiology, University Medical

Center Groningen, University of Groningen, The Netherlands Pim van der Harst & Irene M. Leach * Department of Clinical Sciences/Obstetrics and Gynecology, University of Oulu, Oulu,

90014, Finland Anna-Liisa Hartikainen & Anneli Pouta * Department of Chronic Disease Prevention, National Institute for Health and Welfare, Chronic Disease Epidemiology and Prevention

Unit, Helsinki, 00014, Finland Aki S. Havulinna, Pekka Jousilahti, Seppo Koskinen & Veikko Salomaa * MRC Human Genetics Unit, Institute for Genetics and Molecular Medicine, Western

General Hospital, Edinburgh, EH4 2XU, Scotland, UK Caroline Hayward, Jennifer E. Huffman, Veronique Vitart & Alan F. Wright * Department of Psychiatry, Washington University School of

Medicine, St Louis, MO 63108, USA Andrew C Heath & Pamela A. Madden * Department of Child and Adolescent Psychiatry, University of Duisburg-Essen, Essen, 45147, Germany Johannes

Hebebrand & Anke Hinney * Department of Epidemiology and Preventive Medicine, Regensburg University Medical Center, Regensburg, 93053, Germany Iris M. Heid, Michael F. Leitzmann & 

Thomas W. Winkler * Public Health and Gender Studies, Institute of Epidemiology and Preventive Medicine, Regensburg University Medical Center, Regensburg, Germany Iris M. Heid & Thomas

W. Winkler * Institute of Epidemiology I, Helmholtz Zentrum Mìnchen - German Research Center for Environmental Health, Neuherberg, Germany Iris M. Heid & H-Erich Wichmann * Department of

Internal Medicine, VU University Medical Centre, Amsterdam, The Netherlands Martin den Heijer * Klinik und Poliklinik fìr Innere Medizin II, Universität Regensburg, Regensburg, 93053,

Germany Christian Hengstenberg * Regensburg University Medical Center, Innere Medizin II, Regensburg, 93053, Germany Christian Hengstenberg * Klinik und Poliklinik fìr Innere Medizin II,

Universitätklinikum Regensburg, Regensburg, 93053, Germany Christian Hengstenberg & Klaus Stark * Biocenter Oulu, University of Oulu, Oulu, 90014, Finland Karl-Heinz Herzig, Marjo-Riitta

Jarvelin & Marika Kaakinen * Department of Physiology, Institute of Biomedicine, University of Oulu, Oulu, 90014, Finland Karl-Heinz Herzig * Department of Psychiatry, Kuopio University

Hospital and University of Kuopio, Kuopio, 70210, Finland Karl-Heinz Herzig * Institute of Genetic Medicine, European Academy Bozen/Bolzano (EURAC), Affiliated Institute of the University

of Lìbeck, Lìbeck, Germany, Bolzano/Bozen, 39100, Italy Andrew A. Hicks, Irene Pichler, Peter P. Pramstaller & Claudia B. Volpato * Center for Biomedicine, European Academy Bozen/Bolzano

(EURAC), Affiliated Institute of the University of Lìbeck, Lìbeck, Germany, Bolzano/Bozen, 39100, Italy Andrew A. Hicks & Peter P. Pramstaller * Center for Biomedicine, European Academy

Bozen/Bolzano (EURAC), Affiliated Institute of the University of Lìbeck, Lìbeck, Germany, Bolzano/Bozen, 39100, Italy Andrew A. Hicks & Peter P. Pramstaller * Department of Epidemiology

and Public Health, University College London, 1-19 Torrington Place, London, WC1E 6BT, UK Aroon Hingorani, Mika Kivimaki, Mika Kivmaki, Meena Kumari & Claudia Langenberg * Divisions of

Genetics and Endocrinology and Program in Genomics, Children’s Hospital, Boston, 02115, Massachusetts, USA Joel N. Hirschhorn & Sailaja Vedantam * Department of Genetics, Harvard Medical

School, Boston, 02115, Massachusetts, USA Joel N. Hirschhorn & Sailaja Vedantam * Divisions of Genetics and Endocrinology and Centerfor Basic and Translational Obesity Research,

Children’s Hospital, Boston, 02115, Massachusetts, USA Joel N. Hirschhorn & Sailaja Vedantam * MRC Harwell, Harwell Science and Innovation Campus, Oxfordshire, OX11 0RD, UK Christopher

C. Holmes * Department of Statistics, University of Oxford, Oxford, OX1 3TG, UK Christopher C. Holmes & George Nicholson * Interfaculty Institute for Genetics and Functional Genomics,

Ernst-Moritz-Arndt-University Greifswald, Greifswald, 17487, Germany Georg Homuth & Alexander Teumer * Department of Nutrition, Harvard School of Public Health, Boston, 02115,

Massachusetts, USA Frank B. Hu, David Hunter, Lu Qi & Tsegaselassie Workalemahu * Department of Medicine, Channing Laboratory, Brigham and Women’s Hospital and Harvard Medical School,

Boston, 02115, Massachusetts, USA Frank B. Hu, David Hunter, Lu Qi & Tsegaselassie Workalemahu * Department of Epidemiology, Harvard School of Public Health, Boston, 02115,

Massachusetts, USA Frank B. Hu, David Hunter, Peter Kraft & Liming Liang * Department of Biostatistics andBioinformatics, Emory University, Atlanta, 30322, Georgia, USA Yi-Juan Hu *

School of Population Health, The University of Western Australia, Nedlands, 6009, WA, Australia Jennie Hui & Arthur W. Musk * Department of Internal Medicine, Institute of Clinical

Medicine, University of Oulu, Oulu, 90014, Finland Heikki Huikuri * Cardiovascular Genetics, British Heart Foundation Laboratories, Rayne Building, University College London, London, United

Kingdom Steve E. Humphries * School of Medicine and Pharmacology, The University of Western Australia, Nedlands, 6009, WA, Australia Joseph Hung * Department of Public Health and General

Practice, HUNT Research Centre, Norwegian University of Science and Technology, Levanger, 7600, Norway Kristian Hveem, Kirsti Kvaloy, Kristian Midthjell & Carl G. P. Platou * Centre For

Paediatric Epidemiolgy and Biostatistics/MRC Centre of Epidemiology for Child Health, University College of London Institute of Child Health, London, UK Elina Hyppönen & Chris Power *

Hannover Unified Biobank, Hannover Medical School, Hannover, 30625, Germany Thomas Illig * Division of Research, Kaiser Permanente Northern California, Oakland, 94612, California, USA Carlos

Iribarren * Department of Epidemiology and Biostatistics, University of California, San Francisco, San Francisco, 94107, California, USA Carlos Iribarren * Department of Social Services and

Health Care, Jakobstad, 68601, Finland Bo Isomaa * Core Genotyping Facility, SAIC-Frederick, Inc., NCI-Frederick, Frederick, 21702, Maryland, USA Kevin B. Jacobs & Zhaoming Wang *

School of Medicine and Pharmacology, University of Western Australia, Perth, 6009, Western Australia, Australia Alan L. James * Department of Physiology, Institute of Neuroscience and

Physiology, Sahlgrenska Academy, University of Gothenburg, Gothenburg, 405 30, Sweden John-Olov Jansson * Institute of Medical Biometry and Epidemiology, University of Marburg, Marburg,

35037, Germany Ivonne Jarick & Martina Mìller-Nurasyid * Institute of Health Sciences, University of Oulu, Oulu, 90014, Finland Marjo-Riitta Jarvelin & Marika Kaakinen * National

Institute for Health and Welfare, Oulu, 90101, Finland Marjo-Riitta Jarvelin & Anneli Pouta * Institute for Medical Informatics, Biometry and Epidemiology (IMIBE), University Hospital of

Essen, University of Duisburg-Essen, Essen, Germany Karl-Heinz Jöckel, Susanne Moebus, Sonali Pechlivanis, Carolin Pìtter & Andre Scherag * Department of Cancer Research and Molecular

Medicine, Faculty of Medicine, Norwegian University of Science and Technology (NTNU), Trondheim, N-7489, Norway Åsa Johansson * Uppsala Clinical Research Center, Uppsala university hospital,

Sweden Åsa Johansson * Clinical Pharmacology, William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary, University of London, London, UK Toby

Johnson * Research Centre for Prevention and Health, Glostrup University Hospital, Glostrup, 2600, Denmark Torben Jørgensen * Faculty of Health Science, University of Copenhagen, Copenhagen,

2100, Denmark Torben Jørgensen * Department of Chronic Disease Prevention, National Institute for Health and Welfare, Population Studies Unit, Turku, 20720, Finland Antti Jula * Department

of Epidemiology, School of Public Health, University of North Carolina at Chapel Hill, Chapel Hill, 27514, North Carolina, USA Anne E. Justice, Keri L. Monda & Kari E. North * Department

of Clinical Physiology, University of Tampere and Tampere University Hospital, Tampere, 33520, Finland Mika Kähönen * Hospital for Children and Adolescents, Helsinki University Central

Hospital and University of Helsinki, 00029 HUS, Finland Eero Kajantie * Department of Epidemiology and Medicine, Johns Hopkins Bloomberg School of Public Health, Baltimore, 21205, Maryland,

USA W. H. Linda Kao * Division of Gastroenterology, Massachusetts General Hospital, Boston, 02114, Massachusetts, USA Lee M. Kaplan & Elizabeth K. Speliotes * MGH Weight Center,

Massachusetts General Hospital, Boston, 02114, Massachusetts, USA Lee M. Kaplan * Department of Epidemiology and Population Health, Albert Einstein College of Medicine, Bronx, 10461, New

York, USA Robert C. Kaplan * Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, 00014, Finland Jaakko Kaprio, Johannes Kettunen, Kati Kristiansson, Aarno

Palotie, Niina Pellikka, Leena Peltonen, Markus Perola, Samuli Ripatti, Kaisa Silander, Ida Surakka & Elisabeth Widen * Department of Public Health, Finnish Twin Cohort Study, University

of Helsinki, Helsinki, 00014, Finland Jaakko Kaprio & Kirsi H. Pietiläinen * Department of Mental Health and Substance Abuse Services, National Institute for Health and Welfare, Unit

for Child and Adolescent Mental Health, Helsinki, 00271, Finland Jaakko Kaprio & Robert N. Luben * National Institute for Health and Welfare, Unit for Child and Adolescent Psychiatry,

Helsinki, Finland Jaakko Kaprio * NIHR Oxford Biomedical Research Centre, Churchill Hospital, Oxford, OX3 7LJ, UK Fredrik Karpe & Mark I. McCarthy * Oxford National Institute for Health

Research Biomedical Research Centre, Churchill Hospital, Old Road Headington, Oxford, OX3 7LJ, UK Fredrik Karpe & Mark I. McCarthy * Cardiovascular Research Center and Cardiology

Division, Massachusetts General Hospital, Boston, 02114, Massachusetts, USA Sekar Kathiresan * Center for Human Genetic Research, Massachusetts General Hospital, Boston, 02114,

Massachusetts, USA Sekar Kathiresan, Steven A. McCaroll, Shaun Purcell & Benjamin F. Voight * Program in Medical and Population Genetics, Broad Institute of Harvard and Massachusetts

Institute of Technology, Cambridge, 02142, Massachusetts, USA Sekar Kathiresan, Steven A. McCaroll, James Nemesh & Benjamin F. Voight * UKCRC Centre of Excellence for Public Health (NI)

Queens University, Belfast Frank Kee * Faculty of Medicine, Institute of Health Sciences, University of Oulu, Oulu, Finland Sirkka M. Keinanen-Kiukaanniemi * Unit of General Practice, Oulu

University Hospital, Oulu, Finland Sirkka M. Keinanen-Kiukaanniemi * Department of Public Health and Primary Care, Institute of Public Health, University of Cambridge, Cambridge, CB2 2SR, UK

Kay-Tee Khaw * Department of Epidemiology, Biostatistics and HTA, Radboud University Nijmegen Medical Centre, Nijmegen, 6500, HB, The Netherlands Lambertus A. Kiemeney, Femmie de Vegt &

 Sita H. Vermeulen * Department of Urology, Radboud University Nijmegen Medical Centre, Nijmegen, 6500, HB, The Netherlands Lambertus A. Kiemeney * Comprehensive Cancer Center East,

Nijmegen, 6501, BG, The Netherlands Lambertus A. Kiemeney * National Institute for Health and Welfare, Diabetes Prevention Unit, Helsinki, 00271, Finland Leena Kinnunen, Jaana Lindström, 

Jaakko Tuomilehto & Timo T. Valle * Department of Endocrinology, University Medical Center Groningen, University of Groningen, P.O. Box 30001, Groningen, 9700, RB, The Netherlands

Melanie M. Van der Klauw, Jana V. Van Vliet-Ostaptchouk & Bruce H. R. Wolffenbuttel * LURIC Study nonprofit LLC, Freiburg, Germany Marcus E. Kleber * Mannheim Institute of Public Health,

Social and Preventive Medicine, Medical Faculty of Mannheim, University of Heidelberg, Mannheim, Germany Marcus E. Kleber & Winfried März * Department of Internal Medicine II –

Cardiology, University of Ulm Medical Center, Ulm, Germany Wolfgang Koenig * Andrija Stampar School of Public Health, Medical School, University of Zagreb, Zagreb, 10000, Croatia Ivana

Kolcic, Ozren Polasek & Lina Zgaga * 1st Cardiology Department, Onassis Cardiac Surgery Center 356, Sygrou Ave., Athens, Greece Genovefa Kolovou * Institut fìr Medizinische Biometrie und

Statistik, Universität zu Lìbeck, Universitätsklinikum Schleswig-Holstein, Campus Lìbeck, Lìbeck, 23562, Germany Inke R. König, Christina Loley & Michael Preuss * Interdisciplinary

Centre for Clinical Research, University of Leipzig, Leipzig, 04103, Germany Peter Kovacs * Department of Biostatistics, Harvard School of Public Health, Boston, 02115, Massachusetts, USA

Peter Kraft & Liming Liang * Institut fìr Pharmakologie, Universität Greifswald, Greifswald, 17487, Germany Heyo K. Kroemer * Croatian Centre for Global Health, School of Medicine,

University of Split, Split, 21000, Croatia Vjekoslav Krzelj & Igor Rudan * MRC Unit for Lifelong Health & Ageing, London, UK Diana Kuh, Ken K. Ong & Andrew Wong * Department of

Chronic Disease Prevention, National Institute for Health and Welfare, Chronic Disease Epidemiology and Prevention Unit, Helsinki, 00271, Finland Kari Kuulasmaa, Veikko Salomaa & Jarmo

Virtamo * Department of Medicine, University of Kuopio and Kuopio University Hospital, Kuopio, 70210, Finland Johanna Kuusisto & Markku Laakso * Department of Medicine, University of

Eastern Finland, Kuopio Campus and Kuopio University Hospital, Kuopio, 70210, Finland Johanna Kuusisto & Markku Laakso * Finnish Institute of Occupational Health, Oulu, 90220, Finland

Jaana H. Laitinen * Kuopio Research Institute of Exercise Medicine, Kuopio, Finland Timo A. Lakka & Rainer Rauramaa * Division of Genetic Epidemiology, Department of Medical Genetics,

Molecular and Clinical Pharmacology, Innsbruck Medical University, Innsbruck, 6020, Austria Claudia Lamina * Institut inter-regional pour la sante (IRSA), La Riche, F-37521, France Olivier

Lantieri * Centre National de Genotypage, Evry, Paris, 91057, France G. Mark Lathrop * The Queensland Brain Institute, The University of Queensland, Brisbane, Queensland, Australia Sang Hong

Lee & Peter M. Visscher * Department of Clinical Chemistry, University of Tampere and Tampere University Hospital, Tampere, 33520, Finland Terho Lehtimäki * Department of Clinical

Chemistry, Fimlab Laboratories, University of Tampere and Tampere University Hospital, Tampere, 33520, Finland Terho Lehtimäki * Department of Medicine, Université de Montréal, Montreal, H3T

1J4, Quebec, Canada Guillaume Lettre * Stanford University School of Medicine, Stanford, 93405, California, USA Douglas F. Levinson * Department of Epidemiology, Tulane School of Public

Health and Tropical Medicine, New Orleans, 70112, LA, USA Shengxu Li * Department of Biostatistics, University of North Carolina, Chapel Hill, 27599, NC, USA Dan-Yu Lin * Department of

Medical Sciences, Uppsala University, Akademiska sjukhuset, Uppsala, 751 85, Sweden Lars Lind * Human Genetics, Genome Institute of Singapore, Singapore, 138672, Singapore Jianjun Liu *

Department of Internal Medicine, Istituto Auxologico Italiano, Verbania, Italy Antonio Liuzzi * Transplantation Laboratory, Haartman Institute, University of Helsinki, Helsinki, 00014,

Finland Marja-Liisa Lokki * The Charles Bronfman Institute of Personalized Medicine, Mount Sinai School of Medicine, New York, 10029, NY, USA Ruth J. F. Loos * Child Health and Development

Institute, Mount Sinai School of Medicine, New York, 10029, NY, USA Ruth J. F. Loos * Department of Preventive Medicine, Mount Sinai School of Medicine, New York, 10029, NY, USA Ruth J. F.

Loos * Department of Internal Medicine, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, Gothenburg, 413 45, Sweden Mattias Lorentzon, Claes Ohlsson & Liesbeth

Vandenput * Department of Twin Research and Genetic Epidemiology, King’s College London, London, SE1 7EH, UK Massimo Mangino, Nicole Soranzo & Timothy D. Spector * Università Vita-Salute

San Raffaele, Chair of Nephrology San Raffaele Scientific Institute, OU Nephrology and Dialysis, Milan, 20132, Italy Paolo Manunta * Department of Endocrinology, Diabetology and Nutrition,

Bichat-Claude Bernard University Hospital, Assistance Publique des Hôpitaux de Paris, Paris, F-75018, France Michel Marre * Cardiovascular Genetics Research Unit, Université Henri

Poincaré-Nancy 1, Nancy, 54000, France Michel Marre * Genetic Epidemiology Laboratory, Queensland Institute of Medical Research, 4006, Queensland, Australia Sarah E. Medland & Nicholas

G. Martin * Queensland Institute of Medical Research, 4029, Queensland, Australia Grant W. Montgomery, Sarah E. Medland, Nicholas G. Martin & Jian Yang * Synlab Academy, Mannheim,

Germany Winfried März * Department of Social Medicine, Avon Longitudinal Study of Parents and Children (ALSPAC) Laboratory, University of Bristol, Bristol, BS8 2BN, UK Wendy L. McArdle *

School of Social and Community Medicine, University of Bristol, UK Wendy L. McArdle * Department of Molecular Biology, Massachusetts General Hospital, Boston, 02114, Massachusetts, USA

Steven A. McCaroll & Benjamin F. Voight * Division of Health, Research Board, An Bord Taighde Sláinte, Dublin, 2, Ireland Anne McCarthy * Institute of Human Genetics, Klinikum rechts der

Isar der Technischen Universität Mìnchen, Munich, 81675, Germany Thomas Meitinger * Institute of Human Genetics, Helmholtz Zentrum Mìnchen - German Research Center for Environmental Health,

Neuherberg, 85764, Germany Thomas Meitinger * Department of Clinical Epidemiology and Biostatistics, McMasterUniversity, Hamilton, L8S 4L8, Ontario, Canada David Meyre * Human Genetics,

Leiden University Medical Center, Leiden, 2333, The Netherlands Josine L. Min * Merck Research Laboratories, Merck & Co., Inc., Boston, 02115, Massachusetts, USA Cliona Molony * Center

for Observational Research, Amgen, Thousands Oaks, 91320, CA Keri L. Monda * Molecular Epidemiology Laboratory, Queensland Institute of Medical Research, 4006, Queensland, Australia Grant W.

Montgomery * Genetics Division, GlaxoSmithKline, King of Prussia, Pennsylvania, 19406, USA Vincent Mooser * Medical Research Institute, University of Dundee, Ninewells Hospital and Medical

School, Dundee, DD1 9SY Andrew D. Morris & Colin N. A. Palmer * Institute of Human Genetics, University of Bonn, Bonn, Germany Thomas W. Mìhleisen & Markus M. Nöthen * Department of

Genomics, Life & Brain Center, University of Bonn, Bonn, Germany Thomas W. Mìhleisen & Markus M. Nöthen * Department of Medicine I, University Hospital Grosshadern,

Ludwig-Maximilians-Universität, Munich, Germany Martina Mìller-Nurasyid * Institute of Medical Informatics, Biometry and Epidemiology, Chair of Genetic Epidemiology,

Ludwig-Maximilians-Universität, Munich, Germany Martina Mìller-Nurasyid * Department of Respiratory Medicine, Sir Charles Gairdner Hospital, Nedlands, 6009, Australia Arthur W. Musk *

Analytic and Translational Genetics Unit, Massachusetts General Hospital, Boston, 02114, MA Benjamin M. Neale * MRC Harwell, Harwell, UK George Nicholson * Division of Cardiology,

Cardiovascular Laboratory, Helsinki University Central Hospital, Helsinki, 00029, Finland Markku S. Nieminen & Juha Sinisalo * Department of Community Medicine, Faculty of Health

Sciences, University of Tromsø, Tromsø, Norway Inger Njølstad & Tom Wilsgaard * Department of Clinical Medicine, Faculty of Health Sciences, University of Tromsø, Tromsø, Norway Inger

Njølstad * Department of Public Health, Section of Epidemiology, Aarhus University, Denmark Ellen A. Nohr * Unit of Genetic Epidemiology and Bioinformatics, Dept of Epidemiology, University

Medical Center Groningen, University of Groningen, P.O. Box 30001, Groningen, 9700, RB, The Netherlands Ilja M. Nolte & Harold Snieder * Department of Epidemiology, University of

Groningen, University Medical Center Groningen, The Netherlands Ilja M. Nolte & Ronald P. Stolk * Carolina Center for Genome Sciences, School of Public Health, University of North

Carolina Chapel Hill, Chapel Hill, 27514, North Carolina, USA Kari E. North * Neurogenetics Laboratory, Queensland Institute of Medical Research, 4006, Queensland, Australia Dale R. Nyholt *

Interdisciplinary Center Psychopathology and Emotion Regulation, University of Groningen, University Medical Center Groningen, The Netherlands Albertine J. Oldehinkel * Department of Human

Genetics, Leiden University Medical Center, Leiden, 2333, ZC, The Netherlands Gert-Jan van Ommen * Department of Clinical Genetics, Erasmus MC, Rotterdam, 3015GE, The Netherlands Ben A.

Oostra * Centre for Medical Systems Biology & Netherlands Consortium on Healthy Aging, Leiden, The Netherlands Ben A. Oostra * NHS Blood and Transplant, Cambridge Centre, Cambridge, CB2

0PT, UK Willem H. Ouwehand, Aparna Radhakrishnan, Augusto Rendon, Jennifer G. Sambrook & Jonathan C. Stephens * Department of Pathology and Molecular Medicine, McMaster University,

Hamilton, L8N3Z5, Ontario, Canada Guillaume Paré * Amgen, Cambridge, 02139, Massachusetts, USA Alex N. Parker * Department of Cardiovascular Medicine, University of Oxford, Level 6 West

Wing, John Radcliffe Hospital, Headley Way, Headington, OX3 9DU, Oxford John F. Peden * Illumina Inc. Cambridge, USA John F. Peden * Institute of Biomedical Sciences, University of

Copenhagen, Copenhagen, 2200, Denmark Oluf Pedersen * Faculty of Health Science, University of Aarhus, Aarhus, 8000, Denmark Oluf Pedersen * The Broad Institute of Harvard and MIT,

Cambridge, 02142, Massachusetts, USA Leena Peltonen & Shaun Purcell * Department of Medical Genetics, University of Helsinki, Helsinki, 00014, Finland Leena Peltonen * Department of

Psychiatry/EMGO Institute, VU University Medical Center, Amsterdam, 1081 BT, The Netherlands Brenda Penninx & Jan H. Smit * Department of Psychiatry, Leiden University Medical Centre,

Leiden, 2300 RC, The Netherlands Brenda Penninx * Department of Psychiatry, University Medical Centre Groningen, Groningen, 9713 GZ, The Netherlands Brenda Penninx * Institute of

Epidemiology II, Helmholtz Zentrum Mìnchen - German Research Center for Environmental Health, Neuherberg, Germany Annette Peters & Barbara Thorand * Munich Heart Alliance, Munich,

Germany Annette Peters * Department of Psychiatry, Obesity Research unit, Helsinki University Central Hospital, Helsinki, Finland Kirsi H. Pietiläinen & Aila Rissanen * Department of

Medicine, Levanger Hospital, The Nord-Trøndelag Health Trust, Levanger, 7600, Norway Carl G. P. Platou * Gen-Info Ltd, Zagreb, 10000, Croatia Ozren Polasek * Faculty of Medicine, University

of Split, Croatia Ozren Polasek * Department of Neurology, General Central Hospital, Bolzano, Italy Peter P. Pramstaller * Department of Neurology, University of Lìbeck, Lìbeck, Germany

Peter P. Pramstaller * Departments of Epidemiology, Medicine and Health Services, University of Washington, Seattle, 98195, Washington, USA Bruce M. Psaty * Group Health Research Institute,

Group Health, Seattle, 98101, Washington, USA Bruce M. Psaty * Department of Psychiatry, Harvard Medical School, Boston, 02115, Massachusetts, USA Shaun Purcell * Research Centre of Applied

and Preventive Cardiovascular Medicine, University of Turku, Turku, 20520, Finland Olli Raitakari * The Department of Clinical Physiology, Turku University Hospital, Turku, 20520, Finland

Olli Raitakari * The Department of Clinical Physiology and Nuclear Medicine, Turku University Hospital, Turku, 20520, Finland Olli Raitakari * Department of Clinical Physiology and Nuclear

Medicine, Kuopio University Hospital, Kuopio, Finland Rainer Rauramaa * MRC Biostatistics Unit, Institute of Public Health, Cambridge, UK Augusto Rendon * Department of Clinical Sciences,

Lund University, Malmö, 20502, Sweden Martin Ridderstråle * Finnish Diabetes Association, Kirjoniementie 15, Tampere, 33680, Finland Timo E. Saaristo * Pirkanmaa Hospital District, Tampere,

Finland Timo E. Saaristo * Medizinische Klinik II, Universität zu Lìbeck Ratzeburger Allee 160, Lìbeck, D-23538, Germany Hendrik Sager * Department of Cardiovascular Sciences, University of

Leicester, Glenfield Hospital, Leicester, LE3 9QP, UK Nilesh J. Samani * Leicester NIHR Biomedical Research Unit in Cardiovascular Disease, Glenfield Hospital, Leicester, LE3 9QP, UK Nilesh

J. Samani * South Karelia Central Hospital, Lappeenranta, 53130, Finland Jouko Saramies * Pacific Biosciences, Menlo Park, California, 94025, USA Eric E. Schadt * Sage Bionetworks, Seattle,

98109, Washington, USA Eric E. Schadt * Department of Genetics and Genomic Sciences, Mount Sinai School of Medicine, One Gustave L. Levy Place, Box 1498, New York, 10029-6574, NY, USA Eric

E. Schadt * Institute of Genomics and Multiscale Biology, Mount Sinai School of Medicine, One Gustave L. Levy Place, Box 1498, New York, 10029-6574, NY, USA Eric E. Schadt * Institute for

Community Medicine, University Medicine Greifswald, Greifswald, Germany Sabine Schipf * Laboratory of Genetics, National Institute on Aging, Baltimore, 21224, Maryland, USA David

Schlessinger * Institut fìr Klinische Molekularbiologie, Christian-Albrechts Universität, Kiel, Germany Stefan Schreiber * Department of Medicine III, Prevention and Care of Diabetes,

University of Dresden, Dresden, 01307, Germany Peter E. H. Schwarz * Geriatrics Research and Education Clinical Center, Baltimore Veterans Administration Medical Center, Baltimore, 21201,

Maryland, USA Alan R. Shuldiner * Department of Odontology, Umeå University, Sweden Dmitry Shungin * Azienda ospedaliera di Desio e Vimercate, Milano, Italy Stefano Signorini * Institute of

Preventive Medicine, Bispebjerg University Hospital, Copenhagen, and Novo Nordisk Foundation Center for Basic Metabolic Research, University of Copenhagen, Denmark Thorkild I. A. Sørensen *

Center for Computational Medicine and Bioinformatics, University of Michigan, Ann Arbor, Michigan, USA Elizabeth K. Speliotes * Department of Internal Medicine, Division of Gastroenterology,

University of Michigan, Ann Arbor, Michigan, USA Elizabeth K. Speliotes * University of Eastern Finland and Kuopio University Hospital, Kuopio, 70210, Finland Alena Stančáková * Regensburg

University Medical Center, Clinic and Policlinic for Internal Medicine II, Regensburg, 93053, Germany Klaus Stark * deCODE Genetics, Reykjavik, 101, Iceland Kari Stefansson, Valgerdur

Steinthorsdottir, Gudmar Thorleifsson, Unnur Thorsteinsdottir & G. Bragi Walters * Faculty of Medicine, University of Iceland, Reykjavík, 101, Iceland Kari Stefansson & Unnur

Thorsteinsdottir * Division of Community Health Sciences, St George’s, University of London, London, SW17 0RE, UK David P Strachan * Division of Population Health Sciences and Education, St

George’s, University of London, London, SW17 0RE, UK David P Strachan * Department of Medicine, University of Leipzig, Leipzig, 04103, Germany Michael Stumvoll & Anke Tönjes * LIFE Study

Centre, University of Leipzig, Leipzig, Germany Michael Stumvoll * University of Leipzig, IFB Adiposity Diseases, Leipzig, Germany Michael Stumvoll & Anke Tönjes * Uppsala University /

Dept. of Medical Sciences, Molecular Medicine, Uppsala, 751 85, Sweden Ann-Christine Syvanen * Coordination Centre for Clinical Trials, University of Leipzig, Härtelstr. 16-18, Leipzig,

04103, Germany Anke Tönjes * INSERM UMR_S 937, ICAN Institute, Pierre et Marie Curie Medical School, Paris, 75013, France David-Alexandre Tregouet * Department of Pharmacological Sciences,

University of Milan, Monzino Cardiology Center, IRCCS, Milan, Italy Elena Tremoli * Department of Clinical and Experimental Cardiology, Heart Failure Research Centre, Academic Medical

Center, Amsterdam, The Netherlands Mieke D. Trip * Department of Medicine, Helsinki University Central Hospital, Helsinki, 00290, Finland Tiinamaija Tuomi * Research Program of Molecular

Medicine, University of Helsinki, Helsinki, 00014, Finland Tiinamaija Tuomi * Department of Public Health, Hjelt Institute, University of Helsinki, Helsinki, 00014, Finland Jaakko Tuomilehto

* South Ostrobothnia Central Hospital, Seinajoki, 60220, Finland Jaakko Tuomilehto * Red RECAVA Grupo RD06/0014/0015, Hospital Universitario La Paz, Madrid, 28046, Spain Jaakko Tuomilehto *

Centre for Vascular Prevention, Danube-University Krems, Krems, 3500, Austria Jaakko Tuomilehto * Department of Oncology, University of Cambridge, Cambridge, CB1 8RN, UK Jonathan Tyrer *

Department of Public Health and Clinical Nutrition, University of Eastern Finland, Finland Matti Uusitupa * Research Unit, Kuopio University Hospital, Kuopio, Finland Matti Uusitupa *

Department of Human Genetics, Radboud University Nijmegen Medical Centre, PO Box 9101, Nijmegen, 6500, HB, The Netherlands Sita H. Vermeulen * Department of Medicine, University of Turku and

Turku University Hospital, Turku, 20520, Finland Jorma Viikari * Queensland Statistical Genetics Laboratory, Queensland Institute of Medical Research, 4006, Queensland, Australia Jian Yang 

& Peter M. Visscher * Department of Internal Medicine, Centre Hospitalier Universitaire Vaudois (CHUV) University Hospital, Lausanne, 1011, Switzerland Peter Vollenweider & Gérard

Waeber * Institut fìr Community Medicine, Greifswald, 17489, Germany Henry Völzke * Institute for Community Medicine, Ernst-Moritz-Arndt-University Greifswald, Greifswald, Germany Henry

Völzke * Institut fìr Klinische Chemie und Laboratoriumsmedizin, Universität Greifswald, Greifswald, 17475, Germany Henri Wallaschofski * Institute of Clinical Chemistry and Laboratory

Medicine, University Medicine Greifswald, Greifswald, 17475, Germany Henri Wallaschofski * Department of Preventive Medicine, Keck School of Medicine, University of Southern California, Los

Angeles, 90089, California, USA Richard M. Watanabe * Klinikum Grosshadern, Munich, 81377, Germany H-Erich Wichmann * Ludwig-Maximilians-Universität, Institute of Medical Informatics,

Biometry and Epidemiology, Chair of Epidemiology, Munich, 81377, Germany H-Erich Wichmann * Institute of Medical Informatics, Biometry and Epidemiology, Chair of Epidemiology,

Ludwig-Maximilians-Universität, and Klinikum Grosshadern, Munich, Germany H-Erich Wichmann * Klinikum Grosshadern, Munich, Germany H-Erich Wichmann * Cardiology Group,

Frankfurt-Sachsenhausen, Germany Bernhard R. Winkelmann * Steno Diabetes Center, Gentofte, 2820, Denmark Daniel R. Witte * Centre for Public Health, Queen’s University, Belfast, UK John W.

G. Yarnell * Department of Physiatrics, Lapland Central Hospital, Rovaniemi, 96101, Finland Paavo Zitting * Genetic and Genomic Epidemiology Unit, Wellcome Trust Centre for Human Genetics,

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publications You can also search for this author inPubMed Google Scholar CONSORTIA THE LIFELINES COHORT STUDY * Behrooz Z. Alizadeh * , H. Marike Boezen * , Lude Franke * , Pim van der Harst

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* , Michael Preuss * , Jackie F. Price * , Inga Prokopenko * , Michael A. Province * , Bruce M. Psaty * , Shaun Purcell * , Carolin Pìtter * , Lu Qi * , Thomas Quertermous * , Aparna

Radhakrishnan * , Olli Raitakari * , Joshua C. Randall * , Rainer Rauramaa * , Nigel W. Rayner * , Emil Rehnberg * , Augusto Rendon * , Martin Ridderstråle * , Paul M. Ridker * , Samuli

Ripatti * , Aila Rissanen * , Fernando Rivadeneira * , Carlo Rivolta * , Neil R. Robertson * , Lynda M. Rose * , Igor Rudan * , Timo E. Saaristo * , Hendrik Sager * , Veikko Salomaa * , 

Nilesh J. Samani * , Jennifer G. Sambrook * , Alan R. Sanders * , Camilla Sandholt * , Serena Sanna * , Jouko Saramies * , Eric E. Schadt * , Andre Scherag * , Sabine Schipf * , David

Schlessinger * , Stefan Schreiber * , Heribert Schunkert * , Peter E. H. Schwarz * , Laura J. Scott * , Jianxin Shi * , So-Youn Shin * , Alan R. Shuldiner * , Dmitry Shungin * , Stefano

Signorini * , Kaisa Silander * , Juha Sinisalo * , Boris Skrobek * , Jan H. Smit * , Albert Vernon Smith * , George Davey Smith * , Harold Snieder * , Nicole Soranzo * , Thorkild I. A.

Sørensen * , Ulla Sovio * , Timothy D. Spector * , Elizabeth K. Speliotes * , Alena Stančáková * , Klaus Stark * , Kari Stefansson * , Valgerdur Steinthorsdottir * , Jonathan C. Stephens * ,

 Kathleen Stirrups * , Ronald P. Stolk * , David P Strachan * , Rona J Strawbridge * , Heather M. Stringham * , Michael Stumvoll * , Ida Surakka * , Amy J. Swift * , Ann-Christine Syvanen *

, Mari-Liis Tammesoo * , Maris Teder-Laving * , Tanya M. Teslovich * , Alexander Teumer * , Eirini V. Theodoraki * , Brian Thomson * , Barbara Thorand * , Gudmar Thorleifsson * , Unnur

Thorsteinsdottir * , Nicholas John Timpson * , Anke Tönjes * , David-Alexandre Tregouet * , Elena Tremoli * , Mieke D. Trip * , Tiinamaija Tuomi * , Jaakko Tuomilehto * , Jonathan Tyrer * , 

Manuela Uda * , André G. Uitterlinden * , Gianluca Usala * , Matti Uusitupa * , Timo T. Valle * , Liesbeth Vandenput * , Vincent Vatin * , Sailaja Vedantam * , Femmie de Vegt * , Sita H.

Vermeulen * , Jorma Viikari * , Jarmo Virtamo * , Peter M. Visscher * , Veronique Vitart * , Jana V. Van Vliet-Ostaptchouk * , Benjamin F. Voight * , Peter Vollenweider * , Claudia B.

Volpato * , Henry Völzke * , Gérard Waeber * , Lindsay L. Waite * , Henri Wallaschofski * , G. Bragi Walters * , Zhaoming Wang * , Nicholas J. Wareham * , Richard M. Watanabe * , Hugh

Watkins * , Michael N. Weedon * , Ryan Welch * , Robert J. Weyant * , Eleanor Wheeler * , Charles C. White * , H-Erich Wichmann * , Elisabeth Widen * , Sarah H. Wild * , Gonneke Willemsen *

, Cristen J. Willer * , Tom Wilsgaard * , James F. Wilson * , Sophie van Wingerden * , Bernhard R. Winkelmann * , Thomas W. Winkler * , Daniel R. Witte * , Jacqueline C. M. Witteman * , 

Bruce H. R. Wolffenbuttel * , Andrew Wong * , Andrew R. Wood * , Tsegaselassie Workalemahu * , Alan F. Wright * , Jian Yang * , John W. G. Yarnell * , Lina Zgaga * , Jing Hua Zhao * , M.

Carola Zillikens * , Paavo Zitting *  & Krina T. Zondervan CONTRIBUTIONS M.R.R., J.Y. and P.M.V. conceived and designed the study. M.R.R., A.K. and M.G. analysed the data. M.R.R. devised

and performed the simulations. A.A.E.V., W.J.P., A.A., B.Z., S.M. provided statistical support. 23andMe Inc., The LifeLines cohort, GIANT consortium, G.W.M., N.G.M., M.L., P.L., D.C.,

J.V.V.O., M.B.M., H.S., W.G.I., P.K.E.M, N.L.P, M.McG. and K.E.N. provided study oversight, sample collection and management. M.R.R. and P.M.V. derived the theory and wrote the manuscript.

All collaborators reviewed and approved the final manuscript. CORRESPONDING AUTHORS Correspondence to Matthew R. Robinson or Peter M. Visscher. ETHICS DECLARATIONS COMPETING INTERESTS The

authors declare no competing interests. SUPPLEMENTARY INFORMATION SUPPLEMENTARY INFORMATION Supplementary Figures 1–9, Supplementary Table 1, Supplementary Note, Supplementary Methods,

Supplementary References, Author Lists (PDF 3325 kb) RIGHTS AND PERMISSIONS Reprints and permissions ABOUT THIS ARTICLE CITE THIS ARTICLE Robinson, M., Kleinman, A., Graff, M. _et al._

Genetic evidence of assortative mating in humans. _Nat Hum Behav_ 1, 0016 (2017). https://doi.org/10.1038/s41562-016-0016 Download citation * Received: 10 May 2016 * Accepted: 03 November

2016 * Published: 09 January 2017 * DOI: https://doi.org/10.1038/s41562-016-0016 SHARE THIS ARTICLE Anyone you share the following link with will be able to read this content: Get shareable

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