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Genome-wide association analyses of risk tolerance and risky behaviors in over 1 million individuals identify hundreds of loci and shared genetic influences

Nature Genetics volume 51, pages 245–257 (2019)Cite this article

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Abstract

Humans vary substantially in their willingness to take risks. In a combined sample of over 1 million individuals, we conducted genome-wide association studies (GWAS) of general risk tolerance, adventurousness, and risky behaviors in the driving, drinking, smoking, and sexual domains. Across all GWAS, we identified hundreds of associated loci, including 99 loci associated with general risk tolerance. We report evidence of substantial shared genetic influences across risk tolerance and the risky behaviors: 46 of the 99 general risk tolerance loci contain a lead SNP for at least one of our other GWAS, and general risk tolerance is genetically correlated (\(\left| {\hat r_{\mathrm{g}}} \right|\) ~ 0.25 to 0.50) with a range of risky behaviors. Bioinformatics analyses imply that genes near SNPs associated with general risk tolerance are highly expressed in brain tissues and point to a role for glutamatergic and GABAergic neurotransmission. We found no evidence of enrichment for genes previously hypothesized to relate to risk tolerance.

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Fig. 1: Manhattan plots.
Fig. 2: Genetic correlations with general risk tolerance.
Fig. 3: Results from selected biological analyses.

Data availability

GWAS summary statistics can be downloaded from http://www.thessgac.org/data. SNP-level summary statistics from analyses based entirely or in part on 23andMe data can only be reported for up to 10,000 SNPs. For general risk tolerance, we provide association results for all SNPs that passed quality-control filters in a GWAS meta-analysis of general risk tolerance that excludes the research participants from 23andMe; we also provide association results from the complete GWAS (which includes data from 23andMe) for all lead SNPs identified in our discovery GWAS and MTAG analysis of general risk tolerance and for the 4,000 most significant SNPs in the meta-analysis of the discovery and replication GWAS of risk tolerance. For adventurousness, we provide association results from the complete GWAS (which includes only data from 23andMe) for all lead SNPs and for the next 4,000 most significant SNPs. For automobile speeding propensity, drinks per week, ever smoker, number of sexual partners, and the first PC of the four risky behaviors, we provide association results from the complete GWAS for all SNPs that passed quality-control filters. Contact information for the cohorts included in this paper can be found in the Supplementary Note.

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Acknowledgements

This research was carried out under the auspices of the Social Science Genetic Association Consortium. The research was also conducted using the UK Biobank Resource under application number 11425. The study was supported by funding from the Ragnar Söderberg Foundation (E9/11 and E42/15); the Swedish Research Council (421-2013-1061); the Jan Wallander and Tom Hedelius Foundation; an ERC Consolidator Grant to Philipp Koellinger (647648 EdGe); the Pershing Square Fund of the Foundations of Human Behavior; the Open Philanthropy Project; the National Institute on Aging, National Institutes of Health through grants P01-AG005842, P01-AG005842-20S2, P30-AG012810, and T32-AG000186-23 to the National Bureau of Economic Research and R01-AG042568-02 to the University of Southern California; the government of Canada through Genome Canada and the Ontario Genomics Institute (OGI-152); and the Social Sciences and Humanities Research Council of Canada. We thank the International Cannabis Consortium, the eQTLgen Consortium, and the Psychiatric Genomics Consortium for sharing summary statistics from the GWAS of lifetime cannabis use, eQTL summary statistics, and summary statistics from the GWAS of ADHD, respectively. A full list of acknowledgments is provided in the Supplementary Note.

Author information

Author notes

  1. These authors contributed equally: Richard Karlsson Linnér, Pietro Biroli, Edward Kong, S. Fleur W. Meddens, Robbee Wedow.

  2. These authors jointly supervised this work: Daniel J. Benjamin, Philipp D. Koellinger, Jonathan P. Beauchamp.

  3. A list of members and affiliations appears in the Supplementary Note.

Authors and Affiliations

  1. Department of Complex Trait Genetics, Center for Neurogenomics and Cognitive Research, Amsterdam Neuroscience, VU University Amsterdam, Amsterdam, the Netherlands

    Richard Karlsson Linnér, S. Fleur W. Meddens, Anke R. Hammerschlag, Aysu Okbay, Ronald de Vlaming, Erdogan Taskesen, Aysu Okbay, S. Fleur W. Meddens, Ronald de Vlaming, Christiaan de Leeuw, Danielle Posthuma, Philipp D. Koellinger, Danielle Posthuma & Philipp D. Koellinger

  2. Erasmus University Rotterdam Institute for Behavior and Biology, Erasmus School of Economics, Erasmus University Rotterdam, Rotterdam, the Netherlands

    Richard Karlsson Linnér, S. Fleur W. Meddens, Cornelius A. Rietveld, Ronald de Vlaming, Cornelius A. Rietveld, S. Fleur W. Meddens, Ronald de Vlaming, Patrick J. F. Groenen, A. Roy Thurik, Patrick J. F. Groenen & A. Roy Thurik

  3. Department of Economics, VU University Amsterdam, Amsterdam, the Netherlands

    Richard Karlsson Linnér, S. Fleur W. Meddens, Aysu Okbay, Ronald de Vlaming, Aysu Okbay, S. Fleur W. Meddens, Ronald de Vlaming, Philipp D. Koellinger & Philipp D. Koellinger

  4. Department of Economics, University of Zurich, Zurich, Switzerland

    Pietro Biroli & Christian L. Zünd

  5. Department of Economics, Harvard University, Cambridge, MA, USA

    Edward Kong, Olga Rostapshova, David I. Laibson & David Laibson

  6. Stanley Center for Psychiatric Research, Broad Institute of MIT and Harvard, Cambridge, MA, USA

    Robbee Wedow, Chia-Yen Chen, Patrick Turley, Aarno Palotie & Patrick Turley

  7. Department of Epidemiology, Harvard T. H. Chan School of Public Health, Boston, MA, USA

    Robbee Wedow, Albert Hofman & Albert Hofman

  8. Analytic Translational Genetics Unit, Massachusetts General Hospital and Harvard Medical School, Boston, MA, USA

    Robbee Wedow, Chia-Yen Chen, Patrick Turley & Patrick Turley

  9. Department of Sociology, Harvard University, Cambridge, MA, USA

    Robbee Wedow

  10. Center for the Advancement of Value in Musculoskeletal Care, Hospital for Special Surgery, New York, NY, USA

    Mark Alan Fontana & Mark Alan Fontana

  11. Department of Healthcare Policy and Research, Weill Cornell Medical College, Cornell University, New York, NY, USA

    Mark Alan Fontana & Mark Alan Fontana

  12. Amsterdam School of Economics, University of Amsterdam, Amsterdam, the Netherlands

    Maël Lebreton & Maël P. Lebreton

  13. Amsterdam Brain and Cognition, University of Amsterdam, Amsterdam, the Netherlands

    Maël Lebreton & Maël P. Lebreton

  14. Department of Economics, University of Toronto, Toronto, Ontario, Canada

    Stephen P. Tino, Jonathan P. Beauchamp & Jonathan P. Beauchamp

  15. Biological Psychology, Vrije Universiteit Amsterdam, Amsterdam, the Netherlands

    Abdel Abdellaoui, Michel G. Nivard, Dorret I. Boomsma, Abdel Abdellaoui, Jouke-Jan Hottenga, Gonneke Willemsen, Dorret I. Boomsma, Conor C. Dolan & Dorret I. Boomsma

  16. Department of Applied Economics, Erasmus School of Economics, Erasmus University Rotterdam, Rotterdam, the Netherlands

    Cornelius A. Rietveld & Cornelius A. Rietveld

  17. Department of Epidemiology, Erasmus Medical Center, Rotterdam, the Netherlands

    Cornelius A. Rietveld, Frank J. van Rooij, Cornelius A. Rietveld, Sven J. van der Lee, Najaf Amin, Frank J. van Rooij, Cornelia M. van Duijn, Henning Tiemeier, Albert Hofman, Albert Hofman, Mohammad A. Ikram & Henning Tiemeier

  18. Novo Nordisk Foundation Center for Basic Metabolic Research, University of Copenhagen, Copenhagen, Denmark

    Pascal N. Timshel, Tune H. Pers, Pascal Timshel, Tarunveer S. Ahluwalia, Thorkild I. A. Sørensen & Tune H. Pers

  19. Department of Epidemiology Research, Statens Serum Institut, Copenhagen, Denmark

    Pascal N. Timshel, Tune H. Pers, Pascal Timshel & Tune H. Pers

  20. Institute for Molecular Bioscience, The University of Queensland, Brisbane, Australia

    Maciej Trzaskowski, Yang Wu, Futao Zhang, Grant W. Montgomery, Peter M. Visscher, Jian Yang, Futao Zhang, Grant W. Montgomery, Jacob Gratten, Jian Yang, Peter M. Visscher, Jian Yang & Jacob Gratten

  21. Institute for Social and Economic Research, University of Essex, Colchester, UK

    Yanchun Bao, Melissa C. Smart & Meena Kumari

  22. Seminar for Statistics, Department of Mathematics, ETH Zurich, Zurich, Switzerland

    Laura Buzdugan

  23. Department of Economics, University of Zurich, Zurich, Switzerland

    Laura Buzdugan & Ernst Fehr

  24. Stuyvesant High School, New York, NY, USA

    Ann H. Caplin

  25. Psychiatric & Neurodevelopmental Genetics Unit, Massachusetts General Hospital, Boston, MA, USA

    Chia-Yen Chen & Aarno Palotie

  26. Nuffield Department of Population Health, University of Oxford, Oxford, UK

    Peter Eibich & Peter Eibich

  27. Socio-Economic Panel Study, DIW Berlin, Berlin, Germany

    Peter Eibich, Peter Eibich, Martin Kroh & Gert G. Wagner

  28. Max Planck Institute for Demographic Research, Rostock, Germany

    Peter Eibich & Peter Eibich

  29. Research, 23andMe, Inc., Mountain View, CA, USA

    Pierre Fontanillas, Aaron Kleinman, Michelle Agee, Babak Alipanahi, Robert K. Bell, Katarzyna Bryc, Sarah L. Elson, Nicholas A. Furlotte, Karen E. Huber, Nadia K. Litterman, Jennifer C. McCreight, Matthew H. McIntyre, Joanna L. Mountain, Carrie A. M. Northover, Steven J. Pitts, J. Fah Sathirapongsasuti, Olga V. Sazonova, Janie F. Shelton, Suyash Shringarpure, Chao Tian, Joyce Y. Tung, Vladimir Vacic, Catherine H. Wilson, Nicholas A. Furlotte, David A. Hinds, Joyce Y. Tung, Adam Auton & David A. Hinds

  30. Barcelona Institute for Global Health (ISGlobal), Barcelona, Spain

    Juan R. Gonzalez

  31. Universitat Pompeu Fabra (UPF), Barcelona, Spain

    Juan R. Gonzalez

  32. CIBER Epidemiología y Salud Pública (CIBERESP), Madrid, Spain

    Juan R. Gonzalez

  33. Centre for Global Health Research, Usher Institute for Population Health Sciences and Informatics, University of Edinburgh, Edinburgh, Scotland

    Peter K. Joshi, Katharina E. Schraut, Harry Campbell, Peter K. Joshi, Igor Rudan, Ozren Polasek, James F. Wilson, David W. Clark, Paul R. H. J. Timmers & James F. Wilson

  34. Center for Life Course Health Research, University of Oulu, Oulu, Finland

    Ville Karhunen & Minna Männikkö

  35. Department of Economics, University of California San Diego, La Jolla, CA, USA

    Remy Z. Levin

  36. Genetic and Molecular Epidemiology Group, Lübeck Interdisciplinary Platform for Genome Analytics, Institutes of Neurogenetics & Cardiogenetics, University of Lübeck, Lübeck, Germany

    Christina M. Lill, Lars Bertram & Lars Bertram

  37. Team Loyalty BV, Hoofddorp, the Netherlands

    Gerardus A. Meddens & Gerardus A. Meddens

  38. Departament de Ciències Experimentals i de la Salut, Universitat Pompeu Fabra, Barcelona, Spain

    Gerard Muntané, Carlos Morcillo-Suarez & Arcadi Navarro

  39. Research Department, Hospital Universitari Institut Pere Mata, Institut d’Investigació Sanitària Pere Virgili (IISPV), Reus, Spain

    Gerard Muntané

  40. Centro de Investigación Biomédica en Red en Salud Mental (CIBERSAM), Reus, Spain

    Gerard Muntané

  41. Department of Psychiatry, University of California San Diego, La Jolla, CA, USA

    Sandra Sanchez-Roige, Abraham A. Palmer & Murray B. Stein

  42. Institute for Behavioral Genetics, University of Colorado Boulder, Boulder, CO, USA

    Jason D. Boardman

  43. Institute of Behavioral Science, University of Colorado Boulder, Boulder, CO, USA

    Jason D. Boardman

  44. Department of Sociology, University of Colorado Boulder, Boulder, CO, USA

    Jason D. Boardman

  45. Department of Economics and Finance, Oulu Business School, University of Oulu, Oulu, Finland

    Andrew Conlin & Rauli Svento

  46. Department of Economics, University of Konstanz, Konstanz, Germany

    Urs Fischbacher

  47. Thurgau Institute of Economics, Kreuzlingen, Switzerland

    Urs Fischbacher

  48. Department of Econometrics, Erasmus University Rotterdam, Rotterdam, Rotterdam, the Netherlands

    Patrick J. F. Groenen & Patrick J. F. Groenen

  49. Department of Sociology, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA

    Kathleen Mullan Harris

  50. Carolina Population Center, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA

    Kathleen Mullan Harris

  51. Unit of Psychiatry Research, University of Fribourg, Fribourg, Switzerland

    Gregor Hasler

  52. Family Medicine and Public Health, University of California San Diego, La Jolla, CA, USA

    Sonia Jain & Murray B. Stein

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

    Patrick Sullivan, Robert Karlsson, Paul Lichtenstein, Nancy L. Pedersen, Patrik K. E. Magnusson, Robert Karlsson & Patrik K. E. Magnusson

  54. Department of Health Care Policy, Harvard Medical School, Boston, MA, USA

    Ronald C. Kessler

  55. SURFsara, Amsterdam, the Netherlands

    Maarten Kooyman

  56. Peter Boris Centre for Addictions Research, McMaster University/St. Joseph’s Healthcare Hamilton, Hamilton, Ontario, Canada

    James MacKillop

  57. Homewood Research Institute, Guelph, Ontario, Canada

    James MacKillop

  58. Department of Integrative Physiology, University of Colorado Boulder, Boulder, CO, USA

    Matthew B. McQueen

  59. Department of Economics, University of Munich, Munich, Germany

    Klaus M. Schmidt

  60. Department of Economics, University of Cologne, Cologne, Germany

    Matthias Sutter

  61. Experimental Economics Group, Max Planck Institute for Research into Collective Goods, Bonn, Germany

    Matthias Sutter

  62. Department of Public Finance, University of Innsbruck, Innsbruck, Austria

    Matthias Sutter

  63. Montpellier Business School, Montpellier, France

    A. Roy Thurik & A. Roy Thurik

  64. Department of Internal Medicine, Erasmus Medical Centre, Rotterdam, the Netherlands

    Joyce van Meurs, Joost Verlouw, André G. Uitterlinden & André G. Uitterlinden

  65. UCL Genetics Institute, Department of Genetics, Evolution and Environment, University College London, London, UK

    Jon White

  66. Psychiatry and Behavioral Neuroscience, University of Chicago, Chicago, IL, USA

    Alan R. Sanders, Pablo V. Gejman & Harriet de Wit

  67. Queensland Brain Institute, The University of Queensland, Brisbane, Australia

    Jian Yang, Guo-Bo Chen, Zhihong Zhu, Andrew Bakshi, Riccardo E. Marioni, Anna A. E. Vinkhuyzen, Jian Yang & Jian Yang

  68. Dept of Psychology, University of Olso, Oslo, Norway

    Lars Bertram & Lars Bertram

  69. School of Public Health, Imperial College London, London, UK

    Lars Bertram & Lars Bertram

  70. Estonian Genome Center, University of Tartu, Tartu, Estonia

    Tõnu Esko, Anette Kalnapenkis, Silva Kasela, Jaanika Kronberg-Guzman, Viktorija Kukushkina, Andres Metspalu, Lili Milani, Natalia Pervjakova, Evelin Mihailov, Natalia Pervjakova, Reedik Mägi, Lili Milani, Andres Metspalu, Tõnu Esko & Tõnu Esko

  71. Department of Economics, Stockholm School of Economics, Stockholm, Sweden

    Magnus Johannesson & Magnus Johannesson

  72. Center for Translational Bioethics and Health Care Policy, Geisinger Health System, Danville, PA, USA

    Michelle N. Meyer & Michelle N. Meyer

  73. Centre for Genomic Regulation, Barcelona Institute of Science and Technology, Barcelona, Spain

    Arcadi Navarro

  74. Institució Catalana de Recerca i Estudis Avançats (ICREA), Barcelona, Catalonia, Spain

    Arcadi Navarro

  75. Institute for Genomic Medicine, University of California San Diego, La Jolla, CA, USA

    Abraham A. Palmer

  76. Department of Clinical Genetics, VU Medical Centre, Amsterdam, the Netherlands

    Danielle Posthuma & Danielle Posthuma

  77. Department of Economics, Johannes Gutenberg University, Mainz, Germany

    Daniel Schunk

  78. Behavioral and Health Genomics Center, Center for Economic and Social Research, University of Southern California, Los Angeles, CA, USA

    Patrick Turley, Daniel J. Benjamin, Patrick Turley & Daniel J. Benjamin

  79. Department of Psychiatry, Center for the Study of Traumatic Stress, Uniformed Services University of the Health Sciences, Bethesda, MD, USA

    Robert J. Ursano

  80. Max Planck Institute for Human Development, Berlin, Germany

    Tian Liu & Gert G. Wagner

  81. MRC Human Genetics Unit, Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, Scotland

    Jennifer E. Huffman, Jonathan Marten, Caroline Hayward, Veronique Vitart, James F. Wilson, Alan F. Wright & James F. Wilson

  82. Mater Medical Research Institute, Translational Research Institute, Brisbane, Australia

    Jacob Gratten & Jacob Gratten

  83. Department of Psychology, University of Minnesota Twin Cities, Minneapolis, MN, USA

    James J. Lee, Michael B. Miller, William G. Iacono, Matt McGue, Robert F. Krueger & James J. Lee

  84. Department of Economics, New York University, New York, NY, USA

    Kevin Thom, David Cesarini & David Cesarini

  85. Department of Economics, University of Southern California, Los Angeles, CA, USA

    Daniel J. Benjamin & Daniel J. Benjamin

  86. National Bureau of Economic Research, Cambridge, MA, USA

    Daniel J. Benjamin & Daniel J. Benjamin

  87. German Institute for Economic Research, DIW Berlin, Berlin, Germany

    Philipp D. Koellinger & Philipp D. Koellinger

  88. Computational Biology, Ontario Institute for Cancer Research, Toronto, Canada

    Mawussé Agbessi, Isabel Alves, Philip Awadalla & Marie-Julie Favé

  89. Department of Public Health Sciences, University of Chicago, Chicago, IL, USA

    Habibul Ahsan, Brandon Pierce & Lin Tong

  90. Singapore Immunology Network, Agency for Science, Technology and Research, Singapore, Singapore

    Anand Andiappan, Bernett Lee & Olaf Rotzchke

  91. Department of Computer Science, Johns Hopkins University, Baltimore, MD, USA

    Alexis Battle, Yungil Kim & Ashis Saha

  92. Heart Center Leipzig, Universität Leipzig, Leipzig, Germany

    Frank Beutner

  93. Department of Genetics, University Medical Center Groningen, University of Groningen, Groningen, the Netherlands

    Marc Jan Bonder, Annique Claringbould, Patrick Deelen, Lude Franke, Urmo Võsa, Harm-Jan Westra, Harm-Jan Westra, Lude Franke & Pim van der Harst

  94. Cardiovascular Health Research Unit, Departments of Epidemiology, Medicine, and Health Services, University of Washington, Seattle, WA, USA

    Mark Christiansen & Bruce M. Psaty

  95. Medical School, University of Exeter, Exeter, UK

    Timothy Frayling & Hanieh Yaghootkar

  96. Department of Medicine, University of Washington, Seattle, WA, USA

    Sina A. Gharib

  97. Kaiser Permanente Washington Health Research Institute, Seattle, WA, USA

    Sina A. Gharib

  98. School of Biological Sciences, Georgia Tech, Atlanta, GA, USA

    Gregory Gibson, Urko M. Marigorta & Biao Zeng

  99. Leiden University Medical Centre, Leiden, the Netherlands

    Bastiaan Heijmans

  100. MRC Integrative Epidemiology Unit, University of Bristol, Bristol, UK

    Gibran Hemani, Susan Ring, Beate St Pourcain, David M. Evans, George McMahon, Lavinia Paternoster, Susan M. Ring, Thorkild I. A. Sørensen, Nicholas J. Timpson & George Davey Smith

  101. Vrije Universiteit Amsterdam, Amsterdam, the Netherlands

    Rick Jansen, Michel G. Nivard, Brenda Penninx, Jenny van Dongen & Brenda Penninx

  102. Department of Clinical Physiology, Tampere University Hospital and Faculty of Medicine and Life Sciences, University of Tampere, Tampere, Finland

    Mika Kähönen & Mika Kähönen

  103. University of Helsinki, Helsinki, Finland

    Johannes Kettunen & Samuli Ripatti

  104. Genetics and Genomic Science Department, Icahn School of Medicine at Mount Sinai, New York, NY, USA

    Yungil Kim

  105. Institut für Medizinische Informatik, Statistik und Epidemiologie, Leipzig Research Centre for Civilization Diseases (LIFE), Universität Leipzig, Leipzig, Germany

    Holger Kirsten, Markus Loeffler & Markus Scholz

  106. IFB Adiposity Diseases, Universität Leipzig, Leipzig, Germany

    Peter Kovacs

  107. Interdisciplinary Center for Clinical Research, Faculty of Medicine, Universität Leipzig, Leipzig, Germany

    Knut Krohn

  108. Lausanne University Hospital, Lausanne, Switzerland

    Zoltan Kutalik, Sina Rüeger & Zoltan Kutalik

  109. Department of Clinical Chemistry, Fimlab Laboratories and Faculty of Medicine and Life Sciences, University of Tampere, Tampere, Finland

    Terho Lehtimäki, Ilkka Seppälä & Terho Lehtimäki

  110. Institute of Genetic Epidemiology, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, Germany

    Martina Müller-Nurasyid, Katharina Schramm & Konstantin Strauch

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

    Matthias Nauck

  112. National Institute for Health and Welfare, University of Helsinki, Helsinki, Finland

    Markus Perola, Niina Eklund, Seppo Koskinen, Tomi Mäki-Opas, Veikko Salomaa, Jaakko Kaprio & Markus Perola

  113. Garvan Institute of Medical Research, Garvan-Weizmann Centre for Cellular Genomics, Sydney, New South Wales, Australia

    Joseph Powell

  114. Institute of Human Genetics, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, Germany

    Holger Prokisch, Peter Lichtner & Thomas Meitinger

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

    Olli Raitakari & Olli Raitakari

  116. Population Health Sciences, Bristol Medical School, University of Bristol, Bristol, UK

    Susan Ring

  117. Department of Medicine, Universität Leipzig, Leipzig, Germany

    Michael Stumvoll & Anke Tönjes

  118. Radboud University Medical Center, Radboud University, Nijmegen, the Netherlands

    Peter-Bram t Hoen, Tessel E. Galesloot & Lambertus A. L. M. Kiemeney

  119. Institute for Community Medicine, University Medicine Greifswald, Greifswald, Germany

    Alexander Teumer, Alexander Teumer, Sebastian E. Baumeister, Henry Völzke & Wolfgang Hoffmann

  120. Institute for Laboratory Medicine, Leipzig Research Center for Civilization Diseases (LIFE), Universität Leipzig, Leipzig, Germany

    Joachim Thiery

  121. Interfaculty Institute for Genetics and Functional Genomics, University Medicine Greifswald, Greifswald, Germany

    Uwe Völker, Georg Homuth & Uwe Völker

  122. Icelandic Heart Association, Kopavogur, Iceland

    Valur Emilsson, Albert V. Smith & Vilmundur Gudnason

  123. Faculty of Pharmaceutical Sciences, University of Iceland, Reykjavík, Iceland

    Valur Emilsson

  124. Department of Government, Uppsala University, Uppsala, Sweden

    Sven Oskarsson & Karl-Oskar Lindgren

  125. New York Genome Center, New York, NY, USA

    Joseph K. Pickrell

  126. Copenhagen Prospective Studies on Asthma in Childhood (COPSAC), Herlev and Gentofte Hospital, University of Copenhagen, Copenhagen, Denmark

    Tarunveer S. Ahluwalia, Klaus Bønnelykke, Johannes Waage & Hans Bisgaard

  127. Steno Diabetes Center, Gentofte, Denmark

    Tarunveer S. Ahluwalia & Johannes Waage

  128. Department of Obstetrics and Gynecology, Institute of Clinical Sciences, Sahlgrenska Academy, Gothenburg, Sweden

    Jonas Bacelis & Bo Jacobsson

  129. Research Unit of Molecular Epidemiology, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, Germany

    Clemens Baumbach & Christian Gieger

  130. Institute of Epidemiology II, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, Germany

    Clemens Baumbach & Christa Meisinger

  131. deCODE Genetics/Amgen Inc., Reykjavik, Iceland

    Gyda Bjornsdottir, Augustine Kong, Gudmar Thorleifsson, Bjarni V. Halldórsson, Kari Stefansson & Unnur Thorsteinsdottir

  132. Department of Cell Biology, Erasmus Medical Center, Rotterdam, the Netherlands

    Johannes H. Brandsma & Raymond A. Poot

  133. Istituto di Ricerca Genetica e Biomedica UOS di Sassari, National Research Council of Italy, Sassari, Italy

    Maria Pina Concas, Bjarni Gunnarsson, Simona Vaccargiu & Mario Pirastu

  134. Psychology, University of Illinois, Champaign, IL, USA

    Jaime Derringer

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

    Giorgia Girotto, Dragana Vuckovic, Ilaria Gandin, Paolo Gasparini & Nicola Pirastu

  136. Department of Public Health, University of Helsinki, Helsinki, Finland

    Richa Gupta, Antti Latvala, Anu Loukola & Jaakko Kaprio

  137. Department of Cardiovascular Sciences, University of Leicester, Leicester, UK

    Leanne M. Hall, Christopher P. Nelson & Nilesh J. Samani

  138. NIHR Leicester Cardiovascular Biomedical Research Unit, Glenfield Hospital, Leicester, UK

    Leanne M. Hall, Christopher P. Nelson & Nilesh J. Samani

  139. Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, UK

    Sarah E. Harris, Gail Davies, David C. M. Liewald, Riccardo E. Marioni & Ian J. Deary

  140. Centre for Genomic and Experimental Medicine, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, UK

    Sarah E. Harris & David J. Porteous

  141. Department of Neurology, General Hospital and Medical University Graz, Graz, Austria

    Edith Hofer, Katja E. Petrovic, Helena Schmidt & Reinhold Schmidt

  142. Institute for Medical Informatics, Statistics and Documentation, General Hospital and Medical University Graz, Graz, Austria

    Edith Hofer

  143. Oxford Centre for Diabetes, Endocrinology & Metabolism, University of Oxford, Oxford, UK

    Momoko Horikoshi

  144. Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, UK

    Momoko Horikoshi

  145. Institute of Behavioural Sciences, University of Helsinki, Helsinki, Finland

    Kadri Kaasik, Jari Lahti, Liisa Keltigangas-Järvinen & Katri Räikkönen

  146. Nutrition and Dietetics, Health Science and Education, Harokopio University, Athens, Greece

    Ioanna P. Kalafati & George V. Dedoussis

  147. Folkhälsan Research Centre, Helsingfors, Finland

    Jari Lahti, Katri Räikkönen & Johan G. Eriksson

  148. Institute for Computing and Information Sciences, Radboud University Nijmegen, Nijmegen, the Netherlands

    Christiaan de Leeuw

  149. Quantitative Genetics, QIMR Berghofer Medical Research Institute, Brisbane, Queensland, Australia

    Penelope A. Lind & Sarah E. Medland

  150. Department of Twin Research and Genetic Epidemiology, King’s College London, London, UK

    Massimo Mangino, Lydia Quaye, Cristina Venturini & Tim D. Spector

  151. NIHR Biomedical Research Centre, Guy’s and St. Thomas’ Foundation Trust, London, UK

    Massimo Mangino & Cristina Venturini

  152. Department of Epidemiology, University Medical Center Groningen, University of Groningen, Groningen, the Netherlands

    Peter J. van der Most, Behrooz Z. Alizadeh & Judith M. Vonk

  153. Public Health Stream, Hunter Medical Research Institute, New Lambton, New South Wales, Australia

    Christopher Oldmeadow, Elizabeth G. Holliday & John R. Attia

  154. Faculty of Health and Medicine, University of Newcastle, Newcastle, New South Wales, Australia

    Christopher Oldmeadow, Elizabeth G. Holliday, Rodney J. Scott & John R. Attia

  155. Centre for Integrated Genomic Medical Research, Institute of Population Health, University of Manchester, Manchester, UK

    Antony Payton & William E. R. Ollier

  156. Human Communication and Deafness, School of Psychological Sciences, University of Manchester, Manchester, UK

    Antony Payton

  157. Department of Psychiatry, VU University Medical Center, Amsterdam, the Netherlands

    Wouter J. Peyrot & Yusplitri Milaneschi

  158. GGZ inGeest, Amsterdam, the Netherlands

    Wouter J. Peyrot & Yusplitri Milaneschi

  159. Laboratory of Genetics, National Institute on Aging, Baltimore, MD, USA

    Yong Qian, Jun Ding & David Schlessinger

  160. Department of Medical Genetics, University of Lausanne, Lausanne, Switzerland

    Rico Rueedi

  161. Swiss Institute of Bioinformatics, Lausanne, Switzerland

    Rico Rueedi

  162. Department Of Health Sciences, University of Milan, Milano, Italy

    Erika Salvi & Daniele Cusi

  163. Institute for Medical Informatics, Biometry and Epidemiology, University Hospital of Essen, Essen, Germany

    Börge Schmidt, Lewin Eisele & Karl-Heinz Jöckel

  164. Division of Cancer Epidemiology and Genetics, National Cancer Institute, Bethesda, MD, USA

    Jianxin Shi

  165. Faculty of Medicine, University of Iceland, Reykjavik, Iceland

    Albert V. Smith, Vilmundur Gudnason, Kari Stefansson & Unnur Thorsteinsdottir

  166. School of Oral and Dental Sciences, University of Bristol, Bristol, UK

    Beate St Pourcain

  167. Department of Cardiology, University Medical Center Groningen, University of Groningen, Groningen, the Netherlands

    Niek Verweij, Klaus Berger & Pim van der Harst

  168. Institute of Epidemiology and Social Medicine, University of Münster, Münster, Germany

    Juergen Wellmann

  169. Rush Alzheimer’s Disease Center, Rush University Medical Center, Chicago, IL, USA

    Jingyun Yang, Patricia A. Boyle & David A. Bennett

  170. Department of Neurological Sciences, Rush University Medical Center, Chicago, IL, USA

    Jingyun Yang & David A. Bennett

  171. Department of Epidemiology, University of Michigan, Ann Arbor, MI, USA

    Wei Zhao, Jennifer A. Smith, Erin B. Ware & Sharon L. R. Kardia

  172. Department of Gastroenterology and Hepatology, University Medical Center Groningen, University of Groningen, Groningen, the Netherlands

    Behrooz Z. Alizadeh

  173. Institute of Epidemiology and Preventive Medicine, University of Regensburg, Regensburg, Germany

    Sebastian E. Baumeister

  174. Institute of Molecular Genetics, National Research Council of Italy, Pavia, Italy

    Ginevra Biino

  175. Department of Behavioral Sciences, Rush University Medical Center, Chicago, IL, USA

    Patricia A. Boyle

  176. Warwick Medical School, University of Warwick, Coventry, UK

    Francesco P. Cappuccio

  177. Department of Psychology, University of Edinburgh, Edinburgh, UK

    Gail Davies, David C. M. Liewald & Ian J. Deary

  178. Saïd Business School, University of Oxford, Oxford, UK

    Jan-Emmanuel De Neve

  179. William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London, UK

    Panos Deloukas & Stavroula Kanoni

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

    Panos Deloukas

  181. The Berlin Aging Study II Research Group on Geriatrics, Charité–Universitätsmedizin Berlin, Berlin, Germany

    Ilja Demuth & Elisabeth Steinhagen-Thiessen

  182. Institute of Medical and Human Genetics, Charité–Universitätsmedizin Berlin, Berlin, Germany

    Ilja Demuth

  183. University of Queensland Diamantina Institute, Translational Research Institute, University of Queensland, Brisbane, Queensland, Australia

    David M. Evans

  184. Survey Research Center, Institute for Social Research, University of Michigan, Ann Arbor, MI, USA

    Jessica D. Faul & David R. Weir

  185. Division of Statistical Genomics, Department of Genetics, Washington University School of Medicine, St. Louis, MO, USA

    Mary F. Feitosa, Aldi T. Kraja, Ingrid B Borecki & Michael A. Province

  186. Institute of Human Genetics, University of Bonn, Bonn, Germany

    Andreas J. Forstner

  187. Department of Genomics, Life and Brain Center, University of Bonn, Bonn, Germany

    Andreas J. Forstner

  188. Institute of Biomedical and Neural Engineering, School of Science and Engineering, Reykjavik University, Reykjavik, Iceland

    Bjarni V. Halldórsson

  189. Laboratory of Epidemiology & Population Science, National Institute on Aging, National Institutes of Health, Bethesda, MD, USA

    Tamara B. Harris

  190. Department of Psychiatry, Washington University School of Medicine, St. Louis, MO, USA

    Andrew C. Heath & Pamela A. Madden

  191. Division of Applied Health Sciences, University of Aberdeen, Aberdeen, UK

    Lynne J. Hocking

  192. Manchester Medical School, University of Manchester, Manchester, UK

    Michael A. Horan

  193. Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA

    Philip L. de Jager & Aarno Palotie

  194. Program in Translational NeuroPsychiatric Genomics, Departments of Neurology and Psychiatry, Brigham and Women’s Hospital, Boston, MA, USA

    Philip L. de Jager

  195. Harvard Medical School, Boston, Massachusetts, USA

    Philip L. de Jager

  196. Department of Genes and Environment, Norwegian Institute of Public Health, Oslo, Norway

    Astanand Jugessur, Ronny Myhre & Bo Jacobsson

  197. Department of Genomics of Common Disease, Imperial College London, London, UK

    Marika A. Kaakinen

  198. Public Health, Medical School, University of Split, Split, Croatia

    Ivana Kolcic

  199. Neuroepidemiology Section, National Institute on Aging, National Institutes of Health, Bethesda, MD, USA

    Lenore J. Launer

  200. Department of Psychiatry and Behavioral Sciences, Stanford University, Stanford, CA, USA

    Douglas F. Levinson

  201. Medical Genetics Section, Centre for Genomic and Experimental Medicine, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, UK

    Riccardo E. Marioni

  202. Department of Internal Medicine, Lausanne University Hospital (CHUV), Lausanne, Switzerland

    Pedro Marques-Vidal & Peter Vollenweider

  203. Molecular Epidemiology, QIMR Berghofer Medical Research Institute, Brisbane, Queensland, Australia

    Dale R. Nyholt

  204. Institute of Health and Biomedical Innovation, Queensland Institute of Technology, Brisbane, Queensland, Australia

    Dale R. Nyholt

  205. Analytic and Translational Genetics Unit, Department of Medicine, Massachusetts General Hospital, Boston, MA, USA

    Aarno Palotie

  206. Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland

    Aarno Palotie, Antti-Pekka Sarin & Jaakko Kaprio

  207. Department of Neurology, Massachusetts General Hospital, Boston, MA, USA

    Aarno Palotie

  208. Medical Genetics, Institute for Maternal and Child Health IRCCS ‘Burlo Garofolo’, Trieste, Italy

    Antonietta Robino, Sheila Ulivi & Paolo Gasparini

  209. Social Impact, Arlington, VA, USA

    Olga Rostapshova & Diego Vozzi

  210. Department of Economics, University of Minnesota Twin Cities, Minneapolis, MN, USA

    Aldo Rustichini

  211. Department of Psychiatry and Behavioral Sciences, NorthShore University HealthSystem, Evanston, IL, USA

    Alan R. Sanders & Pablo V. Gejman

  212. Public Health Genomics Unit, National Institute for Health and Welfare, Helsinki, Finland

    Antti-Pekka Sarin

  213. Research Unit for Genetic Epidemiology, Institute of Molecular Biology and Biochemistry, Center of Molecular Medicine, General Hospital and Medical University, Graz, Austria

    Helena Schmidt

  214. Information Based Medicine Stream, Hunter Medical Research Institute, New Lambton, New South Wales, Australia

    Rodney J. Scott

  215. Medical Research Institute, University of Dundee, Dundee, UK

    Blair H. Smith

  216. Research Unit Hypertension and Cardiovascular Epidemiology, Department of Cardiovascular Science, University of Leuven, Leuven, Belgium

    Jan A. Staessen

  217. R&D VitaK Group, Maastricht University, Maastricht, the Netherlands

    Jan A. Staessen

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

    Konstantin Strauch

  219. Department of Geriatrics, Florida State University College of Medicine, Tallahassee, FL, USA

    Antonio Terracciano

  220. Department of Health Sciences and Genetics, University of Leicester, Leicester, UK

    Martin D. Tobin

  221. Research Center for Group Dynamics, Institute for Social Research, University of Michigan, Ann Arbor, MI, USA

    Erin B. Ware

  222. Department of Health Sciences, Community and Occupational Medicine, University Medical Center Groningen, University of Groningen, Groningen, the Netherlands

    Ute Bültmann

  223. Department of Psychology, Union College, Schenectady, NY, USA

    Christopher F. Chabris

  224. Istituto di Ricerca Genetica e Biomedica (IRGB), Consiglio Nazionale delle Ricerche, Cittadella Universitaria di Monserrato, Cagliari, Italy

    Francesco Cucca

  225. Institute of Biomedical Technologies, Italian National Research Council, Milan, Italy

    Daniele Cusi

  226. Department of General Practice and Primary Health Care, University of Helsinki, Helsinki, Finland

    Johan G. Eriksson

  227. Departments of Human Genetics and Psychiatry, Donders Centre for Neuroscience, Nijmegen, the Netherlands

    Barbara Franke

  228. Experimental Genetics Division, Sidra, Doha, Qatar

    Paolo Gasparini

  229. Department of Psychiatry and Psychotherapy, University Medicine Greifswald, Greifswald, Germany

    Hans-Jörgen Grabe

  230. Department of Psychiatry and Psychotherapy, HELIOS-Hospital Stralsund, Stralsund, Germany

    Hans-Jörgen Grabe

  231. Durrer Center for Cardiogenetic Research, ICIN-Netherlands Heart Institute, Utrecht, the Netherlands

    Pim van der Harst

  232. Generation Scotland, Centre for Genomics and Experimental Medicine, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, UK

    Caroline Hayward

  233. Centre for Population Health Research, School of Health Sciences and Sansom Institute, University of South Australia, Adelaide, South Australia, Australia

    Elina Hyppönen

  234. South Australian Health and Medical Research Institute, Adelaide, South Australia, Australia

    Elina Hyppönen

  235. Population, Policy and Practice, UCL Institute of Child Health, London, UK

    Elina Hyppönen & Christine Power

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

    Marjo-Riitta Järvelin

  237. Center for Life Course Epidemiology, Faculty of Medicine, University of Oulu, Oulu, Finland

    Marjo-Riitta Järvelin

  238. Unit of Primary Care, Oulu University Hospital, Oulu, Finland

    Marjo-Riitta Järvelin

  239. Biocenter Oulu, University of Oulu, Oulu, Finland

    Marjo-Riitta Järvelin

  240. Economics, NYU Shanghai, Pudong, China

    Steven F. Lehrer

  241. Policy Studies, Queen’s University, Kingston, Ontario, Canada

    Steven F. Lehrer

  242. Genetic Epidemiology, QIMR Berghofer Medical Research Institute, Brisbane, Queensland, Australia

    Nicholas G. Martin

  243. Centre for Clinical and Cognitive Neuroscience, Institute of Brain Behaviour and Mental Health, Salford Royal Hospital, Manchester, UK

    Neil Pendleton

  244. Manchester Institute for Collaborative Research in Ageing, University of Manchester, Manchester, UK

    Neil Pendleton

  245. Faculty of Medicine, University of Split, Split, Croatia

    Ozren Polasek

  246. Institute of Preventive Medicine, Bispebjerg and Frederiksberg Hospitals, Frederiksberg, Denmark

    Thorkild I. A. Sørensen

  247. Department of Sociology, New York University, New York, NY, USA

    Dalton C. Conley

  248. School of Medicine, New York University, New York, NY, USA

    Dalton C. Conley

Authors
  1. Richard Karlsson Linnér
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Contributions

J.P.B., P.D.K., and D.J.B. designed and oversaw the study. J.P.B. closely supervised the analyses and led the writing of the manuscript. R.K.L. was the lead analyst, responsible for quality control, the meta-analyses, summarizing the overlap across the results of the various GWAS, and the SNP heritability analyses. E.K. conducted the population stratification, replication, and proxy-phenotype analyses. R.W. led the genetic correlation analyses, and M.A.F. contributed to those analyses. P.B. led the polygenic score prediction analyses, and R.K.L., E.K., R.W., A.A., R.d.V., M.A.F., and M.G.N. contributed to those analyses. P.B. and C.L.Z. conducted the MTAG analyses. The bioinformatics analyses were led by S.F.W.M. Analysts who assisted S.F.W.M. include J.G. and M.T. (SMR), A.R.H. (MAGMA), G.A.M. (Gene Network), and P.N.T. (DEPICT). M.L. conducted the review of the literature attempting to link risk tolerance to biological pathways. R.K.L. prepared the majority of figures; E.K. and S.P.T. also prepared some figures. A.O., C.A.R., and S.P.T. helped with several additional analyses. D.C., J.G., and J.J.L. provided helpful advice and feedback on various aspects of the study design. All authors contributed to and critically reviewed the manuscript. P.B., R.K.L., E.K., S.F.W.M., and R.W. made especially major contributions to the writing and editing; these authors contributed equally, and lead authors 2–5 are listed aphabetically in the author list.

Corresponding authors

Correspondence to Richard Karlsson Linnér or Jonathan P. Beauchamp.

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Competing interests

A. Auton, P.F., D.A.H., and A.K. are employees of 23andMe. R.C.K., in the past three years, received support for his epidemiological studies from Sanofi Aventis; was a consultant for Johnson & Johnson Wellness and Prevention, Sage Pharmaceuticals, Shire, and Takeda; and served on an advisory board for the Johnson & Johnson Services Inc. Lake Nona Life Project. R.C.K. is a co-owner of DataStat Inc., a market research firm that carries out healthcare research. J.M. is a principal in BEAM Diagnostics Inc. The authors declare no other competing interests.

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Karlsson Linnér, R., Biroli, P., Kong, E. et al. Genome-wide association analyses of risk tolerance and risky behaviors in over 1 million individuals identify hundreds of loci and shared genetic influences. Nat Genet 51, 245–257 (2019). https://doi.org/10.1038/s41588-018-0309-3

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