About the Authors

Hou-Feng Zheng

Contributed equally to this work with: Hou-Feng Zheng, Jon H. Tobias, Emma Duncan, David M. Evans

Affiliations Department of Medicine, Human Genetics, McGill University, Montreal, Canada, Department of Epidemiology and Biostatistics, Lady Davis Institute for Medical Research, Jewish General Hospital, Montreal, Canada

Jon H. Tobias

Contributed equally to this work with: Hou-Feng Zheng, Jon H. Tobias, Emma Duncan, David M. Evans

Affiliation Musculoskeletal Research Unit, School of Clinical Sciences, University of Bristol, Bristol, United Kingdom

Emma Duncan

Contributed equally to this work with: Hou-Feng Zheng, Jon H. Tobias, Emma Duncan, David M. Evans

Affiliations Human Genetics Group, University of Queensland Diamantina Institute, Princess Alexandra Hospital, University of Queensland, Brisbane, Australia, Endocrinology, Royal Brisbane and Women's Hospital, Brisbane, Australia

David M. Evans

Contributed equally to this work with: Hou-Feng Zheng, Jon H. Tobias, Emma Duncan, David M. Evans

Affiliations Medical Research Council Centre for Causal Analyses in Translational Epidemiology, University of Bristol, Bristol, United Kingdom, School of Social and Community Medicine, University of Bristol, Bristol, United Kingdom

Joel Eriksson

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

Lavinia Paternoster

Affiliations Medical Research Council Centre for Causal Analyses in Translational Epidemiology, University of Bristol, Bristol, United Kingdom, School of Social and Community Medicine, University of Bristol, Bristol, United Kingdom

Laura M. Yerges-Armstrong

Affiliation Department of Medicine, Division of Endocrinology, Diabetes, and Nutrition, University of Maryland School of Medicine, Baltimore, Maryland, United States of America

Terho Lehtimäki

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

Ulrica Bergström

Affiliation Surgical and Perioperative Sciences, Umeå University, Umeå, Sweden

Mika Kähönen

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

Paul J. Leo

Affiliation Human Genetics Group, University of Queensland Diamantina Institute, Princess Alexandra Hospital, University of Queensland, Brisbane, Australia

Olli Raitakari

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

Marika Laaksonen

Affiliation Department of Food and Environmental Sciences, University of Helsinki, Helsinki, Finland

Geoffrey C. Nicholson

Affiliation Rural Clinical School, The University of Queensland, Toowoomba, Australia

Jorma Viikari

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

Martin Ladouceur

Affiliation Montreal Heart Institute, Research Institute, Montreal, Canada

Leo-Pekka Lyytikäinen

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

Carolina Medina-Gomez

Affiliations Department of Internal Medicine, Erasmus University Medical Center, Rotterdam, The Netherlands, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands

Fernando Rivadeneira

Affiliations Department of Internal Medicine, Erasmus University Medical Center, Rotterdam, The Netherlands, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands

Richard L. Prince

Affiliations Endocrinology and Diabetes, Sir Charles Gairdner Hospital, Perth, Australia, School of Medicine and Pharmacology, University of Western Australia, Perth, Australia

Harri Sievanen

Affiliation Bone Research Group, UKK Institute, Tampere, Finland

William D. Leslie

Affiliation Department of Internal Medicine, University of Manitoba, Winnipeg, Canada

Dan Mellström

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

John A. Eisman

Affiliation Garvan Institute of Medical Research, University of New South Wales, Sydney, Australia

Sofia Movérare-Skrtic

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

David Goltzman

Affiliation Department of Medicine, McGill University, Montreal, Canada

David A. Hanley

Affiliation Department of Medicine, University of Calgary, Calgary, Canada

Graeme Jones

Affiliation Menzies Research Institute, University of Tasmania, Hobart, Australia

Beate St. Pourcain

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

Yongjun Xiao

Affiliation Centre for Bone and Periodontal Research, McGill University, Montreal, Canada

Nicholas J. Timpson

Affiliations Medical Research Council Centre for Causal Analyses in Translational Epidemiology, University of Bristol, Bristol, United Kingdom, School of Social and Community Medicine, University of Bristol, Bristol, United Kingdom

George Davey Smith

Affiliations Medical Research Council Centre for Causal Analyses in Translational Epidemiology, University of Bristol, Bristol, United Kingdom, School of Social and Community Medicine, University of Bristol, Bristol, United Kingdom

Ian R. Reid

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

Susan M. Ring

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

Philip N. Sambrook

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

Magnus Karlsson

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

Elaine M. Dennison

Affiliation Medical Research Council Lifecourse Epidemiology Unit, University of Southampton, Southampton, United Kingdom

John P. Kemp

Affiliations Medical Research Council Centre for Causal Analyses in Translational Epidemiology, University of Bristol, Bristol, United Kingdom, School of Social and Community Medicine, University of Bristol, Bristol, United Kingdom

Patrick Danoy

Affiliation Human Genetics Group, University of Queensland Diamantina Institute, Princess Alexandra Hospital, University of Queensland, Brisbane, Australia

Adrian Sayers

Affiliation Musculoskeletal Research Unit, School of Clinical Sciences, University of Bristol, Bristol, United Kingdom

Scott G. Wilson

Affiliations Endocrinology and Diabetes, Sir Charles Gairdner Hospital, Perth, Australia, School of Medicine and Pharmacology, University of Western Australia, Perth, Australia, Twin Research and Genetic Epidemiology, King's College London, London, United Kingdom

Maria Nethander

Affiliation Genomics Core Facility, University of Gothenburg, Gothenburg, Sweden

Eugene McCloskey

Affiliations Academic Unit of Bone Metabolism, Metabolic Bone Centre, University of Sheffield, Sheffield, United Kingdom, NIHR Musculoskeletal Biomedical Research Unit, Sheffield Teaching Hospitals, Sheffield, United Kingdom

Liesbeth Vandenput

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

Richard Eastell

Affiliation Academic Unit of Bone Metabolism, Metabolic Bone Centre, University of Sheffield, Sheffield, United Kingdom

Jeff Liu

Affiliation Lexicon Pharmaceuticals, The Woodlands, Texas, United States of America

Tim Spector

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

Braxton D. Mitchell

Affiliation Department of Medicine, Division of Endocrinology, Diabetes, and Nutrition, University of Maryland School of Medicine, Baltimore, Maryland, United States of America

Elizabeth A. Streeten

Affiliations Department of Medicine, Division of Endocrinology, Diabetes, and Nutrition, University of Maryland School of Medicine, Baltimore, Maryland, United States of America, Geriatric Research and Education Clinical Center (GRECC), Veterans Administration Medical Center, Baltimore, Maryland, United States of America

Robert Brommage

Affiliation Lexicon Pharmaceuticals, The Woodlands, Texas, United States of America

Ulrika Pettersson-Kymmer

Affiliations Department of Pharmacology and Neuroscience, Umeå University, Umeå, Sweden, Department of Public Health and Clinical Medicine, Umeå Unviersity, Umeå, Sweden

Matthew A. Brown

Affiliation Human Genetics Group, University of Queensland Diamantina Institute, Princess Alexandra Hospital, University of Queensland, Brisbane, Australia

Claes Ohlsson

claes.ohlsson@medic.gu.se

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

J. Brent Richards

These authors also contributed equally to this work.

Affiliations Department of Medicine, Human Genetics, McGill University, Montreal, Canada, Department of Epidemiology and Biostatistics, Lady Davis Institute for Medical Research, Jewish General Hospital, Montreal, Canada, Twin Research and Genetic Epidemiology, King's College London, London, United Kingdom

Mattias Lorentzon

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

Competing Interests

R Brommage and J Liu are full-time employees of Lexicon Pharmaceuticals. All other authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: H-F Zheng, E Duncan, J Eriksson, U Pettersson-Kymmer, MA Brown, C Ohlsson, R Brommage, JB Richards, JH Tobias, DM Evans, L Paternoster, T Lehtimäki, U Bergström, M Kähönen, J Viikari, M Laaksonen, H Sievanen, D Mellström, M Karlsson, M Lorentzon. Performed the experiments: Y Xiao, J Liu, R Brommage, JH Tobias, T Lehtimäki, M Kähönen, O Raitakari, J Viikari, M Laaksonen, B St. Pourcain, NJ Timpson, GD Smith, SM Ring, EM Dennison, M Lorentzon. Analyzed the data: H-F Zheng, LM Yerges-Armstrong, PJ Leo, L Vandenput, M Ladouceur, EA Streeten, BD Mitchell, JB Richards, JH Tobias, DM Evans, L Paternoster, U Pettersson-Kymmer, C Medina-Gomez, F Rivadeneira, S Movérare-Skrtic, JP Kemp, A Sayers, M Nethander, M Lorentzon, C Ohlsson, R Brommage, L-P Lyytikäinen. Contributed reagents/materials/analysis tools: GC Nicholson, RL Prince, WD Leslie, J Eriksson, D Goltzman, G Jones, IR Reid, PN Sambrook, E Duncan, P Danoy, T Spector, R Eastell, E McCloskey, SG Wilson, M Lorentzon, C Ohlsson, R Brommage, JH Tobias, DM Evans, L-P Pyytikäinen, T Lehtimäki, H Sievanen, B St. Pourcain, M Laaksonen, U Bergström, M Karlsson, O Raitakari, J Viikari, EM Dennison, SM Ring, JP Kemp, GD Smith, NJ Timpson, D Mellström, DA Hanley, JA Eisman. Wrote the paper: H-F Zheng, JB Richards, M Lorentzon, JH Tobias. Critically revised the manuscript: DA Hanley.