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NIH Public Access Author Manuscript Science. Author manuscript; available in PMC 2013 August 23. NIH-PA Author ManuscriptPublished NIH-PA Author Manuscript in final edited NIH-PA Author Manuscript form as: Science. 2013 June 21; 340(6139): 1467–1471. doi:10.1126/science.1235488. GWAS of 126,559 Individuals Identifies Genetic Variants Associated with Educational Attainment Cornelius A. Rietveld1,2, Sarah E. Medland3, Jaime Derringer4, Jian Yang5, Tõnu Esko6, Nicolas W. Martin3,7, Harm-Jan Westra8, Konstantin Shakhbazov5,9, Abdel Abdellaoui10, Arpana Agrawal11, Eva Albrecht12, Behrooz Z. Alizadeh13, Najaf Amin14, John Barnard15, Sebastian E. Baumeister16, Kelly S. Benke17, Lawrence F. Bielak18, Jeffrey A. Boatman19, Patricia A. Boyle20, Gail Davies21, Christiaan de Leeuw22, Niina Eklund24,25, Daniel S. Evans26, Rudolf Ferhmann8, Krista Fischer6, Christian Gieger12, Håkon K. Gjessing27, Sara Hägg28,29,30, Jennifer R. Harris27, Caroline Hayward31, Christina Holzapfel32,33, Carla A. Ibrahim-Verbaas14,34, Erik Ingelsson28,29,30, Bo Jacobsson27,35, Peter K. Joshi36, Astanand Jugessur27, Marika Kaakinen37,38, Stavroula Kanoni39, Juha Karjalainen8, Ivana Kolcic40, Kati Kristiansson24,25, Zoltán Kutalik41,42, Jari Lahti43, Sang H. Lee3, Peng Lin11, Penelope A. Lind3, Yongmei Liu44, Kurt Lohman45, Marisa Loitfelder46, George McMahon47, Pedro Marques Vidal48, Osorio Meirelles49, Lili Milani6, Ronny Myhre27, Marja-Liisa Nuotio24,25, Christopher J. Oldmeadow50, Katja E. Petrovic51, Wouter J. Peyrot52, Ozren Polašek40, Lydia Quaye53, Eva Reinmaa6, John P. Rice11, Thais S. Rizzi22, Helena Schmidt54, Reinhold Schmidt46, Albert V. Smith55,56, Jennifer A. Smith18, Toshiko Tanaka49, Antonio Terracciano49,57, Matthijs J.H.M. van der Loos1,2, Veronique Vitart31, Henry Völzke16, Jürgen Wellmann58, Lei Yu20, Wei Zhao18, Jüri Allik59, John R. Attia50, Stefania Bandinelli60, François Bastardot61, Jonathan Beauchamp62, David A. Bennett20, Klaus Berger58, Laura J. Bierut11, Dorret I. Boomsma10, Ute Bültmann63, Harry Campbell36, Christopher F. Chabris64, Lynn Cherkas53, Mina K. Chung15, Francesco Cucca65,66, Mariza de Andrade67, Philip L. De Jager68, Jan-Emmanuel De Neve69,70, Ian J. Deary21,71, George V. Dedoussis72, Panos Deloukas39, Maria Dimitriou72, Gudny Eiriksdottir55, Martin F. Elderson73, Johan G. Eriksson74,75,76,77, David M. Evans78, Jessica D. Faul79, Luigi Ferrucci49, Melissa E. Garcia49, Henrik Grönberg30, Vilmundur Gudnason55,56, Per Hall30, Juliette M. Harris53, Tamara B. Harris49, Nicholas D. Hastie31, Andrew C. Heath80, Dena G. Hernandez49, Wolfgang Hoffmann16, Adriaan Hofman81, Rolf Holle83, Elizabeth G. Holliday50, Jouke-Jan Hottenga10, William G. Iacono82, Thomas Illig33,84, Marjo-Riitta Järvelin37,38,85,86,87, Mika Kähönen88, Jaakko Kaprio24,89,90, Robert M. Kirkpatrick82, Matthew Kowgier91, Antti Latvala89,90, Lenore J. Launer49, Debbie A. Lawlor78, Terho Lehtimäki92, Jingmei Li93, Paul Lichtenstein30, Peter Lichtner94, David C. Liewald21, Pamela A. Madden11, Patrik K. E. Magnusson30, Tomi E. Mäkinen95, Marco Masala65, Matt McGue82, Andres Metspalu6, Andreas Mielck83, Michael B. Miller82, Grant W. Montgomery3, Sutapa Mukherjee96,97,98, Dale R. Nyholt3, Ben A. Oostra14, Lyle J. Palmer91, Aarno Palotie24,39,99, Brenda Penninx52, Markus Perola24,25,6, Patricia A. Peyser18, Martin Preisig61, Katri Räikkönen43, Olli T. Raitakari100,101, Anu Realo59, Susan M. Ring47, Samuli †Corresponding author. [email protected] (D.J.B.); [email protected] (D.C.); [email protected] (P.D.K.); [email protected] (P.M.V.). *These authors contributed equally to this work. Supplementary Materials www.sciencemag.org/cgi/content/full/science.1235488/DC1 Materials and Methods Supplementary Text Figs. S1 to S22 Tables S1 to S27 References (23–175) Rietveld et al. Page 2 Ripatti24,25,39, Fernando Rivadeneira2,102, Igor Rudan36, Aldo Rustichini103, Veikko Salomaa104, Antti-Pekka Sarin24, David Schlessinger49, Rodney J. Scott50, Harold 13 78,105 21,106 107 24,25 NIH-PA Author ManuscriptSnieder NIH-PA Author Manuscript, Beate NIH-PA Author Manuscript St Pourcain , John M. Starr , Jae Hoon Sul , Ida Surakka , Rauli Svento108, Alexander Teumer109, The LifeLines Cohort Study, Henning Tiemeier2,110, Frank JAan Rooij2, David R. Van Wagoner15, Erkki Vartiainen111, Jorma Viikari112, Peter Vollenweider61, Judith M. Vonk13, Gérard Waeber61, David R. Weir79, H.-Erich Wichmann113,114,115, Elisabeth Widen24, Gonneke Willemsen10, James F. Wilson36, Alan F. Wright31, Dalton Conley116, George Davey-Smith78, Lude Franke8, Patrick J. F. Groenen121, Albert Hofman2, Magnus Johannesson122, Sharon L.R. Kardia18, Robert F. Krueger82, David Laibson117, Nicholas G. Martin3, Michelle N. Meyer118,119, Danielle Posthuma22,110,120, A. Roy Thurik1,123,124, Nicholas J. Timpson78, André G. Uitterlinden2,102, Cornelia M. van Duijn14,125, Peter M. Visscher3,5,9,†,*, Daniel J. Benjamin126,*,†, David Cesarini127,128,129,*,†, and Philipp D. Koellinger1,2,*,† 1Department of Applied Economics, Erasmus School of Economics, Erasmus University Rotterdam, 3000 DR Rotterdam, The Netherlands 2Department of Epidemiology, Erasmus Medical Center, Rotterdam 3000 CA, The Netherlands 3Queensland Institute of Medical Research, 300 Herston Road, Brisbane, Queensland 4006, Australia 4Institute for Behavioral Genetics, University of Colorado Boulder, Boulder, CO 80309–0447, USA 5University of Queensland Diamantina Institute, The University of Queensland, Princess Alexandra Hospital, Brisbane, Queensland 4102, Australia 6Estonian Genome Center, University of Tartu, Tartu 51010, Estonia 7School of Psychology, University of Queensland, Brisbane, Queensland 4072, Australia 8Department of Genetics, University Medical Center Groningen, University of Groningen, 9713 GZ Groningen, The Netherlands 9Queensland Brain Institute, The University of Queensland, Brisbane, Queensland 4072, Australia 10Department of Biological Psychology, VU University Amsterdam, 1081 BT Amsterdam, The Netherlands 11Department of Psychiatry, Washington University School of Medicine, St. Louis, MO 63110, USA 12Institute of Genetic Epidemiology, Helmholtz Zentrum München, German Research Center for Environmental Health, 85764 Neuherberg, Germany 13Department of Epidemiology, University Medical Center Groningen, University of Groningen, 9700 RB Groningen, The Netherlands 14Genetic Epidemiology Unit, Department of Epidemiology, Erasmus Medical Center, Rotterdam 3000 CA, the Netherlands 15Heart and Vascular and Lerner Research Institutes, Cleveland Clinic, Cleveland, OH 44195, USA 16Institute for Community Medicine, University Medicine Greifswald, Greifswald 17489, Germany 17Samuel Lunenfeld Research Institute, Mount Sinai Hospital, University of Toronto, Toronto, Ontario M5G 1X5, Canada 18Department of Epidemiology, School of Public Health, University of Michigan, Ann Arbor, MI 48109–2029, USA 19Division of Biostatistics, University of Minnesota, Minneapolis, MN 55455, USA 20Rush University Medical Center, Rush Alzheimer’s Disease Center, Chicago, IL 60612, USA 21Centre for Cognitive Aging and Cognitive Epidemiology, The University of Edinburgh, Edinburgh EH8 9JZ, Scotland, UK 22Department of Functional Genomics, VU University Amsterdam and VU Medical Center, 1081 HV Amsterdam, The Netherlands 23Machine Learning Group, Intelligent Systems, Institute for Computing and Information Sciences, Faculty of Science, Radboud University Nijmegen, 6500 GL Nijmegen, The Netherlands 24Institute for Molecular Medicine Finland, University of Helsinki, Helsinki 00014, Finland 25Public Health Genomics Unit, Department of Chronic Disease Prevention, The National Institute for Health and Welfare, Helsinki 00014, Finland 26California Pacific Medical Center Research Institute, San Francisco, CA 94107–1728, USA 27Department of Genes and Environment, Division of Epidemiology, Norwegian Institute of Public Health, Nydalen, N-0403 Oslo, Norway 28Molecular Epidemiology, Department of Medical Sciences, Uppsala University, 751 85 Uppsala, Sweden 29Science for Life Laboratory, Department of Medical Biochemistry and Microbiology, Uppsala University, 751 23 Uppsala, Sweden 30Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, 171 77 Stockholm, Sweden 31Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh EH4 2XU, UK Science. Author manuscript; available in PMC 2013 August 23. Rietveld et al. Page 3 32Else Kroener-Fresenius-Centre for Nutritional Medicine, Technische Universität München, NIH-PA Author Manuscript81675 NIH-PA Author Manuscript Munich, Germany NIH-PA Author Manuscript 33Research Unit of Molecular Epidemiology, Helmholtz Zentrum München, German Research Center for Environmental Health, 85764 Neuherberg, Germany 34Department of Neurology, Erasmus Medical Center, Rotterdam 3000 CA, The Netherlands 35Department of Obstetrics and Gynecology, Institute of Public Health, Sahlgrenska Academy, Sahgrenska University Hospital, Gothenburg, 413 45, Sweden 36Centre for Population Health Sciences, The University of Edinburgh, Edinburgh EH8 9AG, UK 37Institute of Health Sciences, University of Oulu, Oulu 90014, Finland 38Biocenter Oulu, University of Oulu, Oulu 90014, Finland 39Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton CB10 1SA, UK 40Faculty of Medicine, University of Split, 21000 Split, Croatia 41Department of Medical Genetics, University of Lausanne, 1005 Lausanne, Switzerland 42Swiss Institute of Bioinformatics, 1015 Lausanne, Switzerland 43Institute of Behavioral Sciences, University of Helsinki,