About the Authors

Michael A. Nalls

Contributed equally to this work with: Michael A. Nalls, David J. Couper, Toshiko Tanaka, Frank J. A. van Rooij, Ming-Huei Chen, Albert V. Smith, Daniela Toniolo, Neil A. Zakai

nallsm@mail.nih.gov (MAN); sganesh@umich.edu (SKG)

Affiliation Laboratory of Neurogenetics, Intramural Research Program, National Institute on Aging (NIA), National Institutes of Health (NIH), Bethesda, Maryland, United States of America

David J. Couper

Contributed equally to this work with: Michael A. Nalls, David J. Couper, Toshiko Tanaka, Frank J. A. van Rooij, Ming-Huei Chen, Albert V. Smith, Daniela Toniolo, Neil A. Zakai

Affiliation Collaborative Studies Coordinating Center, Department of Biostatistics, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, United States of America

Toshiko Tanaka

Contributed equally to this work with: Michael A. Nalls, David J. Couper, Toshiko Tanaka, Frank J. A. van Rooij, Ming-Huei Chen, Albert V. Smith, Daniela Toniolo, Neil A. Zakai

Affiliation Longitudinal Studies Section, Clinical Research Branch, NIA, NIH, Baltimore, Maryland, United States of America

Frank J. A. van Rooij

Contributed equally to this work with: Michael A. Nalls, David J. Couper, Toshiko Tanaka, Frank J. A. van Rooij, Ming-Huei Chen, Albert V. Smith, Daniela Toniolo, Neil A. Zakai

Affiliations Department of Epidemiology, Erasmus MC, Rotterdam, The Netherlands, Netherlands Consortium for Healthy Aging (NGI-NCHA), The Netherlands Genomics Initiative, Leiden, The Netherlands

Ming-Huei Chen

Contributed equally to this work with: Michael A. Nalls, David J. Couper, Toshiko Tanaka, Frank J. A. van Rooij, Ming-Huei Chen, Albert V. Smith, Daniela Toniolo, Neil A. Zakai

Affiliations National Heart, Lung, and Blood Institute's Framingham Heart Study, Framingham, Massachusetts, United States of America, Department of Neurology, Boston University School of Medicine, Boston, Massachusetts, United States of America

Albert V. Smith

Contributed equally to this work with: Michael A. Nalls, David J. Couper, Toshiko Tanaka, Frank J. A. van Rooij, Ming-Huei Chen, Albert V. Smith, Daniela Toniolo, Neil A. Zakai

Affiliations Icelandic Heart Association, Kopavogur, Iceland, University of Iceland, Reykjavik, Iceland

Daniela Toniolo

Contributed equally to this work with: Michael A. Nalls, David J. Couper, Toshiko Tanaka, Frank J. A. van Rooij, Ming-Huei Chen, Albert V. Smith, Daniela Toniolo, Neil A. Zakai

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

Neil A. Zakai

Contributed equally to this work with: Michael A. Nalls, David J. Couper, Toshiko Tanaka, Frank J. A. van Rooij, Ming-Huei Chen, Albert V. Smith, Daniela Toniolo, Neil A. Zakai

Affiliations Department of Medicine University of Vermont College of Medicine, Burlington, Vermont, United States of America, Department of Pathology University of Vermont College of Medicine, Burlington, Vermont, United States of America

Qiong Yang

Affiliations National Heart, Lung, and Blood Institute's Framingham Heart Study, Framingham, Massachusetts, United States of America, Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts, United States of America

Andreas Greinacher

Affiliation Institute of Immunology and Transfusion Medicine, Ernst-Moritz-Arndt-University Greifswald, Greifswald, Germany

Andrew R. Wood

Affiliation Genetics of Complex Traits, Peninsula College of Medicine and Dentistry, University of Exeter, United Kingdom

Melissa Garcia

Affiliation Laboratory for Epidemiology, Demography, and Biometry, NIA, NIH, Bethesda, Maryland, United States of America

Paolo Gasparini

Affiliation Medical Genetics, IRCCS–Burlo Garofolo/University of Trieste, Trieste, Italy

Yongmei Liu

Affiliation Department of Epidemiology and Prevention, Division of Public Health Sciences, Wake Forest University, Winston-Salem, North Carolina, United States of America

Thomas Lumley

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

Aaron R. Folsom

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

Alex P. Reiner

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

Christian Gieger

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

Vasiliki Lagou

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

Janine F. Felix

Affiliations Department of Epidemiology, Erasmus MC, Rotterdam, The Netherlands, Netherlands Consortium for Healthy Aging (NGI-NCHA), The Netherlands Genomics Initiative, Leiden, The Netherlands, Division of Clinical Epidemiology and Aging Research, German Cancer Research Center, Heidelberg, Germany

Henry Völzke

Affiliation Community Medicine, Ernst-Moritz-Arndt-University Greifswald, Greifswald, Germany

Natalia A. Gouskova

Affiliation University of North Carolina, School of Public Health, United States of America

Alessandro Biffi

Affiliations Center for Human Genetic Research, Department of Neurology, Massachusetts General Hospital, Boston, Massachusetts, United States of America, Program in Medical and Population Genetics, Broad Institute, Cambridge, Massachusetts, United States of America

Angela Döring

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

Uwe Völker

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

Sean Chong

Affiliation Laboratory of Neurogenetics, Intramural Research Program, National Institute on Aging (NIA), National Institutes of Health (NIH), Bethesda, Maryland, United States of America

Kerri L. Wiggins

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

Augusto Rendon

Affiliation Department of Haematology, University of Cambridge and National Health Service Blood and Transplant, Cambridge, United Kingdom

Abbas Dehghan

Affiliations Department of Epidemiology, Erasmus MC, Rotterdam, The Netherlands, Netherlands Consortium for Healthy Aging (NGI-NCHA), The Netherlands Genomics Initiative, Leiden, The Netherlands

Matt Moore

Affiliation Laboratory of Neurogenetics, Intramural Research Program, National Institute on Aging (NIA), National Institutes of Health (NIH), Bethesda, Maryland, United States of America

Kent Taylor

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

James G. Wilson

Affiliation Department of Physiology and Biophysics, University of Mississippi Medical Center, Jackson, Mississippi, United States of America

Guillaume Lettre

Affiliation Montreal Heart Institute and Universite de Montreal, Montreal, Canada

Albert Hofman

Affiliations Department of Epidemiology, Erasmus MC, Rotterdam, The Netherlands, Netherlands Consortium for Healthy Aging (NGI-NCHA), The Netherlands Genomics Initiative, Leiden, The Netherlands

Joshua C. Bis

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

Nicola Pirastu

Affiliation Medical Genetics, IRCCS–Burlo Garofolo/University of Trieste, Trieste, Italy

Caroline S. Fox

Affiliations National Heart, Lung, and Blood Institute's Framingham Heart Study, Framingham, Massachusetts, United States of America, Division of Endocrinology, Brigham and Women's Hospital and Harvard Medical School, Boston, United States of America

Christa Meisinger

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

Jennifer Sambrook

Affiliation Department of Haematology, University of Cambridge and National Health Service Blood and Transplant, Cambridge, United Kingdom

Sampath Arepalli

Affiliation Laboratory of Neurogenetics, Intramural Research Program, National Institute on Aging (NIA), National Institutes of Health (NIH), Bethesda, Maryland, United States of America

Matthias Nauck

Affiliation Institute for Clinical Chemistry and Laboratory Medicine, Ernst-Moritz-Arndt-University Greifswald, Greifswald, Germany

Holger Prokisch

Affiliations Institute of Human Genetics, Klinikum rechts der Isar, Technical University Munich, Munich, Germany, Institute of Human Genetics, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, Germany

Jonathan Stephens

Affiliation Department of Haematology, University of Cambridge and National Health Service Blood and Transplant, Cambridge, United Kingdom

Nicole L. Glazer

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

L. Adrienne Cupples

Affiliations National Heart, Lung, and Blood Institute's Framingham Heart Study, Framingham, Massachusetts, United States of America, Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts, United States of America

Yukinori Okada

Affiliations Laboratory for Statistical Analysis, Center for Genomic Medicine (CGM), Institute of Physical and Chemical Research (RIKEN), Yokohama, Japan, Department of Allergy and Rheumatology, Graduate School of Medicine, University of Tokyo, Tokyo, Japan

Atsushi Takahashi

Affiliation Laboratory for Statistical Analysis, Center for Genomic Medicine (CGM), Institute of Physical and Chemical Research (RIKEN), Yokohama, Japan

Yoichiro Kamatani

Affiliation Centre d'Etude du Polymorphisme Humain (CEPH), Paris, France

Koichi Matsuda

Affiliation Laboratory of Molecular Medicine, Human Genome Center, Institute of Medical Science, University of Tokyo, Tokyo, Japan

Tatsuhiko Tsunoda

Affiliation Laboratory for Medical Informatics, CGM, RIKEN, Yokohama, Japan

Toshihiro Tanaka

Affiliation Laboratory for Cardiovascular Diseases, CGM, RIKEN, Yokohama, Japan

Michiaki Kubo

Affiliation Laboratory for Genotyping Development, CGM, RIKEN, Yokohama, Japan

Yusuke Nakamura

Affiliation Laboratory of Molecular Medicine, Human Genome Center, Institute of Medical Science, University of Tokyo, Tokyo, Japan

Kazuhiko Yamamoto

Affiliation Department of Allergy and Rheumatology, Graduate School of Medicine, University of Tokyo, Tokyo, Japan

Naoyuki Kamatani

Affiliation Laboratory for Statistical Analysis, Center for Genomic Medicine (CGM), Institute of Physical and Chemical Research (RIKEN), Yokohama, Japan

Michael Stumvoll

Affiliations Department of Medicine, University of Leipzig, Leipzig, Germany, LIFE Study Centre, University of Leipzig, Leipzig, Germany

Anke Tönjes

Affiliation Department of Medicine, University of Leipzig, Leipzig, Germany

Inga Prokopenko

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

Thomas Illig

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

Kushang V. Patel

Affiliation Laboratory for Epidemiology, Demography, and Biometry, NIA, NIH, Bethesda, Maryland, United States of America

Stephen F. Garner

Affiliation Department of Haematology, University of Cambridge and National Health Service Blood and Transplant, Cambridge, United Kingdom

Brigitte Kuhnel

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

Massimo Mangino

Affiliation Department of Twin Research and Genetic Epidemiology, King's College London, London, United Kingdom

Ben A. Oostra

Affiliations Department of Epidemiology, Erasmus MC, Rotterdam, The Netherlands, Netherlands Consortium for Healthy Aging (NGI-NCHA), The Netherlands Genomics Initiative, Leiden, The Netherlands, Department of Clinical Genetics, Erasmus MC, Rotterdam, The Netherlands

Swee Lay Thein

Affiliation Molecular Haematology, King's College London, London, United Kingdom

Josef Coresh

Affiliation Department of Epidemiology, Johns Hopkins University, Baltimore, Maryland, United States of America

H.-Erich Wichmann

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

Stephan Menzel

Affiliation Molecular Haematology, King's College London, London, United Kingdom

JingPing Lin

Affiliation Office of Biostatistical Research, Division of Cardiovascular Sciences, NHLBI, NIH, Bethesda, Maryland, United States of America

Giorgio Pistis

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

André G. Uitterlinden

Affiliations Department of Epidemiology, Erasmus MC, Rotterdam, The Netherlands, Netherlands Consortium for Healthy Aging (NGI-NCHA), The Netherlands Genomics Initiative, Leiden, The Netherlands, Department of Internal Medicine, Erasmus MC, Rotterdam, The Netherlands

Tim D. Spector

Affiliation Department of Twin Research and Genetic Epidemiology, King's College London, London, United Kingdom

Alexander Teumer

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

Gudny Eiriksdottir

Affiliation Icelandic Heart Association, Kopavogur, Iceland

Vilmundur Gudnason

Affiliations Icelandic Heart Association, Kopavogur, Iceland, University of Iceland, Reykjavik, Iceland

Stefania Bandinelli

Affiliation Geriatric Unit ASF, Firenze, Italy

Timothy M. Frayling

Affiliation Genetics of Complex Traits, Peninsula College of Medicine and Dentistry, University of Exeter, United Kingdom

Aravinda Chakravarti

Affiliation McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University, Baltimore, Maryland, United States of America

Cornelia M. van Duijn

Affiliations Department of Epidemiology, Erasmus MC, Rotterdam, The Netherlands, Netherlands Consortium for Healthy Aging (NGI-NCHA), The Netherlands Genomics Initiative, Leiden, The Netherlands

David Melzer

Affiliations Epidemiology and Public Health, Peninsula College of Medicine and Dentistry, University of Exeter, Exeter, United Kingdom, The European Centre for Environment and Human Health, PCMD, Truro, United Kingdom

Willem H. Ouwehand

Affiliations Department of Haematology, University of Cambridge and National Health Service Blood and Transplant, Cambridge, United Kingdom, Wellcome Trust Sanger Institute, Hinxton, United Kingdom

Daniel Levy

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

Eric Boerwinkle

Affiliation Human Genetics Center, University of Texas Health Science Center, Houston, Texas, United States of America

Andrew B. Singleton

Affiliation Laboratory of Neurogenetics, Intramural Research Program, National Institute on Aging (NIA), National Institutes of Health (NIH), Bethesda, Maryland, United States of America

Dena G. Hernandez

These authors also contributed equally to this work.

Affiliations Laboratory of Neurogenetics, Intramural Research Program, National Institute on Aging (NIA), National Institutes of Health (NIH), Bethesda, Maryland, United States of America, Department of Molecular Neuroscience and Reta Lila Laboratories, Institute of Neurology, University College London, London, United Kingdom

Dan L. Longo

These authors also contributed equally to this work.

Affiliation Clinical Research Branch, National Institute on Aging, Baltimore, Maryland, United States of America

Nicole Soranzo

These authors also contributed equally to this work.

Affiliation Wellcome Trust Sanger Institute, Hinxton, United Kingdom

Jacqueline C. M. Witteman

These authors also contributed equally to this work.

Affiliations Department of Epidemiology, Erasmus MC, Rotterdam, The Netherlands, Netherlands Consortium for Healthy Aging (NGI-NCHA), The Netherlands Genomics Initiative, Leiden, The Netherlands

Bruce M. Psaty

These authors also contributed equally to this work.

Affiliations Departments of Epidemiology, Medicine and Health Services, University of Washington, Seattle, Washington, United States of America, Group Health Research Institute, Group Health, Seattle, Washington, United States of America

Luigi Ferrucci

These authors also contributed equally to this work.

Affiliation Longitudinal Studies Section, Clinical Research Branch, NIA, NIH, Baltimore, Maryland, United States of America

Tamara B. Harris

These authors also contributed equally to this work.

Affiliation Laboratory for Epidemiology, Demography, and Biometry, NIA, NIH, Bethesda, Maryland, United States of America

Christopher J. O'Donnell

These authors also contributed equally to this work.

Affiliations National Heart, Lung, and Blood Institute's Framingham Heart Study, Framingham, Massachusetts, United States of America, Division of Intramural Research, National Heart, Lung, and Blood Institute (NHLBI), Bethesda, Maryland, United States of America, Cardiology Division, Massachusetts General Hospital, Harvard Medical School, Boston, Massachusetts, United States of America

Santhi K. Ganesh

nallsm@mail.nih.gov (MAN); sganesh@umich.edu (SKG)

Affiliation Division of Cardiovascular Medicine, Department of Internal Medicine, University of Michigan, Ann Arbor, Michigan, United States of America

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: Michael A Nalls, Dan L Longo, Luigi Ferrucci, Tamara B Harris, Christopher J O'Donnell, Santhi K Ganesh. Performed the experiments: Andrew R Wood, Yongmei Liu, Thomas Lumley, Aaron R Folsom, Janine F Felix, Henry Völzke, Natalia A Gouskova, Angela Döring, Uwe Völker, Sean Chong, Kerri L Wiggins, Matt Moore, Kent Taylor, James G Wilson, Albert Hoffman, Joshua C Bis, Nicola Pirastu, Caroline S Fox, Christa Meisinger, Jennifer Sambrook, Sampath Arepalli, Matthias Nauck, Holger Prokisch, Jonathan Stephens, Nicole L Glazer, L Adrienne Cupples, Atsushi Takahashi, Yoichiro Kamatani, Koichi Matsuda, Tatsuhiko Tsunoda, Toshihiro Tanaka, Michiaki Kubo, Yusuke Nakamura, Kazuhiko Yamamoto, Naoyuki Kamatani, Michael Stumvoll, Thomas Illig, Kushang V Patel, Stephen F Garner, Brigitte Kuhnel, Massimo Mangino, Ben A Oostra, Josef Coresh, H-Erich Wichmann, Stephan Menzel, JingPing Lin, Giorgio Pistis, Andre G Utterlinden, Tim D Spector, Alexander Teumer, Gudny Eriksdottir, Vilmundur Gudnason, Stefania Bandinelli, Timothy M Frayling, Aravinda Chakravarti, Cornelia M van Duijn, David Melzer, Willem H Ouwehand, Daniel Levy, Eric Boerwinkle, Andrew B Singleton, Dena G Hernandez, Dan L Longo, Nicole Soranzo, Jacqueline CM Witteman, Bruce M Psaty, Luigi Ferrucci, Tamara B Harris, Christopher J O'Donnell, Santhi K Ganesh. Analyzed the data: Michael A Nalls, David J Couper, Toshiko Tanaka, Frank JA van Rooji, Ming-Huei Chen, Albert V Smith, Daniela Tonolio, Neil A Zakai, Qiong Yang, Andreas Greinacher, Melissa Garcia, Paolo Gasparini, Alex P Reiner, Christian Gieger, Vasiliki Lagou, Alessandro Biffi, Augusto Rendon, Abbas Dehghan, Guillaume Lettre, Joshua C Bis, Yukinori Okada, Atsushi Tojnes, Inga Prokopenko, Swee L Thein, David Melzer, Nicole Soranzo, Santhi K Ganesh. Contributed reagents/materials/analysis tools: Andrew R Wood, Yongmei Liu, Thomas Lumley, Aaron R Folsom, Janine F Felix, Henry Völzke, Natalia A Gouskova, Angela Döring, Uwe Völker, Sean Chong, Kerri L Wiggins, Matt Moore, Kent Taylor, James G Wilson, Albert Hoffman, Joshua C Bis, Nicola Pirastu, Caroline S Fox, Christa Meisinger, Jennifer Sambrook, Sampath Arepalli, Matthias Nauck, Holger Prokisch, Jonathan Stephens, Nicole L Glazer, L Adrienne Cupples, Atsushi Takahashi, Yoichiro Kamatani, Koichi Matsuda, Tatsuhiko Tsunoda, Toshihiro Tanaka, Michiaki Kubo, Yusuke Nakamura, Kazuhiko Yamamoto, Naoyuki Kamatani, Michael Stumvoll, Thomas Illig, Kushang V Patel, Stephen F Garner, Brigitte Kuhnel, Massimo Mangino, Ben A Oostra, Josef Coresh, H-Erich Wichmann, Stephan Menzel, JingPing Lin, Giorgio Pistis, Andre G Utterlinden, Tim D Spector, Alexander Teumer, Gudny Eriksdottir, Vilmundur Gudnason, Stefania Bandinelli, Timothy M Frayling, Aravinda Chakravarti, Cornelia M van Duijn, David Melzer, Willem H Ouwehand, Daniel Levy, Eric Boerwinkle, Andrew B Singleton, Dena G Hernandez, Dan L Longo, Nicole Soranzo, Jacqueline CM Witteman, Bruce M Psaty, Luigi Ferrucci, Tamara B Harris, Christopher J O'Donnell, Santhi K Ganesh. Wrote the paper: Michael A Nalls, Santhi K Ganesh.