About the Authors

John P. Kemp

Contributed equally to this work with: John P. Kemp, Carolina Medina-Gomez

j.p.kemp@bristol.ac.uk

Affiliations MRC Integrative Epidemiology Unit at the University of Bristol, Bristol, United Kingdom, University of Queensland Diamantina Institute, Translational Research Institute, Brisbane, Queensland, Australia

Carolina Medina-Gomez

Contributed equally to this work with: John P. Kemp, Carolina Medina-Gomez

Affiliations Department of Internal Medicine, Erasmus University Medical Center, Rotterdam, The Netherlands, The Generation R Study Group, Erasmus University Medical Center, Rotterdam, The Netherlands, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands, Netherlands Genomics Initiative (NGI)-sponsored Netherlands Consortium for Healthy Aging (NCHA), The Netherlands

Karol Estrada

Affiliation Analytic and Translational Genetics Unit, Massachusetts General Hospital, Boston, Massachusetts, United States of America

Beate St Pourcain

Affiliations MRC Integrative Epidemiology Unit at the University of Bristol, Bristol, United Kingdom, School of Oral and Dental Sciences, University of Bristol, Bristol, United Kingdom, School of Experimental Psychology, University of Bristol, Bristol, United Kingdom

Denise H. M. Heppe

Affiliations The Generation R Study Group, Erasmus University Medical Center, Rotterdam, The Netherlands, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands, Department of Paediatrics, Erasmus University Medical Center, Rotterdam, The Netherlands

Nicole M. Warrington

Affiliation University of Queensland Diamantina Institute, Translational Research Institute, Brisbane, Queensland, Australia

Ling Oei

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

Susan M. Ring

Affiliation MRC Integrative Epidemiology Unit at the University of Bristol, Bristol, United Kingdom

Claudia J. Kruithof

Affiliations The Generation R Study Group, Erasmus University Medical Center, Rotterdam, The Netherlands, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands

Nicholas J. Timpson

Affiliation MRC Integrative Epidemiology Unit at the University of Bristol, Bristol, United Kingdom

Lisa E. Wolber

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

Sjur Reppe

Affiliation Department of Medical Biochemistry, Oslo University Hospital, Ullevaal, Oslo, Norway

Kaare Gautvik

Affiliations Department of Medical Biochemistry, Oslo University Hospital, Ullevaal, Oslo, Norway, Department of Medical Biochemistry, Oslo Deacon Hospital, Oslo, Norway

Elin Grundberg

Affiliations Department of Human Genetics, McGill University, Montréal, Canada, McGill University and Genome Québec Innovation Centre, Montréal, Canada

Bing Ge

Affiliation McGill University and Genome Québec Innovation Centre, Montréal, Canada

Bram van der Eerden

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

Jeroen van de Peppel

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

Matthew A. Hibbs

Affiliations Department of Computer Science, Trinity University, San Antonio, Texas, United States of America, The Jackson Laboratory, Bar Harbor, Maine, United States of America

Cheryl L. Ackert-Bicknell

Affiliation The Jackson Laboratory, Bar Harbor, Maine, United States of America

Kwangbom Choi

Affiliation The Jackson Laboratory, Bar Harbor, Maine, United States of America

Daniel L. Koller

Affiliation Department of Medical and Molecular Genetics, Indiana University School of Medicine, Indianapolis, Indiana, United States of America

Michael J. Econs

Affiliations Department of Medical and Molecular Genetics, Indiana University School of Medicine, Indianapolis, Indiana, United States of America, Department of Medicine, Indiana University School of Medicine, Indianapolis, Indiana, United States of America

Frances M. K. Williams

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

Tatiana Foroud

Affiliation Department of Medical and Molecular Genetics, Indiana University School of Medicine, Indianapolis, Indiana, United States of America

M. Carola Zillikens

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

Claes Ohlsson

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

Albert Hofman

Affiliations The Generation R Study Group, Erasmus University Medical Center, Rotterdam, The Netherlands, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands, Netherlands Genomics Initiative (NGI)-sponsored Netherlands Consortium for Healthy Aging (NCHA), The Netherlands

André G. Uitterlinden

Affiliations Department of Internal Medicine, Erasmus University Medical Center, Rotterdam, The Netherlands, The Generation R Study Group, Erasmus University Medical Center, Rotterdam, The Netherlands, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands, Netherlands Genomics Initiative (NGI)-sponsored Netherlands Consortium for Healthy Aging (NCHA), The Netherlands

George Davey Smith

Affiliation MRC Integrative Epidemiology Unit at the University of Bristol, Bristol, United Kingdom

Vincent W. V. Jaddoe

Affiliations The Generation R Study Group, Erasmus University Medical Center, Rotterdam, The Netherlands, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands, Department of Paediatrics, Erasmus University Medical Center, Rotterdam, The Netherlands

Jonathan H. Tobias

JHT, FR and DME are joint senior authors on this work.

Affiliation School of Clinical Sciences, University of Bristol, Bristol, United Kingdom

Fernando Rivadeneira

JHT, FR and DME are joint senior authors on this work.

Affiliations Department of Internal Medicine, Erasmus University Medical Center, Rotterdam, The Netherlands, The Generation R Study Group, Erasmus University Medical Center, Rotterdam, The Netherlands, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands, Netherlands Genomics Initiative (NGI)-sponsored Netherlands Consortium for Healthy Aging (NCHA), The Netherlands

David M. Evans

JHT, FR and DME are joint senior authors on this work.

Affiliations MRC Integrative Epidemiology Unit at the University of Bristol, Bristol, United Kingdom, University of Queensland Diamantina Institute, Translational Research Institute, Brisbane, Queensland, Australia

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: JPK CMG AGU GDS VWVJ JvdP CLAB TF JHT FR DME. Performed the experiments: JPK CMG NMW LO CJK SR KG EG BG BvdE JvdP MAH KC DLK FR JHT DME. Analyzed the data: JPK CMG KE BSP DHMH NJT NMW LO CJK LEW FMKW MCZ AH AGU SR KG EG BG BvdE JvdP MAH CLAB KC DLK MJE TF CO VWVJ JHT FR DME. Contributed reagents/materials/analysis tools: JPK CMG KE BSP DHMH NJT NMW LO SMR CJK LEW FMKW MCZ AH AGU SR KG EG BG BvdE JvdP MAH CLAB KC DLK MJE TF CO GDS VWVJ JHT FR DME. Wrote the paper: JPK CMG KE BSP DHMH NJT NMW LO SMR CJK LEW FMKW MCZ AH AGU SR KG EG BG BvdE JvdP MAH CLAB KC DLK MJE TF CO GDS VWVJ JHT FR DME.