About the Authors

Andrea D. Coviello

Contributed equally to this work with: Andrea D. Coviello, Robin Haring, Melissa Wellons, Dhananjay Vaidya, Terho Lehtimäki, Sarah Keildson

Affiliations Section of Preventive Medicine and Epidemiology, Boston University School of Medicine, Boston, Massachusetts, United States of America, Section of Endocrinology, Diabetes, and Nutrition, Boston University School of Medicine, Boston, Massachusetts, United States of America, National Heart, Lung, and Blood Institute's The Framingham Heart Study, Framingham, Massachusetts, United States of America

Robin Haring

Contributed equally to this work with: Andrea D. Coviello, Robin Haring, Melissa Wellons, Dhananjay Vaidya, Terho Lehtimäki, Sarah Keildson

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

Melissa Wellons

Contributed equally to this work with: Andrea D. Coviello, Robin Haring, Melissa Wellons, Dhananjay Vaidya, Terho Lehtimäki, Sarah Keildson

Affiliation Department of Medicine and Department of Obstetrics and Gynecology, The University of Alabama at Birmingham, Birmingham, Alabama, United States of America

Dhananjay Vaidya

Contributed equally to this work with: Andrea D. Coviello, Robin Haring, Melissa Wellons, Dhananjay Vaidya, Terho Lehtimäki, Sarah Keildson

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

Terho Lehtimäki

Contributed equally to this work with: Andrea D. Coviello, Robin Haring, Melissa Wellons, Dhananjay Vaidya, Terho Lehtimäki, Sarah Keildson

Affiliation Department of Clinical Chemistry, Fimlab Laboratories, Tampere University Hospital and University of Tampere School of Medicine, Tampere, Finland

Sarah Keildson

Contributed equally to this work with: Andrea D. Coviello, Robin Haring, Melissa Wellons, Dhananjay Vaidya, Terho Lehtimäki, Sarah Keildson

Affiliation Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom

Kathryn L. Lunetta

Affiliation Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts, United States of America

Chunyan He

Affiliations Department of Public Health, Indiana University School of Medicine, Indianapolis, Indiana, United States of America, Melvin and Bren Simon Cancer Center, Indiana University, Indianapolis, Indiana, United States of America

Myriam Fornage

Affiliation University of Texas Health Sciences Center at Houston, Houston, Texas, United States of America

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

Massimo Mangino

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

N. Charlotte Onland-Moret

Affiliation Julius Center for Health Sciences and Primary Care, University Medical Center Utrecht, Utrecht, The Netherlands

Brian Chen

Affiliation Program on Genomics and Nutrition and the Center for Metabolic Disease Prevention, School of Public Health, University of California Los Angeles, Los Angeles, California, United States of America

Joel Eriksson

Affiliation Center for Bone and Arthritis Research, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, Gothenburg, Sweden

Melissa Garcia

Affiliation Laboratory of Epidemiology, Demography, and Biometry, National Institute on Aging, Bethesda, Maryland, United States of America

Yong Mei Liu

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

Annemarie Koster

Affiliation Laboratory of Epidemiology, Demography, and Biometry, National Institute on Aging, Bethesda, Maryland, United States of America

Kurt Lohman

Affiliation Wake Forest University School of Medicine, Winston-Salem, North Carolina, United States of America

Leo-Pekka Lyytikäinen

Affiliation Department of Clinical Chemistry, Fimlab Laboratories, Tampere University Hospital and University of Tampere School of Medicine, Tampere, Finland

Ann-Kristin Petersen

Affiliation Institute of Genetic Epidemiology, Helmholtz Zentrum München, Neuherberg, Germany

Jennifer Prescott

Affiliations Program in Molecular and Genetic Epidemiology, Department of Epidemiology, Harvard School of Public Health, Boston, Massachusetts, United States of America, Channing Laboratory, Department of Medicine, Brigham and Women's Hospital, and Harvard Medical School, Boston, Massachusetts, United States of America

Lisette Stolk

Affiliations Department of Internal Medicine, Erasmus MC, Rotterdam, The Netherlands, Netherlands Consortium of Healthy Aging, Rotterdam, The Netherlands

Liesbeth Vandenput

Affiliation Center for Bone and Arthritis Research, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, Gothenburg, Sweden

Andrew R. Wood

Affiliation Genetics of Complex Traits, Peninsula Medical School, University of Exeter, Exeter, United Kingdom

Wei Vivian Zhuang

Affiliation Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts, United States of America

Aimo Ruokonen

Affiliation Institute of Diagnostics, University of Oulu, Oulu, Finland

Anna-Liisa Hartikainen

Affiliation Department of Obstetrics and Gynecology, University Hospital of Oulu, Oulu, Finland

Anneli Pouta

Affiliation National Institute for Health and Welfare and Institute of Health Sciences, University of Oulu, Oulu, Finland

Stefania Bandinelli

Affiliation Geriatric Unit, Azienda Sanitaria di Firenze, Florence, Italy

Reiner Biffar

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

Georg Brabant

Affiliation Experimental and Clinical Endocrinology, University of Lübeck, Lübeck, Germany

David G. Cox

Affiliations Cancer Research Center of Lyon, INSERM U1052, Lyon, France, Department of Epidemiology and Biostatistics, School of Public Health, Imperial College, London, United Kingdom

Yuhui Chen

Affiliation Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom

Steven Cummings

Affiliation California Pacific Medical Center, San Francisco, California, United States of America

Luigi Ferrucci

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

Marc J. Gunter

Affiliation Department of Epidemiology and Biostatistics, School of Public Health, Imperial College, London, United Kingdom

Susan E. Hankinson

Affiliations Channing Laboratory, Department of Medicine, Brigham and Women's Hospital, and Harvard Medical School, Boston, Massachusetts, United States of America, Division of Biostatistics and Epidemiology, University of Massachusetts, Amherst, Massachusetts, United States of America, Department of Epidemiology, Harvard School of Public Health, Boston, Massachusetts, United States of America

Hannu Martikainen

Affiliation Department of Obstetrics and Gynecology, University Hospital of Oulu, Oulu, Finland

Albert Hofman

Affiliations Netherlands Consortium of Healthy Aging, Rotterdam, The Netherlands, Department of Epidemiology, Erasmus MC, Rotterdam, The Netherlands

Georg Homuth

Affiliation Interfaculty Institute for Genetics and Functional Genomics, University of Greifswald, Greifswald, Germany

Thomas Illig

Affiliations Research Unit of Molecular Epidemiology, Helmholtz Zentrum München, Neuherberg, Germany, Hannover Unified Biobank, Hannover Medical School, Hannover, Germany

John-Olov Jansson

Affiliation Center for Bone and Arthritis Research, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, Gothenburg, Sweden

Andrew D. Johnson

Affiliation National Heart, Lung, and Blood Institute's The Framingham Heart Study, Framingham, Massachusetts, United States of America

David Karasik

Affiliation Hebrew SeniorLife Institute for Aging Research and Harvard Medical School, Boston, Massachusetts, United States of America

Magnus Karlsson

Affiliation Clinical and Molecular Osteoporosis Research Unit, Department of Clinical Sciences and Department of Orthopaedics, Lund University, Malmö, Sweden

Johannes Kettunen

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

Douglas P. Kiel

Affiliation Hebrew SeniorLife Institute for Aging Research and Harvard Medical School, Boston, Massachusetts, United States of America

Peter Kraft

Affiliation Program in Molecular and Genetic Epidemiology, Department of Epidemiology, Harvard School of Public Health, Boston, Massachusetts, United States of America

Jingmin Liu

Affiliation Women's Health Initiative Clinical Coordinating Center, Division of Public Health Sciences, Fred Hutchinson Cancer Research Center, Seattle, Washington, United States of America

Östen Ljunggren

Affiliation Department of Medical Sciences, University of Uppsala, Uppsala, Sweden

Mattias Lorentzon

Affiliation Center for Bone and Arthritis Research, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, Gothenburg, Sweden

Marcello Maggio

Affiliation Department of Internal Medicine and Biomedical Sciences, Section of Geriatrics, University of Parma, Parma, Italy

Marcello R. P. Markus

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

Dan Mellström

Affiliation Center for Bone and Arthritis Research, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, Gothenburg, Sweden

Iva Miljkovic

Affiliation University of Pittsburgh, Pittsburgh, Pennsylvania, United States of America

Daniel Mirel

Affiliation Gene Environment Initiative, Program in Medical and Population Genetics, Broad Institute of Harvard and MIT, Boston, Massachusetts, United States of America

Sarah Nelson

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

Laure Morin Papunen

Affiliation Department of Obstetrics and Gynecology, University Hospital of Oulu, Oulu, Finland

Petra H. M. Peeters

Affiliation Julius Center for Health Sciences and Primary Care, University Medical Center Utrecht, Utrecht, The Netherlands

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

Leslie Raffel

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

Martin Reincke

Affiliation Medizinische Klinik and Poliklinik IV, Ludwig-Maximilians University, Munich, Germany

Alex P. Reiner

Affiliation Division of Public Health Sciences, Fred Hutchinson Cancer Research Center, Seattle, Washington, United States of America

Kathryn Rexrode

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

Fernando Rivadeneira

Affiliations Department of Internal Medicine, Erasmus MC, Rotterdam, The Netherlands, Netherlands Consortium of Healthy Aging, Rotterdam, The Netherlands

Stephen M. Schwartz

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

David Siscovick

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

Nicole Soranzo

Affiliations Department of Twin Research and Genetic Epidemiology, King's College London, London, United Kingdom, Human Genetics, Wellcome Trust Sanger Institute, Hinxton, United Kingdom

Doris Stöckl

Affiliations Institute of Epidemiology II, Helmholtz Zentrum München, Neuherberg, Germany, Department of Obstetrics and Gynaecology, Ludwig-Maximilians-University, Munich, Germany

Shelley Tworoger

Affiliations Channing Laboratory, Department of Medicine, Brigham and Women's Hospital, and Harvard Medical School, Boston, Massachusetts, United States of America, Department of Epidemiology, Harvard School of Public Health, Boston, Massachusetts, United States of America

André G. Uitterlinden

Affiliations Department of Internal Medicine, Erasmus MC, Rotterdam, The Netherlands, Netherlands Consortium of Healthy Aging, Rotterdam, The Netherlands, Department of Epidemiology, Erasmus MC, Rotterdam, The Netherlands

Carla H. van Gils

Affiliation Julius Center for Health Sciences and Primary Care, University Medical Center Utrecht, Utrecht, The Netherlands

Ramachandran S. Vasan

Affiliations Section of Preventive Medicine and Epidemiology, Boston University School of Medicine, Boston, Massachusetts, United States of America, National Heart, Lung, and Blood Institute's The Framingham Heart Study, Framingham, Massachusetts, United States of America

H.-Erich Wichmann

Affiliations Institute of Epidemiology I, Helmholtz Zentrum München, Neuherberg, Germany, Institute of Medical Informatics, Biometry, and Epidemiology, Ludwig-Maximilians-Universität, Munich, Germany, Klinikum Großhadern, Munich, Germany

Guangju Zhai

Affiliations Department of Twin Research and Genetic Epidemiology, King's College London, London, United Kingdom, Discipline of Genetics, Faculty of Medicine, Memorial University of Newfoundland, St. John's, Newfoundland, Canada

Shalender Bhasin

Affiliation Section of Endocrinology, Diabetes, and Nutrition, Boston University School of Medicine, Boston, Massachusetts, United States of America

Martin Bidlingmaier

Affiliation Medizinische Klinik and Poliklinik IV, Ludwig-Maximilians University, Munich, Germany

Stephen J. Chanock

Affiliation Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, Maryland, United States of America

Immaculata De Vivo

Affiliations Program in Molecular and Genetic Epidemiology, Department of Epidemiology, Harvard School of Public Health, Boston, Massachusetts, United States of America, Channing Laboratory, Department of Medicine, Brigham and Women's Hospital, and Harvard Medical School, Boston, Massachusetts, United States of America

Tamara B. Harris

Affiliation Laboratory of Epidemiology, Demography, and Biometry, National Institute on Aging, Bethesda, Maryland, United States of America

David J. Hunter

Affiliations Program in Molecular and Genetic Epidemiology, Department of Epidemiology, Harvard School of Public Health, Boston, Massachusetts, United States of America, Channing Laboratory, Department of Medicine, Brigham and Women's Hospital, and Harvard Medical School, Boston, Massachusetts, United States of America

Mika Kähönen

Affiliation Department of Clinical Physiology, Tampere University Hospital and University of Tampere School of Medicine, Tampere, Finland

Simin Liu

Affiliation Program on Genomics and Nutrition, Department of Epidemiology, University of California Los Angeles, Los Angeles, California, United States of America

Pamela Ouyang

Affiliation Division of Cardiology, Johns Hopkins Bayview Medical Center, Baltimore, Maryland, United States of America

Tim D. Spector

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

Yvonne T. van der Schouw

Affiliation Julius Center for Health Sciences and Primary Care, University Medical Center Utrecht, Utrecht, The Netherlands

Jorma Viikari

Affiliation Department of Medicine, Turku University Hospital and University of Turku, Turku, Finland

Henri Wallaschofski

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

Mark I. McCarthy

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

Timothy M. Frayling

Affiliation Genetics of Complex Traits, Peninsula Medical School, University of Exeter, Exeter, United Kingdom

Anna Murray

Affiliation Genetics of Complex Traits, Peninsula Medical School, University of Exeter, Exeter, United Kingdom

Steve Franks

Affiliation Institute of Reproductive and Developmental Biology, Imperial College London, London, United Kingdom

Marjo-Riitta Järvelin

These authors also contributed equally to this work.

Affiliations Department of Biostatistics and Epidemiology, School of Public Health, MRC-HPA Centre for Environment and Health, Faculty of Medicine, Imperial College London, London, United Kingdom, Institute of Health Sciences, University of Oulu, Oulu, Finland, Biocenter Oulu, University of Oulu, Oulu, Finland, National Institute of Health and Welfare, University of Oulu, Oulu, Finland

Frank H. de Jong

These authors also contributed equally to this work.

Affiliation Department of Internal Medicine, Erasmus MC, Rotterdam, The Netherlands

Olli Raitakari

These authors also contributed equally to this work.

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

Alexander Teumer

These authors also contributed equally to this work.

Affiliation Interfaculty Institute for Genetics and Functional Genomics, University of Greifswald, Greifswald, Germany

Claes Ohlsson

These authors also contributed equally to this work.

Affiliation Center for Bone and Arthritis Research, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, Gothenburg, Sweden

Joanne M. Murabito

murabito@bu.edu (JM Murabito); John.perry@pms.ac.uk (JRB Perry)

These authors also contributed equally to this work.

Affiliations National Heart, Lung, and Blood Institute's The Framingham Heart Study, Framingham, Massachusetts, United States of America, Section of General Internal Medicine, Boston University School of Medicine, Boston, Massachusetts, United States of America

John R. B. Perry

murabito@bu.edu (JM Murabito); John.perry@pms.ac.uk (JRB Perry)

These authors also contributed equally to this work.

Affiliations Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom, Department of Twin Research and Genetic Epidemiology, King's College London, London, United Kingdom, Genetics of Complex Traits, Peninsula Medical School, University of Exeter, Exeter, United Kingdom

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: FH de Jong, O Raitakari, A Teumer, C Ohlsson, JM Murabito, JRB Perry. Analyzed the data: JRB Perry, AD Coviello, R Haring, M Wellons, D Vaidya, T Lehtimäki, S Keildson, KL Lunetta, C He, M-R Järvelin. Wrote the paper: AD Coviello, R Haring, M Wellons, D Vaidya, T Lehtimäki, S Keildson, KL Lunetta, C He, TM Frayling, A Murray, S Franks, M-R Järvelin, FH de Jong, O Raitakari, A Teumer, C Ohlsson, JM Murabio, JRB Perry. Individual cohort phenotyping/genotyping: KL Lunetta, C He, M Fornage, V Lagou, M Mangino, NC Onland-Moret, B Chen, J Eriksson, M Garcia, YM Liu, A Koster, K Lohman, L-P Lyytikäinen, A-K Petersen, J Prescott, L Stolk, L Vandenput, AR Wood, WV Zhuang, A Ruokonen, A-L Hartikainen, A Pouta, S Bandinelli, AD Coviello, R Biffar, G Brabant, DG Cox, Y Chen, S Cummings, L Ferrucci, MJ Gunter, SE Hankinson, H Martikainen, A Hofman, G Homuth, T Illig, J-O Jansson, AD Johnson, D Karasik, M Karlsson, J Kettunen, DP Kiel, P Kraft, J Liu, Ö Ljunggren, M Lorentzon, M Maggio, MRP Markus, D Mellström, I Miljkovic, D Mirel, S Nelson, L Morin Papunen, PHM Peeters, I Prokopenko, L Raffel, M Reincke, AP Reiner, K Rexrode, F Rivadeneira, SM Schwartz, D Siscovick, N Soranzo, D Stöckl, S Tworoger, AG Uitterlinden, CH van Gils, RS Vasan, H-E Wichmann, G Zhai, S Bhasin, M Bidlingmaier, SJ Chanock, I De Vivo, TB Harris, DJ Hunter, M Kähönen, S Liu, P Ouyang, TD Spector, YT van der Schouw, J Viikari, H Wallaschofski, MI McCarthy, C Ohlsson, TM Frayling, A Murray, S Franks, M-R Järvelin.