About the Authors

David Melzer

Contributed equally to this work with: David Melzer, John R. B. Perry, Dena Hernandez

Affiliations Department of Epidemiology and Public Health, Institute of Biomedical and Clinical Sciences, Peninsula College of Medicine and Dentistry, University of Exeter, Devon, United Kingdom, Genetics of Complex Traits, Institute of Biomedical and Clinical Sciences, Peninsula College of Medicine and Dentistry, University of Exeter, Devon, United Kingdom

John R. B. Perry

Contributed equally to this work with: David Melzer, John R. B. Perry, Dena Hernandez

Affiliation Genetics of Complex Traits, Institute of Biomedical and Clinical Sciences, Peninsula College of Medicine and Dentistry, University of Exeter, Devon, United Kingdom

Dena Hernandez

Contributed equally to this work with: David Melzer, John R. B. Perry, Dena Hernandez

Affiliation Laboratory of Neurogenetics, National Institute of Aging, Porter Neuroscience Research Center, Bethesda, Maryland, United States of America

Anna-Maria Corsi

Affiliation Tuscany Regional Health Agency, I.O.T. and Department of Medical and Surgical Critical Care, University of Florence, Florence, Italy

Kara Stevens

Affiliations Department of Epidemiology and Public Health, Institute of Biomedical and Clinical Sciences, Peninsula College of Medicine and Dentistry, University of Exeter, Devon, United Kingdom, Genetics of Complex Traits, Institute of Biomedical and Clinical Sciences, Peninsula College of Medicine and Dentistry, University of Exeter, Devon, United Kingdom

Ian Rafferty

Affiliation Laboratory of Neurogenetics, National Institute of Aging, Porter Neuroscience Research Center, Bethesda, Maryland, United States of America

Fulvio Lauretani

Affiliation Tuscany Regional Health Agency, I.O.T. and Department of Medical and Surgical Critical Care, University of Florence, Florence, Italy

Anna Murray

Affiliations Department of Epidemiology and Public Health, Institute of Biomedical and Clinical Sciences, Peninsula College of Medicine and Dentistry, University of Exeter, Devon, United Kingdom, Genetics of Complex Traits, Institute of Biomedical and Clinical Sciences, Peninsula College of Medicine and Dentistry, University of Exeter, Devon, United Kingdom

J. Raphael Gibbs

Affiliation Laboratory of Neurogenetics, National Institute of Aging, Porter Neuroscience Research Center, Bethesda, Maryland, United States of America

Giuseppe Paolisso

Affiliation Department of Geriatric Medicine and Metabolic Diseases, Second University of Naples, Naples, Italy

Sajjad Rafiq

Affiliations Department of Epidemiology and Public Health, Institute of Biomedical and Clinical Sciences, Peninsula College of Medicine and Dentistry, University of Exeter, Devon, United Kingdom, Genetics of Complex Traits, Institute of Biomedical and Clinical Sciences, Peninsula College of Medicine and Dentistry, University of Exeter, Devon, United Kingdom

Javier Simon-Sanchez

Affiliation Laboratory of Neurogenetics, National Institute of Aging, Porter Neuroscience Research Center, Bethesda, Maryland, United States of America

Hana Lango

Affiliation Genetics of Complex Traits, Institute of Biomedical and Clinical Sciences, Peninsula College of Medicine and Dentistry, University of Exeter, Devon, United Kingdom

Sonja Scholz

Affiliation Laboratory of Neurogenetics, National Institute of Aging, Porter Neuroscience Research Center, Bethesda, Maryland, United States of America

Michael N. Weedon

Affiliation Genetics of Complex Traits, Institute of Biomedical and Clinical Sciences, Peninsula College of Medicine and Dentistry, University of Exeter, Devon, United Kingdom

Sampath Arepalli

Affiliation Laboratory of Neurogenetics, National Institute of Aging, Porter Neuroscience Research Center, Bethesda, Maryland, United States of America

Neil Rice

Affiliations Department of Epidemiology and Public Health, Institute of Biomedical and Clinical Sciences, Peninsula College of Medicine and Dentistry, University of Exeter, Devon, United Kingdom, Genetics of Complex Traits, Institute of Biomedical and Clinical Sciences, Peninsula College of Medicine and Dentistry, University of Exeter, Devon, United Kingdom

Nicole Washecka

Affiliation Laboratory of Neurogenetics, National Institute of Aging, Porter Neuroscience Research Center, Bethesda, Maryland, United States of America

Alison Hurst

Affiliations Department of Epidemiology and Public Health, Institute of Biomedical and Clinical Sciences, Peninsula College of Medicine and Dentistry, University of Exeter, Devon, United Kingdom, Genetics of Complex Traits, Institute of Biomedical and Clinical Sciences, Peninsula College of Medicine and Dentistry, University of Exeter, Devon, United Kingdom

Angela Britton

Affiliation Laboratory of Neurogenetics, National Institute of Aging, Porter Neuroscience Research Center, Bethesda, Maryland, United States of America

William Henley

Affiliation School of Mathematics and Statistics, University of Plymouth, Plymouth, United Kingdom

Joyce van de Leemput

Affiliation Laboratory of Neurogenetics, National Institute of Aging, Porter Neuroscience Research Center, Bethesda, Maryland, United States of America

Rongling Li

Affiliation Department of Preventive Medicine and Center for Genomics and Bioinformatics, College of Medicine, University of Tennessee Health Science Center, Memphis, Tennessee, United States of America

Anne B. Newman

Affiliation University of Pittsburgh, Graduate School of Public Health, Departments of Epidemiology and Medicine, Pittsburgh, Pennsylvania, United States of America

Greg Tranah

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

Tamara Harris

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

Vijay Panicker

Affiliations Genetics of Complex Traits, Institute of Biomedical and Clinical Sciences, Peninsula College of Medicine and Dentistry, University of Exeter, Devon, United Kingdom, Henry Wellcome Laboratories for Integrative Neurosciences and Endocrinology, University of Bristol, Bristol, United Kingdom

Colin Dayan

Affiliation Henry Wellcome Laboratories for Integrative Neurosciences and Endocrinology, University of Bristol, Bristol, United Kingdom

Amanda Bennett

Affiliation Oxford Centre for Diabetes, Endocrinology and Metabolism, Headington, Oxford, United Kingdom

Mark I. McCarthy

Affiliations Oxford Centre for Diabetes, Endocrinology and Metabolism, Headington, Oxford, United Kingdom, The Wellcome Trust Centre for Human Genetics, Roosevelt Drive, Oxford, United Kingdom

Aimo Ruokonen

Affiliation Department of Clinical Chemistry, University of Oulu, Oulu, Finland

Marjo-Riitta Jarvelin

Affiliations Department of Public Health, Science, and General Practice, University of Oulu, Oulu, Finland, Department of Epidemiology and Public Health, Imperial College London, London, United Kingdom

Jack Guralnik

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

Stefania Bandinelli

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

Timothy M. Frayling

tim.frayling@pms.ac.uk

Affiliation Genetics of Complex Traits, Institute of Biomedical and Clinical Sciences, Peninsula College of Medicine and Dentistry, University of Exeter, Devon, United Kingdom

Andrew Singleton

Affiliation Laboratory of Neurogenetics, National Institute of Aging, Porter Neuroscience Research Center, Bethesda, Maryland, United States of America

Luigi Ferrucci

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

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: D Melzer, J Perry, D Hernandez, T Frayling, A Singleton, L Ferrucci. Performed the experiments: D Hernandez, I Rafferty, J Gibbs, J Simon-Sanchez, S Scholz, S Arepalli, N Washecka, A Britton, J van de Leemput, A Singleton. Analyzed the data: J Perry, K Stevens, A Murray, S Rafiq, H Lango, M Weedon, N Rice, A Hurst, W Hurst, A Singleton. Contributed reagents/materials/analysis tools: A Corsi, F Lauretani, G Paolisso, R Layan, A Newman, G Tranah, T Harris, V Panicker, C D A Bennett, M McCarthy, A Ruokonen, M Järvelin, J Guralnik, S Bandinelli, A Singleton, L Ferrucci. Wrote the paper: D Melzer, J Perry, D Hernandez, T. Frayling, L Ferrucci.