About the Authors

Ruth McQuillan

Affiliation Centre for Population Health Sciences, University of Edinburgh, Edinburgh, Scotland, United Kingdom

Niina Eklund

Affiliations Institute for Molecular Medicine Finland (FIMM), Helsinki, Finland, Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, Finland

Nicola Pirastu

Affiliation Institute for Maternal and Child Health, IRCCS “Burlo Garofolo,” University of Trieste, Trieste, Italy

Maris Kuningas

Affiliation Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands

Brian P. McEvoy

Affiliation Queensland Institute of Medical Research, Brisbane, Queensland, Australia

Tõnu Esko

Affiliations Estonian Genome Center, University of Tartu, Tartu, Estonia, Institute of Molecular and Cell Biology, University of Tartu, Tartu, Estonia

Tanguy Corre

Affiliation Division of Genetics and Cell Biology, San Raffaele Research Institute, Milano, Italy

Gail Davies

Affiliation Department of Psychology, University of Edinburgh, Edinburgh, Scotland, United Kingdom

Marika Kaakinen

Affiliations Biocenter Oulu, University of Oulu, Oulu, Finland, Institute of Health Sciences, University of Oulu, Oulu, Finland

Leo-Pekka Lyytikäinen

Affiliations Department of Clinical Chemistry, Tampere University Hospital, Tampere, Finland, Department of Clinical Chemistry, University of Tampere School of Medicine, Tampere, Finland

Kati Kristiansson

Affiliations Institute for Molecular Medicine Finland (FIMM), Helsinki, Finland, Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, Finland

Aki S. Havulinna

Affiliation Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, Finland

Martin Gögele

Affiliation Centre for Biomedicine, European Academy Bozen/Bolzano (EURAC), Bolzano, Italy

Veronique Vitart

Affiliation MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, Scotland, United Kingdom

Albert Tenesa

Affiliations MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, Scotland, United Kingdom, The Roslin Institute, University of Edinburgh, Easter Bush, Midlothian, Scotland, United Kingdom

Yurii Aulchenko

Affiliation Genetic Epidemiology Unit, Department of Epidemiology, Erasmus MC University Medical Center, Rotterdam, The Netherlands

Caroline Hayward

Affiliation MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, Scotland, United Kingdom

Åsa Johansson

Affiliation Uppsala Clinical Research, Uppsala University, Uppsala, Sweden

Mladen Boban

Affiliation Department of Pharmacology, Faculty of Medicine, University of Split, Split, Croatia

Sheila Ulivi

Affiliation Institute for Maternal and Child Health, IRCCS “Burlo Garofolo,” Trieste, Italy

Antonietta Robino

Affiliation Institute for Maternal and Child Health, IRCCS “Burlo Garofolo,” University of Trieste, Trieste, Italy

Vesna Boraska

Affiliation Department of Biology, Faculty of Medicine, University of Split, Split, Croatia

Wilmar Igl

Affiliation Department of Immunology, Genetics and Pathology, SciLifeLab Uppsala, Rudbeck Laboratory, Uppsala University, Uppsala, Sweden

Sarah H. Wild

Affiliation Centre for Population Health Sciences, University of Edinburgh, Edinburgh, Scotland, United Kingdom

Lina Zgaga

Affiliations Centre for Population Health Sciences, University of Edinburgh, Edinburgh, Scotland, United Kingdom, Andrija Stampar School of Public Health, Medical School, University of Zagreb, Zagreb, Croatia

Najaf Amin

Affiliation Genetic Epidemiology Unit, Department of Epidemiology, Erasmus MC University Medical Center, Rotterdam, The Netherlands

Evropi Theodoratou

Affiliation Centre for Population Health Sciences, University of Edinburgh, Edinburgh, Scotland, United Kingdom

Ozren Polašek

Affiliations Department of Public Health, Faculty of Medicine, University of Split, Split, Croatia, Centre for Global Health, University of Split, Split, Croatia

Giorgia Girotto

Affiliation Institute for Maternal and Child Health, IRCCS “Burlo Garofolo,” University of Trieste, Trieste, Italy

Lorna M. Lopez

Affiliations Department of Psychology, University of Edinburgh, Edinburgh, Scotland, United Kingdom, Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, Scotland, United Kingdom

Cinzia Sala

Affiliation Division of Genetics and Cell Biology, San Raffaele Research Institute, Milano, Italy

Jari Lahti

Affiliation Institute of Behavioural Sciences, University of Helsinki, Helsinki, Finland

Tiina Laatikainen

Affiliation Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, Finland

Inga Prokopenko

Affiliations Oxford Centre for Diabetes, Endocrinology, and Metabolism, University of Oxford, Churchill Hospital, Headington, Oxford, United Kingdom, Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom

Mart Kals

Affiliation Estonian Genome Center, University of Tartu, Tartu, Estonia

Jorma Viikari

Affiliations Department of Medicine, Turku University Hospital, Turku, Finland, Department of Medicine, University of Turku, Turku, Finland

Jian Yang

Affiliation Queensland Institute of Medical Research, Brisbane, Queensland, Australia

Anneli Pouta

Affiliation National Institute for Health and Welfare, Oulu, Finland

Karol Estrada

Affiliations Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands, Department of Internal Medicine, Erasmus University Medical Center, Rotterdam, The Netherlands, Netherlands Genomics Initiative (NGI)–sponsored Netherlands Consortium for Healthy Aging (NCHA), Leiden, The Netherlands

Albert Hofman

Affiliations Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands, Netherlands Genomics Initiative (NGI)–sponsored Netherlands Consortium for Healthy Aging (NCHA), Leiden, The Netherlands

Nelson Freimer

Affiliations Brain Research Institute, University of California Los Angeles, Los Angeles, California, United States of America, UCLA Center for Neurobehavioral Genetics, University of California Los Angeles, Los Angeles, California, United States of America, Semel Institute for Neuroscience and Human Behavior, University of California Los Angeles, Los Angeles, California, United States of America

Nicholas G. Martin

Affiliation Queensland Institute of Medical Research, Brisbane, Queensland, Australia

Mika Kähönen

Affiliations Department of Clinical Physiology, Tampere University Hospital, Tampere, Finland, Department of Clinical Physiology, University of Tampere School of Medicine, Tampere, Finland

Lili Milani

Affiliation Estonian Genome Center, University of Tartu, Tartu, Estonia

Markku Heliövaara

Affiliation Department of Health and Functional Capacity, National Public Health Institute–Helsinki and Turku, Turku, Finland

Erkki Vartiainen

Affiliation Division of Welfare and Health Promotion, National Institute for Health and Welfare, Helsinki, Finland

Katri Räikkönen

Affiliation Institute of Behavioural Sciences, University of Helsinki, Helsinki, Finland

Corrado Masciullo

Affiliation Division of Genetics and Cell Biology, San Raffaele Research Institute, Milano, Italy

John M. Starr

Affiliations Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, Scotland, United Kingdom, Geriatric Medicine Unit, University of Edinburgh, Royal Victoria Hospital, Edinburgh, Scotland, United Kingdom

Andrew A. Hicks

Affiliation Centre for Biomedicine, European Academy Bozen/Bolzano (EURAC), Bolzano, Italy

Laura Esposito

Affiliation CBM scrl – Genomics, Area Science Park, Basovizza, Trieste, Italy

Ivana Kolčić

Affiliations Department of Public Health, Faculty of Medicine, University of Split, Split, Croatia, Centre for Global Health, University of Split, Split, Croatia

Susan M. Farrington

Affiliation Colon Cancer Genetics Group, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, Scotland, United Kingdom

Ben Oostra

Affiliation Genetic Epidemiology Unit, Department of Clinical Genetics, Erasmus MC University Medical Center, Rotterdam, The Netherlands

Tatijana Zemunik

Affiliation Department of Biology, Faculty of Medicine, University of Split, Split, Croatia

Harry Campbell

Affiliation Centre for Population Health Sciences, University of Edinburgh, Edinburgh, Scotland, United Kingdom

Mirna Kirin

Affiliation Centre for Population Health Sciences, University of Edinburgh, Edinburgh, Scotland, United Kingdom

Marina Pehlic

Affiliation Department of Biology, Faculty of Medicine, University of Split, Split, Croatia

Flavio Faletra

Affiliation Institute for Maternal and Child Health, IRCCS “Burlo Garofolo,” Trieste, Italy

David Porteous

Affiliations Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, Scotland, United Kingdom, Medical Genetics Section, The University of Edinburgh Molecular Medicine Centre, Institute of Genetics and Molecular Medicine, Western General Hospital, Edinburgh, Scotland, United Kingdom

Giorgio Pistis

Affiliation Division of Genetics and Cell Biology, San Raffaele Research Institute, Milano, Italy

Elisabeth Widén

Affiliation Institute for Molecular Medicine Finland (FIMM), Helsinki, Finland

Veikko Salomaa

Affiliation Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, Finland

Seppo Koskinen

Affiliation Department of Health and Functional Capacity, National Public Health Institute, Helsinki, Finland

Krista Fischer

Affiliation Estonian Genome Center, University of Tartu, Tartu, Estonia

Terho Lehtimäki

Affiliations Department of Clinical Chemistry, Tampere University Hospital, Tampere, Finland, Department of Clinical Chemistry, University of Tampere School of Medicine, Tampere, Finland

Andrew Heath

Affiliation Department of Psychiatry, Washington University St. Louis, Missouri, United States of America

Mark I. McCarthy

Affiliations Oxford Centre for Diabetes, Endocrinology, and Metabolism, University of Oxford, Churchill Hospital, Headington, Oxford, United Kingdom, Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom, Oxford NIHR Biomedical Research Centre, Churchill Hospital, Headington, Oxford, United Kingdom

Fernando Rivadeneira

Affiliations Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands, Department of Internal Medicine, Erasmus University Medical Center, Rotterdam, The Netherlands, Netherlands Genomics Initiative (NGI)–sponsored Netherlands Consortium for Healthy Aging (NCHA), Leiden, The Netherlands

Grant W. Montgomery

Affiliation Queensland Institute of Medical Research, Brisbane, Queensland, Australia

Henning Tiemeier

Affiliations Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands, Netherlands Genomics Initiative (NGI)–sponsored Netherlands Consortium for Healthy Aging (NCHA), Leiden, The Netherlands, Department of Psychiatry, Erasmus Medical Center, Rotterdam, The Netherlands

Anna-Liisa Hartikainen

Affiliation Institute of Clinical Medicine/Obstetrics and Gynecology, University of Oulu, Oulu, Finland

Pamela A. F. Madden

Affiliation Department of Psychiatry, Washington University St. Louis, Missouri, United States of America

Pio d'Adamo

Affiliation Institute for Maternal and Child Health, IRCCS “Burlo Garofolo,” University of Trieste, Trieste, Italy

Nicholas D. Hastie

Affiliation MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, Scotland, United Kingdom

Ulf Gyllensten

Affiliation Department of Immunology, Genetics and Pathology, SciLifeLab Uppsala, Rudbeck Laboratory, Uppsala University, Uppsala, Sweden

Alan F. Wright

Affiliation MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, Scotland, United Kingdom

Cornelia M. van Duijn

Affiliations Genetic Epidemiology Unit, Department of Epidemiology, Erasmus MC University Medical Center, Rotterdam, The Netherlands, Netherlands Genomics Initiative (NGI)–sponsored Netherlands Consortium for Healthy Aging (NCHA), Leiden, The Netherlands

Malcolm Dunlop

Affiliation Colon Cancer Genetics Group, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, Scotland, United Kingdom

Igor Rudan

Affiliation Centre for Population Health Sciences, University of Edinburgh, Edinburgh, Scotland, United Kingdom

Paolo Gasparini

Affiliation Institute for Maternal and Child Health, IRCCS “Burlo Garofolo,” University of Trieste, Trieste, Italy

Peter P. Pramstaller

Affiliations Centre for Biomedicine, European Academy Bozen/Bolzano (EURAC), Bolzano, Italy, Department of Neurology, General Central Hospital, Bolzano, Italy, Department of Neurology, University of Lübeck, Lübeck, Germany

Ian J. Deary

Affiliations Department of Psychology, University of Edinburgh, Edinburgh, Scotland, United Kingdom, Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, Scotland, United Kingdom

Daniela Toniolo

Affiliations Division of Genetics and Cell Biology, San Raffaele Research Institute, Milano, Italy, Institute of Molecular Genetics–CNR, Pavia, Italy

Johan G. Eriksson

Affiliations Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, Finland, Department of General Practice and Primary Health Care, University of Helsinki, Helsinki, Finland, Unit of General Practice, Helsinki University Central Hospital, Helsinki, Finland, Folkhälsan Research Center, Helsinki, Finland

Antti Jula

Affiliation Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, Finland

Olli T. Raitakari

Affiliations Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku, Finland, Department of Clinical Physiology, Turku University Hospital, Turku, Finland

Andres Metspalu

Affiliations Estonian Genome Center, University of Tartu, Tartu, Estonia, Institute of Molecular and Cell Biology, University of Tartu, Tartu, Estonia

Markus Perola

Affiliations Institute for Molecular Medicine Finland (FIMM), Helsinki, Finland, Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, Finland, Estonian Genome Center, University of Tartu, Tartu, Estonia

Marjo-Riitta Järvelin

Affiliations Biocenter Oulu, University of Oulu, Oulu, Finland, Institute of Health Sciences, University of Oulu, Oulu, Finland, National Institute for Health and Welfare, Oulu, Finland, Department of Epidemiology and Biostatistics, School of Public Health, Imperial College London, United Kingdom

André Uitterlinden

Affiliations Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands, Department of Internal Medicine, Erasmus University Medical Center, Rotterdam, The Netherlands, Netherlands Genomics Initiative (NGI)–sponsored Netherlands Consortium for Healthy Aging (NCHA), Leiden, The Netherlands

Peter M. Visscher

Affiliation Queensland Institute of Medical Research, Brisbane, Queensland, Australia

James F. Wilson

jim.wilson@ed.ac.uk

Affiliation Centre for Population Health Sciences, University of Edinburgh, Edinburgh, Scotland, United Kingdom

on behalf of the ROHgen Consortium

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Analyzed the data: R McQuillan, N Eklund, N Pirastu, BP McEvoy, M Kuningas, T Esko, C Torre, G Davies, L-P Lyytikäinen, K Kristiansson, M Kirin, M Gögele, V Vitart, Y Aulchenko, C Hayward, Å Johansson, W Igl, N Amin, LM Lopez, K Estrada, C Masciullo, G Pistis, T Lhetimäki, J Yang, F Rivadeneira, PM Visscher, OT Raitakari, T Corre, A Tenesa, L Milani, M Kals, K Fischer. Wrote the paper: R McQuillan, JF Wilson. Conceived and designed the experiments: D Porteous, K Fischer, A Metspalu. Established and managed the population resources: AS Havulinna, C Hayward, Å Johansson, M Boban, S Ulivi, V Boraska, W Igl, SH Wild, L Zgaga, E Theodoratou, O Polašek, J Lahti, T Laatikainen, I Prokopenko, J Viikari, AA Hicks, M Kähönen, E Vartiainen, K Räikkönen, JM Starr, A Heath, I Kolčić, SM Farrington, B Oostra, T Zemunik, H Campbell, M Pehlic, E Widén, V Salomaa, T Lehtimäki, H Tiemeier, P d'Adamo, ND Hastie, U Gyllensten, AF Wright, CM van Duijn, M Dunlop, I Rudan, P Gasparini, PP Pramstaller, IJ Deary, D Toniolo, JG Eriksson, M Perola, A Jula, M-R Järvelin, NG Martin, GW Montgomery, A Hofman, PAF Madden, PM Visscher, OT Raitakari, A Uitterlinden, JF Wilson, M Heliövaara, S Koskinen, A Metspalu. Collected the data: AS Havulinna, M Gögele, V Vitart, C Hayward, Å Johansson, M Boban, S Ulivi, A Robino, V Boraska, W Igl, SH Wild, L Zgaga, E Theodoratou, O Polašek, G Girotto, C Sala, J Lahti, I Prokopenko, J Viikari, AA Hicks, M Kähönen, K Räikkönen, JM Starr, A Heath, L Esposito, I Kolčić, B Oostra, T Zemunik, M Pehlic, F Faletra, E Widén, V Salomaa, T Lhetimäki, F Rivadeneira, H Tiemeier, U Gyllensten, CM van Duijn, I Rudan, PP Pramstaller, NG Martin, GW Montgomery, A Hofman, PAF Madden, IJ Deary, D Toniolo, JG Eriksson, M Perola, A Jula, OT Raitakari, A Uitterlinden, M Kaakinen, A Pouta, N Freimer, MI McCarthy, A-L Hartikainen, A Metspalu, JF Wilson. Conducted meta-analysis: R McQuillan.