Genome-wide association meta-analysis of human longevity identifies a novel locus conferring survival beyond 90 years of age

Qihua Tan9,11, Yoichiro Kamatani13, Anna M. Bennet14, Riin Tamm15,16, Stella Trompet4,5, Dan´ıel F. Guðbjartsson17, Friederike Flachsbart18, Giuseppina Rose20, Alexander Viktorin14, Krista Fischer15, Marianne Nygaard9,11, Heather J. Cordell8, Paolina Crocco20, Erik B. van den Akker1,21, Stefan Bo¨ hringer3, Quinta Helmer3, Christopher P. Nelson22,23, Gary I. Saunders24, Maris Alver15,16, Karen Andersen-Ranberg9, Marie E. Breen25,26, Ruud van der Breggen1,{ , Amke Caliebe19, Miriam Capri27, Elisa Cevenini27,{ , Joanna C. Collerton29,{ , Serena Dato20,{ , Karen Davies29,{ , Ian Ford30, Jutta Gampe32,{ , Paolo Garagnani27, Eco J.C. de Geus33,34, Jennifer Harrow24, Diana van Heemst5, Bastiaan T. Heijmans1,2, Femke-Anouska Heinsen18, Jouke-Jan Hottenga33, Albert Hofman2,6, Bernard Jeune9,{ , Palmi V. Jonsson35,36, Mark Lathrop13,34,38,{ , Doris Lechner37, Carmen Martin-Ruiz29, Susan E. Mcnerlan25,39,{ , Evelin Mihailov15,40, Alberto Montesanto20, Simon P. Mooijaart2,5, Anne Murphy25,{ , Ellen A. Nohr41,10, Lavinia Paternoster42, Iris Fernando Rivadeneira2,6,7, Owen A. Ross25,43, Stefano Salvioli27, Naveed Sattar44, Stefan Schreiber18,45,{ , Hreinn Stefa´nsson17, David J. Stott31, Henning Tiemeier2,6,46, Andre´ G. Uitterlinden2,6,7, Rudi G.J. Westendorp2,5,{ , Gonneke Willemsen33, Nilesh J. Pilar Galan47, Thorkild I.A. Sørensen48,49, Dorret I. Boomsma33, J. Wouter Jukema4,50, Irene Maeve Rea25,{ , Giuseppe Passarino20,{ , Anton J.M. de Craen5,{ , Kaare Christensen9,11,12,{ , Almut Nebel18,{ , Ka´ri Stefa´nsson17, Andres Metspalu15,16,40, Patrik Magnusson14, He´le`ne Blanche´13,{ , Lene Christiansen9,11, Thomas B.L. Kirkwood29,{ , Cornelia M. van Duijn2,6,

1Department of Molecular Epidemiology, 2Netherlands Consortium for Healthy Ageing, 3Department of Medical Statistics and Bioinformatics, 4Department of Cardiology and 5Department of Gerontology and Geriatrics, Leiden University Medical Center, Leiden 2300 RC, The Netherlands, 6Department of Epidemiology and 7Department of Internal Medicine, Erasmus Medical Center, Rotterdam 3000 CA, The Netherlands 8Institute of Genetic Medicine, International Centre for Life, Newcastle University, Newcastle upon Tyne NE1 3BZ, UK, 9Epidemiology, Institute of Public Health and 10Department of Gynecology and Obstetrics, Institute of Clinical Research, University of Southern Denmark, Odense C DK-5000, Denmark, 11Department of Clinical Genetics and 12Clinical Biochemistry and Pharmacology, Odense University Hospital, Odense C DK-5000, Denmark, 13Fondation Jean Dausset-CEPH, Paris 75010, France, 14Department of Medical Epidemiology and Biostatistics, Karolinska Institute, Stockholm SE-171 77, Sweden, 15Estonian Genome Center and 16Institute of Molecular and Cell Biology, University of Tartu, Tartu 51010, Estonia, 17Population Genomics,

To whom correspondence should be addressed at: Molecular Epidemiology Section, Leiden University Medical Center, Zone S5-P, PO Box 9600, 2300 RC Leiden, The Netherlands. Tel: +31 715269731/69730; Fax: +31 715268280; Email: p.slagboom@lumc.nl †On behalf of the GEHA consortium. A full list of consortium members is provided in Supplementary Material.

The Author 2014. Published by Oxford University Press. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/3.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited.

Genetics, Reykjav´ık 101, Iceland, 18Institute of Clinical Molecular Biology and 19Institute of Medical Informatics Statistics, Christian-Albrechts-University, Kiel 24105, Germany, 20Department of Biology, Ecology and Earth University of Calabria, Rende 87036, Italy, 21Delft Bioinformatics Lab, Delft University of Technology, Delft 2600