About the Authors

Jens Baumert

Contributed equally to this work with: Jens Baumert, Jie Huang, Barbara McKnight, Maria Sabater-Lleal, Maristella Steri

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

Jie Huang

Contributed equally to this work with: Jens Baumert, Jie Huang, Barbara McKnight, Maria Sabater-Lleal, Maristella Steri

Affiliations National Heart, Lung and Blood Institute's Framingham Heart Study, Framingham, Massachusetts, United States of America, National Heart, Lung and Blood Institute Division of Intramural Research, Bethesda, Maryland, United States of America, Department of Human Genetics, Wellcome Trust Sanger Institute, Hinxton, Cambridge, United Kingdom

Barbara McKnight

Contributed equally to this work with: Jens Baumert, Jie Huang, Barbara McKnight, Maria Sabater-Lleal, Maristella Steri

Affiliation Department of Biostatistics, University of Washington, Seattle, Washington, United States of America

Maria Sabater-Lleal

Contributed equally to this work with: Jens Baumert, Jie Huang, Barbara McKnight, Maria Sabater-Lleal, Maristella Steri

Affiliation Cardiovascular Genetics and Genomics Group, Atherosclerosis Research Unit, Department of Medicine, Karolinska Institutet, Karolinska University Hospital Solna, Stockholm, Sweden

Maristella Steri

Contributed equally to this work with: Jens Baumert, Jie Huang, Barbara McKnight, Maria Sabater-Lleal, Maristella Steri

Affiliation Istituto di Ricerca Genetica e Biomedica, Consiglio Nazionale delle Ricerche, Cagliari, Italy

Audrey Y. Chu

Affiliation Division of Preventive Medicine, Brigham and Women's Hospital, Boston, Massachusetts, United States of America

Stella Trompet

Affiliations Department of Cardiology, Leiden University Medical Center, Leiden, the Netherlands, Department of Gerontology and Geriatrics, Leiden University Medical Center, Leiden, the Netherlands

Lorna M. Lopez

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

Myriam Fornage

Affiliation Brown Foundation Institute of Molecular Medicine, Division of Epidemiology, School of Public Health, University of Texas Health Science Center at Houston, Houston, Texas, United States of America

Alexander Teumer

Affiliation Interfaculty Institute for Genetics and Functional Genomics, Ernst-Moritz-Arndt-University Greifswald, Greifswald, Germany

Weihong Tang

Affiliation Division of Epidemiology and Community Health, University of Minnesota, Minneapolis, Minnesota, United States of America

Alicja R. Rudnicka

Affiliation Division of Population Health Sciences & Education, St George's, University of London, Cranmer Terrace, London, United Kingdom

Anders Mälarstig

Affiliation Cardiovascular Genetics and Genomics Group, Atherosclerosis Research Unit, Department of Medicine, Karolinska Institutet, Karolinska University Hospital Solna, Stockholm, Sweden

Jouke-Jan Hottenga

Affiliation Department of Biological Psychology, VU University & EMGO+ institute, VU Medical Centre, Amsterdam, the Netherlands

Maryam Kavousi

Affiliations Department of Epidemiology, Erasmus Medical Center, Rotterdam, the Netherlands, Netherlands Genomics Initiative (NGI)-Sponsored Netherlands Consortium for Healthy Aging (NCHA), Rotterdam, the Netherlands

Jari Lahti

Affiliations Institute of Behavioural Sciences, University of Helsinki, Helsinki, Finland, Folkhalsan Research Centre, Helsinki, Finland

Toshiko Tanaka

Affiliation Translational Gerontology Branch, National Institute on Aging, Baltimore, Maryland, United States of America

Caroline Hayward

Affiliation MRC Human Genetics Unit, Institute of Genetics and Molecular Medicine, Western General Hospital, Edinburgh, Scotland, United Kingdom

Jennifer E. Huffman

Affiliation MRC Human Genetics Unit, Institute of Genetics and Molecular Medicine, Western General Hospital, Edinburgh, Scotland, United Kingdom

Pierre-Emmanuel Morange

Affiliation INSERM UMR_S 1062, Aix-Marseille Université, Marseille, France

Lynda M. Rose

Affiliation Division of Preventive Medicine, Brigham and Women's Hospital, Boston, Massachusetts, United States of America

Saonli Basu

Affiliation Division of Biostatistics, University of Minnesota, Minneapolis, Minnesota, United States of America

Ann Rumley

Affiliation Division of Cardiovascular and Medical Sciences, University of Glasgow, Glasgow, United Kingdom

David J. Stott

Affiliation Institute of Cardiovascular and Medical Sciences, Faculty of Medicine, University of Glasgow, Glasgow, United Kingdom

Brendan M. Buckley

Affiliation Department of Pharmacology and Therapeutics, University College Cork, Cork, Ireland

Anton J. M. de Craen

Affiliation Department of Gerontology and Geriatrics, Leiden University Medical Center, Leiden, the Netherlands

Serena Sanna

Affiliation Istituto di Ricerca Genetica e Biomedica, Consiglio Nazionale delle Ricerche, Cagliari, Italy

Marco Masala

Affiliation Istituto di Ricerca Genetica e Biomedica, Consiglio Nazionale delle Ricerche, Cagliari, Italy

Reiner Biffar

Affiliation Department of Prosthetic Dentistry, Gerostomatology and Dental Materials, University Medicine Greifswald, Greifswald, Germany

Georg Homuth

Affiliation Interfaculty Institute for Genetics and Functional Genomics, Ernst-Moritz-Arndt-University Greifswald, Greifswald, Germany

Angela Silveira

Affiliation Cardiovascular Genetics and Genomics Group, Atherosclerosis Research Unit, Department of Medicine, Karolinska Institutet, Karolinska University Hospital Solna, Stockholm, Sweden

Bengt Sennblad

Affiliations Cardiovascular Genetics and Genomics Group, Atherosclerosis Research Unit, Department of Medicine, Karolinska Institutet, Karolinska University Hospital Solna, Stockholm, Sweden, Science for Life Laboratory, Karolinska Insitutet, Stockholm, Sweden

Anuj Goel

Affiliations Department of Cardiovascular Medicine, University of Oxford, John Radcliffe Hospital, Headington, Oxford, United Kingdom, Department of Cardiovascular Medicine, The Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom

Hugh Watkins

Affiliations Department of Cardiovascular Medicine, University of Oxford, John Radcliffe Hospital, Headington, Oxford, United Kingdom, Department of Cardiovascular Medicine, The Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom

Martina Müller-Nurasyid

Affiliations Institute of Genetic Epidemiology, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, Germany, Institute of Medical Informatics, Biometry and Epidemiology, Chair of Genetic Epidemiology, Ludwig-Maximilians-Universität München, Munich, Germany, Department of Medicine I, University Hospital Grosshadern, Ludwig-Maximilians-Universität München, Munich, Germany

Regina Rückerl

Affiliations Institute of Epidemiology II, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, Germany, ESC-Environmental Science Center, University of Augsburg, Augsburg, Germany

Kent Taylor

Affiliation Medical Genetics Institute, Cedars-Sinai Medical Center, Los Angeles, California, United States of America

Ming-Huei Chen

Affiliation Department of Biostatistics, Boston University, Boston, Massachusetts, United States of America

Eco J. C. de Geus

Affiliation Department of Haematology, Erasmus Medical Center, Rotterdam, the Netherlands

Albert Hofman

Affiliations Department of Epidemiology, Erasmus Medical Center, Rotterdam, the Netherlands, Netherlands Genomics Initiative (NGI)-Sponsored Netherlands Consortium for Healthy Aging (NCHA), Rotterdam, the Netherlands

Jacqueline C. M. Witteman

Affiliations Department of Epidemiology, Erasmus Medical Center, Rotterdam, the Netherlands, Netherlands Genomics Initiative (NGI)-Sponsored Netherlands Consortium for Healthy Aging (NCHA), Rotterdam, the Netherlands

Moniek P. M. de Maat

Affiliation Department of Haematology, Erasmus Medical Center, Rotterdam, the Netherlands

Aarno Palotie

Affiliations Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, United Kingdom, Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland, Department of Medical Genetics, University of Helsinki and University Central Hospital, Helsinki, Finland

Gail Davies

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

David S. Siscovick

Affiliation Department of Epidemiology, University of Washington, Seattle, Washington, United States of America

Ivana Kolcic

Affiliation Department of Public Health, University of Split Medical School, Split, Croatia

Sarah H. Wild

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

Jaejoon Song

Affiliation Division of Biostatistics, University of Minnesota, Minneapolis, Minnesota, United States of America

Wendy L. McArdle

Affiliation School of Social and Community Medicine, University of Bristol, Bristol, United Kingdom

Ian Ford

Affiliation Robertson Center for Biostatistics, University of Glasgow, Glasgow, United Kingdom

Naveed Sattar

Affiliation BHF Glasgow Cardiovascular Research Centre, Faculty of Medicine, Glasgow, United Kingdom

David Schlessinger

Affiliation Intramural Research Program, National Institute on Aging, Baltimore, Maryland, United States of America

Anne Grotevendt

Affiliation Institute for Clinical Chemistry and Laboratory Medicine, University Medicine Greifswald, Greifswald, Germany

Maria Grazia Franzosi

Affiliation Department of Cardiovascular Research, IRCCS Istituto di Ricerche Farmacologiche Mario Negri, Milan, Italy

Thomas Illig

Affiliations Research Unit of Molecular Epidemiology, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, Germany, Hannover Unified Biobank, Hannover Medical School, Hannover, Germany

Melanie Waldenberger

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

Thomas Lumley

Affiliation Department of Statistics, University of Auckland, Auckland, New Zealand

Geoffrey H. Tofler

Affiliation Royal North Shore Hospital, University of Sydney, Sydney, Australia

Gonneke Willemsen

Affiliation Department of Biological Psychology, VU University & EMGO+ institute, VU Medical Centre, Amsterdam, the Netherlands

André G. Uitterlinden

Affiliations Department of Epidemiology, Erasmus Medical Center, Rotterdam, the Netherlands, Netherlands Genomics Initiative (NGI)-Sponsored Netherlands Consortium for Healthy Aging (NCHA), Rotterdam, the Netherlands, Department of Internal Medicine, Erasmus Medical Center, Rotterdam, the Netherlands

Fernando Rivadeneira

Affiliation Division of Preventive Medicine, Division of Genetics, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts, United States of America

Katri Räikkönen

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

Daniel I. Chasman

Affiliation Division of Preventive Medicine, Division of Genetics, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts, United States of America

Aaron R. Folsom

Affiliation Division of Epidemiology and Community Health, University of Minnesota, Minneapolis, Minnesota, United States of America

Gordon D. Lowe

Affiliation Institute of Cardiovascular & Medical Sciences, University of Glasgow, Glasgow, United Kingdom

Rudi G. J. Westendorp

Affiliation Department of Gerontology and Geriatrics, Leiden University Medical Center, Leiden, the Netherlands

P. Eline Slagboom

Affiliation Department of Molecular Epidemiology, Leiden University Medical Center, Leiden, the Netherlands

Francesco Cucca

Affiliation Istituto di Ricerca Genetica e Biomedica, Consiglio Nazionale delle Ricerche, Cagliari, Italy

Henri Wallaschofski

Affiliations Intramural Research Program, National Institute on Aging, Baltimore, Maryland, United States of America, DZHK (German Centre for Cardiovascular Research), partner site Greifswald, Greifswald, Germany

Rona J. Strawbridge

Affiliation Cardiovascular Genetics and Genomics Group, Atherosclerosis Research Unit, Department of Medicine, Karolinska Institutet, Karolinska University Hospital Solna, Stockholm, Sweden

Udo Seedorf

Affiliation Leibniz-Institut für Arterioskleroseforschung an der Universität Münster, Münster, Germany

Wolfgang Koenig

Affiliation Department of Internal Medicine II - Cardiology, University of Ulm Medical Center, Ulm, Germany

Joshua C. Bis

Affiliation Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, Washington, United States of America

Kenneth J. Mukamal

Affiliations Harvard Medical School, Boston, Massachusetts, United States of America, Division of General Medicine and Primary Care, Beth Israel Deaconess Medical Center, Boston, Massachusetts, United States of America

Jenny van Dongen

Affiliation Department of Biological Psychology, VU University & EMGO+ institute, VU Medical Centre, Amsterdam, the Netherlands

Elisabeth Widen

Affiliation Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, United Kingdom

Oscar H. Franco

Affiliations Department of Epidemiology, Erasmus Medical Center, Rotterdam, the Netherlands, Netherlands Genomics Initiative (NGI)-Sponsored Netherlands Consortium for Healthy Aging (NCHA), Rotterdam, the Netherlands

John M. Starr

Affiliations Centre for Cognitive Ageing and Cognitive Epidemiology, The University of Edinburgh, Edinburgh, United Kingdom, Alzheimer Scotland Dementia Research Centre, The University of Edinburgh, Edinburgh, United Kingdom

Kiang Liu

Affiliation Department of Preventive Medicine, Northwestern University Medical School, Chicago, Illinois, United States of America

Luigi Ferrucci

Affiliation Translational Gerontology Branch, National Institute on Aging, Baltimore, Maryland, United States of America

Ozren Polasek

Affiliation Department of Epidemiology, University of Washington, Seattle, Washington, United States of America

James F. Wilson

Affiliation Department of Public Health, University of Split Medical School, Split, Croatia

Tiphaine Oudot-Mellakh

Affiliations INSERM, UMR_S 1166, Paris, France, Sorbonne Universités, UPMC Univ Paris 06, UMR_S 1166, Team Genomics & Pathophysiology of Cardiovascular Diseases, Paris, France, ICAN Institute for Cardiometabolism and Nutrition, Paris, France

Harry Campbell

Affiliation Department of Public Health, University of Split Medical School, Split, Croatia

Pau Navarro

Affiliation MRC Human Genetics Unit, Institute of Genetics and Molecular Medicine, Western General Hospital, Edinburgh, Scotland, United Kingdom

Stefania Bandinelli

Affiliation Geriatric Unit, Azienda Sanitaria Firenze (ASF), Florence, Italy

Johan Eriksson

Affiliations Folkhalsan Research Centre, Helsinki, Finland, National Institute for Health and Welfare, Helsinki, Finland, Department of General Practice and Primary Health Care, University of Helsinki, Helsinki, Finland, Helsinki University Central Hospital, Unit of General Practice, Helsinki, Finland

Dorret I. Boomsma

Affiliation Department of Biological Psychology, VU University & EMGO+ institute, VU Medical Centre, Amsterdam, the Netherlands

Abbas Dehghan

Affiliations Department of Epidemiology, Erasmus Medical Center, Rotterdam, the Netherlands, Netherlands Genomics Initiative (NGI)-Sponsored Netherlands Consortium for Healthy Aging (NCHA), Rotterdam, the Netherlands

Robert Clarke

Affiliation Clinical Trial Service Unit, University of Oxford, Oxford, United Kingdom

Anders Hamsten

Affiliation Cardiovascular Genetics and Genomics Group, Atherosclerosis Research Unit, Department of Medicine, Karolinska Institutet, Karolinska University Hospital Solna, Stockholm, Sweden

Eric Boerwinkle

Affiliation Human Genetics Center and Institute of Molecular Medicine, University of Texas Health Science Center, Houston, Texas, United States of America

J. Wouter Jukema

Affiliations Department of Cardiology, Leiden University Medical Center, Leiden, the Netherlands, Durrer Center for Cardiogenetic Research, Amsterdam, the Netherlands, Interuniversity Cardiology Institute of the Netherlands, Utrecht, the Netherlands

Silvia Naitza

Affiliation Istituto di Ricerca Genetica e Biomedica, Consiglio Nazionale delle Ricerche, Cagliari, Italy

Paul M. Ridker

Affiliation Division of Preventive Medicine, Division of Cardiology, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts, United States of America

Henry Völzke

Affiliation Institute for Community Medicine, University Medicine Greifswald, Greifswald, Germany

Ian J. Deary

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

Alexander P. Reiner

Affiliation Department of Medical Genetics, University of Helsinki and University Central Hospital, Helsinki, Finland

David-Alexandre Trégouët

These authors also contributed equally to this work as last authors.

Affiliations INSERM, UMR_S 1166, Paris, France, Sorbonne Universités, UPMC Univ Paris 06, UMR_S 1166, Team Genomics & Pathophysiology of Cardiovascular Diseases, Paris, France, ICAN Institute for Cardiometabolism and Nutrition, Paris, France

Christopher J. O'Donnell

These authors also contributed equally to this work as last authors.

Affiliations National Heart, Lung and Blood Institute's Framingham Heart Study, Framingham, Massachusetts, United States of America, National Heart, Lung and Blood Institute Division of Intramural Research, Bethesda, Maryland, United States of America

David P. Strachan

These authors also contributed equally to this work as last authors.

Affiliation Division of Population Health Sciences & Education, St George's, University of London, Cranmer Terrace, London, United Kingdom

Annette Peters

peters@helmholtz-muenchen.de (A. Peters); nlsmith@u.washington.edu (NLS)

These authors also contributed equally to this work as last authors.

Affiliations Institute of Epidemiology II, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, Germany, DZHK (German Centre for Cardiovascular Research), partner site Munich, Munich, Germany

Nicholas L. Smith

peters@helmholtz-muenchen.de (A. Peters); nlsmith@u.washington.edu (NLS)

These authors also contributed equally to this work as last authors.

Affiliations Department of Medical Genetics, University of Helsinki and University Central Hospital, Helsinki, Finland, Group Health Research Institute, Group Health Cooperative, Seattle, Washington, United States of America, Seattle Epidemiologic Research & Information Center, Veterans Affairs Office of Research & Development, Seattle, Washington, United States of America

Competing Interests

The authors have the following interests: The Framingham Heart Study (FHS) was supported by the National Heart, Lung and Blood Institute's Framingham Heart Study (Contract No. N01-HC-25195) and its contract with Affymetrix, Inc for genotyping services (Contract No. N02-HL-6-4278). The PROSPER/PHASE study was supported by an investigator initiated grant obtained from Bristol-Myers Squibb. Dr. Kavousi is supported by AXA Research Fund. Oscar H. Franco works in ErasmusAGE, a center for aging research across the life course funded by Nestle Nutrition (Nestec Ltd), Metagenics Inc, and AXA. Part of the SHIP study was funded by a joint grant from Siemens Healthcare, Erlangen, Germany and the Federal State of Mecklenburg West Pomerania. The University of Greifswald is a member of the ‘Center of Knowledge Interchange’ program of the Siemens AG and the Caché Campus program of the InterSystems GmbH. There are no patents, products in development or marketed products to declare. This does not alter the authors' adherence to all the PLOS ONE policies on sharing data and materials, as detailed online in the guide for authors at the time of submission.

Author Contributions

Conceived and designed the experiments: ARF EB (ARIC) DPS (B58C) APR (CARDIA) NLS (CHS) CH IK OP (CROATIA-Vis) CJO (FHS) JE (HBCS) LF S. Bandinelli (InCHIANTI) A. Peters TI WK (KORA) IJD JMS (LBC) DAT (MARTHA) GW EJCDG DIB (NTR) SHW JFW HC (ORCADES) H. Watkins US A. Hamsten (PROCARDIS) IF RGJW JWJ DJS BMB AJMdC PES (PROSPER) AGU AD (RS) DS FC SN (SardiNIA) HV (SHIP) DIC PMR (WGHS). Wrote the paper: WT S. Basu ARF EB (ARIC) DPS (B58C) APR MF (CARDIA) BM KT TL KJM JCB NLS (CHS) CH IK OP PN (CROATIA-Vis) JH CJO (FHS) JL KR EW A. Palotie JE (HBCS) TT (InCHIANTI) JB RR WK A. Peters (KORA) LML GD JMS IJD (LBC) PEM DAT (MARTHA) JJH EJCDG DIB (NTR) JEH SHW JFW HC (ORCADES) MS-L H. Watkins US A. Hamsten (PROCARDIS) ST RGJW JWJ DJS NS AJMdC PES (PROSPER) MK AD (RS) MS SS SN (SardiNIA) AT RB A. Grotevendt H. Wallaschofski HV (SHIP) AYC DIC PMR (WGHS). Performed or contributed data measurements: ARF EB (ARIC) AR WLM GDL DPS (B58C) MF KL DSS (CARDIA) KT KJM (CHS) CH IK OP PN (CROATIA-Vis) MHC GHT (FHS) JL EW A Palotie KR JE (HBCS) LF S. Bandinelli (InCHIANTI) MMN TI MW WK (KORA) GD IJD JMS (LBC) PEM (MARTHA) JJH GW EJCDG DIB (NTR) SHW JFW HC (ORCADES) AM RS AS BS A. Goel MGF RC (PROCARDIS) IF RGJW JWJ DJS BMB (PROSPER) A. Hofman JCMW AGU MPMM FR OHF (RS) MM DS FC (SardiNIA) A. Grotevendt H. Wallaschofski HV (SHIP) LMR DIC PMR (WGHS). Performed data analyses: JS WT S. Basu (ARIC) ARR DPS (B58C) MF (CARDIA) BM JCB (CHS) CH (CROATIA-Vis) JH (FHS) JL (HBCS) TT (InCHIANTI) JB (KORA) LML (LBC) TOM DAT (MARTHA) JJH JVD (NTR) JEH (ORCADES) MS-L (PROCARDIS) ST (PROSPER) MK (RS) MS SS (SardiNIA) AT GH (SHIP) AYC LMR (WGHS). Drafted the manuscript: JB (KORA).