About the Authors

Valentina Escott-Price

Contributed equally to this work with: Valentina Escott-Price, Céline Bellenguez, Li-San Wang, Seung-Hoan Choi, Sudha Seshadri, Gerard D. Schellenberg, Philippe Amouyel, Julie Williams

Affiliation Institute of Psychological Medicine and Clinical Neurosciences, MRC Centre for Neuropsychiatric Genetics & Genomics, Cardiff University, Cardiff, United Kingdom

Céline Bellenguez

Contributed equally to this work with: Valentina Escott-Price, Céline Bellenguez, Li-San Wang, Seung-Hoan Choi, Sudha Seshadri, Gerard D. Schellenberg, Philippe Amouyel, Julie Williams

Affiliations Inserm U744, Lille, France, Université Lille 2, Lille, France, Institut Pasteur de Lille, Lille, France

Li-San Wang

Contributed equally to this work with: Valentina Escott-Price, Céline Bellenguez, Li-San Wang, Seung-Hoan Choi, Sudha Seshadri, Gerard D. Schellenberg, Philippe Amouyel, Julie Williams

Affiliation Department of Pathology and Laboratory Medicine, University of Pennsylvania Perelman School of Medicine, Philadelphia, Pennsylvania, United States of America

Seung-Hoan Choi

Contributed equally to this work with: Valentina Escott-Price, Céline Bellenguez, Li-San Wang, Seung-Hoan Choi, Sudha Seshadri, Gerard D. Schellenberg, Philippe Amouyel, Julie Williams

Affiliation Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts, United States of America

Denise Harold

Affiliation Institute of Psychological Medicine and Clinical Neurosciences, MRC Centre for Neuropsychiatric Genetics & Genomics, Cardiff University, Cardiff, United Kingdom

Lesley Jones

Affiliation Institute of Psychological Medicine and Clinical Neurosciences, MRC Centre for Neuropsychiatric Genetics & Genomics, Cardiff University, Cardiff, United Kingdom

Peter Holmans

Affiliation Institute of Psychological Medicine and Clinical Neurosciences, MRC Centre for Neuropsychiatric Genetics & Genomics, Cardiff University, Cardiff, United Kingdom

Amy Gerrish

Affiliation Institute of Psychological Medicine and Clinical Neurosciences, MRC Centre for Neuropsychiatric Genetics & Genomics, Cardiff University, Cardiff, United Kingdom

Alexey Vedernikov

Affiliation Institute of Psychological Medicine and Clinical Neurosciences, MRC Centre for Neuropsychiatric Genetics & Genomics, Cardiff University, Cardiff, United Kingdom

Alexander Richards

Affiliation Institute of Psychological Medicine and Clinical Neurosciences, MRC Centre for Neuropsychiatric Genetics & Genomics, Cardiff University, Cardiff, United Kingdom

Anita L. DeStefano

Affiliation Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts, United States of America

Jean-Charles Lambert

Affiliations Inserm U744, Lille, France, Université Lille 2, Lille, France, Institut Pasteur de Lille, Lille, France

Carla A. Ibrahim-Verbaas

Affiliation Department of Epidemiology and Neurology, Erasmus MC University Medical Center, Rotterdam, the Netherlands

Adam C. Naj

Affiliation Department of Biostatistics and Epidemiology and Center for Clinical Epidemiology and Biostatistics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, United States of America

Rebecca Sims

Affiliation Institute of Psychological Medicine and Clinical Neurosciences, MRC Centre for Neuropsychiatric Genetics & Genomics, Cardiff University, Cardiff, United Kingdom

Gyungah Jun

Affiliations Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts, United States of America, Department of Medicine (Biomedical Genetics), Boston University School of Medicine, Boston, Massachusetts, United States of America, Department of Ophthalmology, Boston University School of Medicine, Boston, Massachusetts, United States of America

Joshua C. Bis

Affiliation Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, Washington, United States of America

Gary W. Beecham

Affiliations The John P. Hussman Institute for Human Genomics, University of Miami, Miami, Florida, United States of America, Dr. John T. Macdonald Foundation Department of Human Genetics, University of Miami, Miami, Florida, United States of America

Benjamin Grenier-Boley

Affiliations Inserm U744, Lille, France, Université Lille 2, Lille, France, Institut Pasteur de Lille, Lille, France

Giancarlo Russo

Affiliation Functional Genomics Center Zurich, ETH/University of Zurich, Zurich, Switzerland

Tricia A. Thornton-Wells

Affiliation Department of Molecular Physiology and Biophysics, Vanderbilt University, Nashville, Tennessee, United States of America

Nicola Denning

Affiliation Institute of Psychological Medicine and Clinical Neurosciences, MRC Centre for Neuropsychiatric Genetics & Genomics, Cardiff University, Cardiff, United Kingdom

Albert V. Smith

Affiliations University of Iceland, Faculty of Medicine, Reykjavik, Iceland, Icelandic Heart Association, Kopavogur, Iceland

Vincent Chouraki

Affiliations Inserm U744, Lille, France, Université Lille 2, Lille, France, Institut Pasteur de Lille, Lille, France, Department of Neurology, Boston University School of Medicine, Boston, Massachusetts, United States of America

Charlene Thomas

Affiliation Institute of Psychological Medicine and Clinical Neurosciences, MRC Centre for Neuropsychiatric Genetics & Genomics, Cardiff University, Cardiff, United Kingdom

M. Arfan Ikram

Affiliations Departments of Epidemiology, Neurology and Radiology, Erasmus MC University Medical Center, Rotterdam, the Netherlands, Netherlands Consortium for Healthy Aging, Leiden, The Netherlands

Diana Zelenika

Affiliation Centre National de Genotypage, Institut Genomique, Commissariat à l'énergie Atomique, Evry, France

Badri N. Vardarajan

Affiliations Department of Medicine (Biomedical Genetics), Boston University School of Medicine, Boston, Massachusetts, United States of America, Taub Institute on Alzheimer's Disease and the Aging Brain, Department of Neurology, Columbia University New York, New York, United States of America, Gertrude H. Sergievsky Center, Department of Neurology, Columbia University, New York, New York, United States of America

Yoichiro Kamatani

Affiliation Fondation Jean Dausset- CEPH, Paris, France

Chiao-Feng Lin

Affiliation Department of Pathology and Laboratory Medicine, University of Pennsylvania Perelman School of Medicine, Philadelphia, Pennsylvania, United States of America

Helena Schmidt

Affiliation Institute for Molecular Biology and Biochemistry, Medical University of Graz, Graz, Austria

Brian Kunkle

Affiliation The John P. Hussman Institute for Human Genomics, University of Miami, Miami, Florida, United States of America

Melanie L. Dunstan

Affiliation Institute of Psychological Medicine and Clinical Neurosciences, MRC Centre for Neuropsychiatric Genetics & Genomics, Cardiff University, Cardiff, United Kingdom

Maria Vronskaya

Affiliation Institute of Psychological Medicine and Clinical Neurosciences, MRC Centre for Neuropsychiatric Genetics & Genomics, Cardiff University, Cardiff, United Kingdom

the United Kingdom Brain Expression Consortium

Membership of the UK Brain Expression consortium is provided in Materials S1.

Affiliation Reta Lila Weston Research Laboratories, Department of Molecular Neuroscience, UCL Institute of Neurology, London, United Kingdom

Andrew D. Johnson

Affiliation NHLBI Cardiovascular Epidemiology and Human Genomics Branch, The Framingham Heart Study, Framingham, Massachusetts, United States of America

Agustin Ruiz

Affiliation Memory Clinic of Fundació ACE. Institut Català de Neurociències Aplicades, Barcelona, Spain

Marie-Thérèse Bihoreau

Affiliation Centre National de Genotypage, Institut Genomique, Commissariat à l'énergie Atomique, Evry, France

Christiane Reitz

Affiliations Taub Institute on Alzheimer's Disease and the Aging Brain, Department of Neurology, Columbia University New York, New York, United States of America, Gertrude H. Sergievsky Center, Department of Neurology, Columbia University, New York, New York, United States of America

Florence Pasquier

Affiliations Université Lille 2, Lille, France, CNR-MAJ, Centre Hospitalier Régional Universitaire de Lille, Lille, France

Paul Hollingworth

Affiliation Institute of Psychological Medicine and Clinical Neurosciences, MRC Centre for Neuropsychiatric Genetics & Genomics, Cardiff University, Cardiff, United Kingdom

Olivier Hanon

Affiliation University Paris Descartes, Sorbonne Paris V, Broca Hospital, Geriatrics department, Paris, France

Annette L. Fitzpatrick

Affiliations Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, Washington, United States of America, Departments of Epidemiology and Global Health, University of Washington, Seattle, Washington, United States of America

Joseph D. Buxbaum

Affiliations Department of Neuroscience, Mount Sinai School of Medicine, New York, New York, United States of America, Department of Psychiatry, Mount Sinai School of Medicine, New York, New York, United States of America, Departments of Genetics and Genomic Sciences, Mount Sinai School of Medicine, New York, New York, United States of America

Dominique Campion

Affiliation CNR-MAJ, Inserm U1079, Rouen University Hospital, 76031 France, Rouen, France

Paul K. Crane

Affiliation Department of Medicine, University of Washington, Seattle, Washington, United States of America

Clinton Baldwin

Affiliation Department of Medicine (Biomedical Genetics), Boston University School of Medicine, Boston, Massachusetts, United States of America

Tim Becker

Affiliation German Center for Neurodegenerative Diseases (DZNE), Bonn, and Institute for Medical Biometry, Informatics and Epidemiology, University of Bonn, Bonn, Germany

Vilmundur Gudnason

Affiliations University of Iceland, Faculty of Medicine, Reykjavik, Iceland, Icelandic Heart Association, Kopavogur, Iceland

Carlos Cruchaga

Affiliation Department of Psychiatry and Hope Center Program on Protein Aggregation and Neurodegeneration, Washington University School of Medicine, St. Louis, Missouri, United States of America

David Craig

Affiliation Ageing Group, Centre for Public Health, School of Medicine, Dentistry and Biomedical Sciences, Queen's University Belfast, Belfast, United Kingdom

Najaf Amin

Affiliation Department of Epidemiology, Erasmus MC University Medical Center, Rotterdam, the Netherlands

Claudine Berr

Affiliation INSERM U1061, Faculty of Medicine, Hôpital La Colombière, Montpellier, France

Oscar L. Lopez

Affiliation Departments of Neurology, University of Pittsburgh School of Medicine, Pittsburgh, Pennsylvania, United States of America

Philip L. De Jager

Affiliations Program in Translational NeuroPsychiatric Genomics, Institute for the Neurosciences, Department of Neurology & Psychiatry, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts, United States of America, Program in Medical and Population Genetics, Broad Institute, Boston, Massachusetts, United States of America

Vincent Deramecourt

Affiliations Université Lille 2, Lille, France, CNR-MAJ, Centre Hospitalier Régional Universitaire de Lille, Lille, France

Janet A. Johnston

Affiliation Ageing Group, Centre for Public Health, School of Medicine, Dentistry and Biomedical Sciences, Queen's University Belfast, Belfast, United Kingdom

Denis Evans

Affiliation Rush Institute for Healthy Aging, Department of Internal Medicine, Rush University Medical Center, Chicago, Illinois, United States of America

Simon Lovestone

Affiliation King's College London, Institute of Psychiatry, Department of Neuroscience, De Crespigny Park, Denmark Hill, London, United Kingom

Luc Letenneur

Affiliation Inserm U897, Victor Segalen University, F-33076, Bordeaux, France

Isabel Hernández

Affiliation Memory Clinic of Fundació ACE. Institut Català de Neurociències Aplicades, Barcelona, Spain

David C. Rubinsztein

Affiliation Cambridge Institute for Medical Research, University of Cambridge, Cambridge, United Kingdom

Gudny Eiriksdottir

Affiliation Icelandic Heart Association, Kopavogur, Iceland

Kristel Sleegers

Affiliations Neurodegenerative Brain Diseases Group, Department of Molecular Genetics, VIB, Antwerp, Belgium, Laboratory of Neurogenetics, Institute Born-Bunge, University of Antwerp, Antwerp, Belgium

Alison M. Goate

Affiliation Department of Psychiatry and Hope Center Program on Protein Aggregation and Neurodegeneration, Washington University School of Medicine, St. Louis, Missouri, United States of America

Nathalie Fiévet

Affiliations Inserm U744, Lille, France, Institut Pasteur de Lille, Lille, France

Matthew J. Huentelman

Affiliation Neurogenomics Division, Translational Genomics Research Institute, Phoenix, Arizona, United States of America

Michael Gill

Affiliation Discipline of Psychiatry, Trinity College, Dublin, Ireland

Kristelle Brown

Affiliation Institute of Genetics, Queen's Medical Centre, University of Nottingham, Nottingham, United Kingdom

M. Ilyas Kamboh

Affiliations Department of Human Genetics, University of Pittsburgh, Pittsburgh, Pennsylvania, United States of America, Alzheimer's Disease Research Center, University of Pittsburgh, Pittsburgh, Pennsylvania, United States of America

Lina Keller

Affiliation Aging Reasearch Center, Department of Neurobiology, Care Sciences and Society, Karolinska Institutet and Stockholm University, Stockholm, Sweden

Pascale Barberger-Gateau

Affiliation King's College London, Institute of Psychiatry, Department of Neuroscience, De Crespigny Park, Denmark Hill, London, United Kingom

Bernadette McGuinness

Affiliation Ageing Group, Centre for Public Health, School of Medicine, Dentistry and Biomedical Sciences, Queen's University Belfast, Belfast, United Kingdom

Eric B. Larson

Affiliations Department of Medicine, University of Washington, Seattle, Washington, United States of America, Group Health Research Institute, Group Health, Seattle, Washington, United States of America

Amanda J. Myers

Affiliation Department of Psychiatry and Behavioral Sciences, Miller School of Medicine, University of Miami, Miami, Florida, United States of America

Carole Dufouil

Affiliation Inserm U897, Victor Segalen University, F-33076, Bordeaux, France

Stephen Todd

Affiliation Ageing Group, Centre for Public Health, School of Medicine, Dentistry and Biomedical Sciences, Queen's University Belfast, Belfast, United Kingdom

David Wallon

Affiliation CNR-MAJ, Inserm U1079, Rouen University Hospital, 76031 France, Rouen, France

Seth Love

Affiliation University of Bristol Institute of Clinical Neurosciences, School of Clinical Sciences, Frenchay Hospital, Bristol, United Kingdom

Ekaterina Rogaeva

Affiliation Tanz Centre for Research in Neurodegenerative Disease, University of Toronto, Toronto, Ontario, Canada

John Gallacher

Affiliation Institute of Primary Care and Public Health, Cardiff University, Neuadd Meirionnydd, University Hospital of Wales, Heath Park, Cardiff, United Kingdom

Peter St George-Hyslop

Affiliations Tanz Centre for Research in Neurodegenerative Disease, University of Toronto, Toronto, Ontario, Canada, Cambridge Institute for Medical Research and Department of Clinical Neurosciences, University of Cambridge, Cambridge, United Kingdom

Jordi Clarimon

Affiliations Neurology Department. IIB Sant Pau. Sant Pau Hospital. Universitat Autònoma de Barcelona, Barcelona, Spain, Center for Networker Biomedical Research in Neurodegenerative Diseases (CIBERNED), Barcelona, Spain

Alberto Lleo

Affiliations Neurology Department. IIB Sant Pau. Sant Pau Hospital. Universitat Autònoma de Barcelona, Barcelona, Spain, Center for Networker Biomedical Research in Neurodegenerative Diseases (CIBERNED), Barcelona, Spain

Anthony Bayer

Affiliation Institute of Primary Care and Public Health, Cardiff University, Neuadd Meirionnydd, University Hospital of Wales, Heath Park, Cardiff, United Kingdom

Debby W. Tsuang

Affiliation Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle, Washington, United States of America

Lei Yu

Affiliation Department of Neurological Sciences, Rush University Medical Center, Chicago, Illinois, United States of America

Magda Tsolaki

Affiliation 3rd Department of Neurology, Aristotle University of Thessaloniki, Thessaloniki, Greece

Paola Bossù

Affiliation Clinical and Behavioral Neurology, Fondazione Santa Lucia, Roma, Italy

Gianfranco Spalletta

Affiliation Clinical and Behavioral Neurology, Fondazione Santa Lucia, Roma, Italy

Petra Proitsi

Affiliation King's College London, Institute of Psychiatry, Department of Neuroscience, De Crespigny Park, Denmark Hill, London, United Kingom

John Collinge

Affiliation MRC Prion Unit, Department of Neurodegenerative Disease, UCL Institute of Neurology, London, United Kingdom

Sandro Sorbi

Affiliations NEUROFARBA Department of Neuroscience, Psychology, Drug Research and Child Health, University of Florence, Florence, Italy, Centro di Ricerca, Trasferimento e Alta Formazione DENOTHE, University of Florence, Florence, Italy

Florentino Sanchez Garcia

Affiliation Department of Immunology, Hospital Universitario Dr. Negrin, Las Palmas de Gran Canaria, Spain

Nick C. Fox

Affiliation Dementia Research Center, Department of Neurodegenerative Disease, UCL Institute of Neurology, London, United Kingdom

John Hardy

Affiliation Department of Molecular Neuroscience and Reta Lilla Weston Laboratories, Institute of Neurology, London, United Kingdom

Maria Candida Deniz Naranjo

Affiliation Department of Immunology, Hospital Universitario Dr. Negrin, Las Palmas de Gran Canaria, Spain

Paolo Bosco

Affiliation IRCCS Associazione Oasi Maria SS, Troina, Italy

Robert Clarke

Affiliation Oxford Healthy Aging Project (OHAP), Clinical Trial Service Unit, University of Oxford, Oxford, United Kingdom

Carol Brayne

Affiliation Cognitive Function and Ageing Study (CFAS), Institute of Public Health, University of Cambridge, Cambridge, United Kingdom

Daniela Galimberti

Affiliation University of Milan, Fondazione Cà Granda, IRCCS Ospedale Policlinico, Milan, Italy

Elio Scarpini

Affiliation University of Milan, Fondazione Cà Granda, IRCCS Ospedale Policlinico, Milan, Italy

Ubaldo Bonuccelli

Affiliation Neurological Clinic, University of Pisa, Pisa, Italy

Michelangelo Mancuso

Affiliation Neurological Clinic, University of Pisa, Pisa, Italy

Gabriele Siciliano

Affiliation Neurological Clinic, University of Pisa, Pisa, Italy

Susanne Moebus

Affiliation Urban Epidemiology, Institute for Medical Informatics, Biometry and Epidemiology, University Hospital Essen, University Duisburg-Essen, Essen, Germany

Patrizia Mecocci

Affiliation Section of Gerontology and Geriatrics, Department of Clinical and Experimental Medicine, University of Perugia, Perugia, Italy

Maria Del Zompo

Affiliation Section of Neuroscience and Clinical Pharmacology, Department of Biomedical Sciences, University of Cagliari, Cagliari, Italy

Wolfgang Maier

Affiliation Department of Psychiatry and Psychotherapy, University of Bonn, Germany and German Center for Neurodegenerative Diseases (DZNE, Bonn), Bonn, Germany

Harald Hampel

Affiliations Department of Psychiatry, University of Frankfurt am Main, Frankfurt am Main, Germany (H.H.), Department of Psychiatry, Ludwig-Maximilians University, Munich, Germany

Alberto Pilotto

Affiliation Gerontology and Geriatrics Research Laboratory, I.R.C.C.S. Casa Sollievo della Sofferenza, San Giovanni Rotondo (FG), Italy

Ana Frank-García

Affiliations Centro de Biología Molecular Severo Ochoa (CSIC-UAM); Madrid, Spain, Centro de Investigación Biomédica en Red sobre Enfermedades Neurodegenerativas (CIBERNED), Madrid, Spain, Instituto de Investigación Sanitaria “Hospital la Paz” (IdIPaz), Madrid, Spain

Francesco Panza

Affiliation Department of Geriatrics,Center for Aging Brain,University of Bari, Bari, Italy

Vincenzo Solfrizzi

Affiliation Department of Geriatrics,Center for Aging Brain,University of Bari, Bari, Italy

Paolo Caffarra

Affiliations Department of Neuroscience-University of Parma, Parma, Italy, Center for Cognitive Disorders AUSL, Parma, Italy

Benedetta Nacmias

Affiliations NEUROFARBA Department of Neuroscience, Psychology, Drug Research and Child Health, University of Florence, Florence, Italy, Centro di Ricerca, Trasferimento e Alta Formazione DENOTHE, University of Florence, Florence, Italy

William Perry

Affiliations The John P. Hussman Institute for Human Genomics, University of Miami, Miami, Florida, United States of America, Dr. John T. Macdonald Foundation Department of Human Genetics, University of Miami, Miami, Florida, United States of America

Manuel Mayhaus

Affiliation Department Of Psychiatry, University Hospital, Saarland, Germany

Lars Lannfelt

Affiliation Department of Public Health/Geriatrics, Uppsala University, Uppsala, Sweden

Hakon Hakonarson

Affiliation Center for Applied Genomics, Children's Hospital of Philadelphia, Philadelphia, Pennsylvania, United States of America

Sabrina Pichler

Affiliation Department Of Psychiatry, University Hospital, Saarland, Germany

Minerva M. Carrasquillo

Affiliation Department of Neuroscience, Mayo Clinic, Jacksonville, Florida, United States of America

Martin Ingelsson

Affiliation Department of Public Health/Geriatrics, Uppsala University, Uppsala, Sweden

Duane Beekly

Affiliation National Alzheimer's Coordinating Center, University of Washington, Seattle, Washington, United States of America

Victoria Alvarez

Affiliation Genetica molecular-Huca-Oviedo, Oviedo, Spain

Fanggeng Zou

Affiliation Department of Neuroscience, Mayo Clinic, Jacksonville, Florida, United States of America

Otto Valladares

Affiliation Department of Pathology and Laboratory Medicine, University of Pennsylvania Perelman School of Medicine, Philadelphia, Pennsylvania, United States of America

Steven G. Younkin

Affiliation Department of Neuroscience, Mayo Clinic, Jacksonville, Florida, United States of America

Eliecer Coto

Affiliation Genetica molecular-Huca-Oviedo, Oviedo, Spain

Kara L. Hamilton-Nelson

Affiliation The John P. Hussman Institute for Human Genomics, University of Miami, Miami, Florida, United States of America

Wei Gu

Affiliation Department of Psychiatry, University Hospital, Saarland, Germany

Cristina Razquin

Affiliation Neurogenetics Laboratory, Division of Neurosciences, Center for Applied Medical Research, University of Navarra School of Medicine, Pamplona, Spain

Pau Pastor

Affiliations Neurogenetics Laboratory, Division of Neurosciences, Center for Applied Medical Research, University of Navarra School of Medicine, Pamplona, Spain, CIBERNED, Centro de Investigación Biomédica en Red de Enfermedades Neurodegenerativas, Instituto de Salud Carlos III, Madrid, Spain

Ignacio Mateo

Affiliation Neurology Service and CIBERNED, "Marqués de Valdecilla" University Hospital (University of Cantabria and IFIMAV), Santander, Spain

Michael J. Owen

Affiliation Institute of Psychological Medicine and Clinical Neurosciences, MRC Centre for Neuropsychiatric Genetics & Genomics, Cardiff University, Cardiff, United Kingdom

Kelley M. Faber

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

Palmi V. Jonsson

Affiliations University of Iceland, Faculty of Medicine, Reykjavik, Iceland, Landspitali University Hospital, Reykjavik, Iceland

Onofre Combarros

Affiliation Neurology Service and CIBERNED, "Marqués de Valdecilla" University Hospital (University of Cantabria and IFIMAV), Santander, Spain

Michael C. O'Donovan

Affiliation Institute of Psychological Medicine and Clinical Neurosciences, MRC Centre for Neuropsychiatric Genetics & Genomics, Cardiff University, Cardiff, United Kingdom

Laura B. Cantwell

Affiliation Department of Pathology and Laboratory Medicine, University of Pennsylvania Perelman School of Medicine, Philadelphia, Pennsylvania, United States of America

Hilkka Soininen

Affiliations Institute of Clinical Medicine - Neurology, University of Eastern Finland, Kuopio, Finland, Department of Neurology, Kuopio University Hospital, Kuopio, Finland

Deborah Blacker

Affiliations Department of Epidemiology, Harvard School of Public Health, Boston, Massachusetts, United States of America, Department of Psychiatry, Massachusetts General Hospital/Harvard Medical School, Boston, Massachusetts, United States of America

Simon Mead

Affiliation MRC Prion Unit, Department of Neurodegenerative Disease, UCL Institute of Neurology, London, United Kingdom

Thomas H. Mosley Jr

Affiliation Department of Medicine (Geriatrics), University of Mississippi Medical Center, Jackson, Mississippi, United States of America

David A. Bennett

Affiliations Department of Neurological Sciences, Rush University Medical Center, Chicago, Illinois, United States of America, Rush Alzheimer's Disease Center, Rush University Medical Center, Chicago, Illinois, United States of America

Tamara B. Harris

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

Laura Fratiglioni

Affiliations Aging Research Center, Department Neurobiology, Care Sciences and Society, Karolinska Institutet and Stockholm University, Stockholm, Sweden, Department Geriatric Medicine, Genetics Unit, Karolinska University Hospital Huddinge, Stockholm, Sweden

Clive Holmes

Affiliation Division of Clinical Neurosciences, School of Medicine, University of Southampton, Southampton, United Kingdom

Renee F. A. G. de Bruijn

Affiliation Departments of Neurology and Epidemiology, Erasmus MC University Medical Center, Rotterdam, the Netherlands

Peter Passmore

Affiliation Ageing Group, Centre for Public Health, School of Medicine, Dentistry and Biomedical Sciences, Queen's University Belfast, Belfast, United Kingdom

Thomas J. Montine

Affiliation Department of Pathology, University of Washington, Seattle, Washington, United States of America

Karolien Bettens

Affiliations Neurodegenerative Brain Diseases Group, Department of Molecular Genetics, VIB, Antwerp, Belgium, Laboratory of Neurogenetics, Institute Born-Bunge, University of Antwerp, Antwerp, Belgium

Jerome I. Rotter

Affiliation Institute for Translational Genomics and Population Sciences, Los Angeles Biomedical Research Institute at Harbor-UCLA Medical Center, Torrance, California, United States of America

Alexis Brice

Affiliations INSERM UMR_S975-CNRS UMR 7225, Université Pierre et Marie Curie, Centre de recherche de l'Institut du Cerveau et de la Moëlle épinière-CRICM, Hôpital de la Salpêtrière, Paris France, AP-HP, Hôpital de la Pitié-Salpêtrière, Paris, France

Kevin Morgan

Affiliation Institute of Genetics, Queen's Medical Centre, University of Nottingham, Nottingham, United Kingdom

Tatiana M. Foroud

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

Walter A. Kukull

Affiliation Department of Epidemiology, University of Washington, Seattle, Washington, United States of America

Didier Hannequin

Affiliation CNR-MAJ, Inserm U1079, Rouen University Hospital, 76031 France, Rouen, France

John F. Powell

Affiliation King's College London, Institute of Psychiatry, Department of Neuroscience, De Crespigny Park, Denmark Hill, London, United Kingom

Michael A. Nalls

Affiliation Laboratory of Neurogenetics, Intramural Research Program, National Institute on Aging, Bethesda, Maryland, United States of America

Karen Ritchie

Affiliations INSERM U1061, Faculty of Medicine, Hôpital La Colombière, Montpellier, France, Imperial College, London, United Kingdom

Kathryn L. Lunetta

Affiliation Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts, United States of America

John S. K. Kauwe

Affiliation Department of Biology, Brigham Young University, Provo, Utah, United States of America

Eric Boerwinkle

Affiliations Human Genome Sequencing Center, Baylor College of Medicine, Houston, Texas, United States of America, Human Genetics Center and Div. of Epidemiology, University of Texas Health Sciences Center at Houston, Houston, Texas, United States of America

Matthias Riemenschneider

Affiliation Department of Psychiatry, University Hospital, Saarland, Germany

Mercè Boada

Affiliations Memory Clinic of Fundació ACE. Institut Català de Neurociències Aplicades, Barcelona, Spain, Hospital Universitari Vall d'Hebron - Institut de Recerca, Universitat Autònoma de Barcelona. (VHIR-UAB), Barcelona, Spain

Mikko Hiltunen

Affiliations Institute of Clinical Medicine - Neurology, University of Eastern Finland, Kuopio, Finland, Department of Neurology, Kuopio University Hospital, Kuopio, Finland

Eden R. Martin

Affiliations The John P. Hussman Institute for Human Genomics, University of Miami, Miami, Florida, United States of America, Dr. John T. Macdonald Foundation Department of Human Genetics, University of Miami, Miami, Florida, United States of America

Reinhold Schmidt

Affiliation Department of Neurology, Medical University Graz, Graz, Austria

Dan Rujescu

Affiliation Department of Psychiatry, Ludwig-Maximilians University, Munich, Germany

Jean-François Dartigues

Affiliations Inserm U897, Victor Segalen University, F-33076, Bordeaux, France, Centre de Mémoire de Ressources et de Recherche de Bordeaux, CHU de Bordeaux, Bordeaux, France

Richard Mayeux

Affiliations Taub Institute on Alzheimer's Disease and the Aging Brain, Department of Neurology, Columbia University New York, New York, United States of America, Gertrude H. Sergievsky Center, Department of Neurology, Columbia University, New York, New York, United States of America

Christophe Tzourio

Affiliation Inserm U708, Victor Segalen University, Bordeaux, France

Albert Hofman

Affiliations Departments of Epidemiology, Neurology and Radiology, Erasmus MC University Medical Center, Rotterdam, the Netherlands, Netherlands Consortium for Healthy Aging, Leiden, The Netherlands

Markus M. Nöthen

Affiliation Institute of Human Genetics, Department of Genomics, Life and Brain Center, University of Bonn, and German Center for Neurodegenerative Diseases (DZNE, Bonn), Bonn, Germany

Caroline Graff

Affiliations Department Geriatric Medicine, Genetics Unit, Karolinska University Hospital Huddinge, Stockholm, Sweden, Karolinska Institutet, Department of Neurobiology, Care Sciences and Society, KIADRC, Stockholm, Sweden

Bruce M. Psaty

Affiliations Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, Washington, United States of America, Group Health Research Institute, Group Health Cooperative, Seattle, Washington, United States of America

Jonathan L. Haines

Affiliations Vanderbilt Center for Human Genetics Research, Vanderbilt University, Nashville, Tennessee, United States of America, Department of Epidemiology & Biostatistics, Case Western Reserve University, Cleveland, Ohio, United States of America

Mark Lathrop

Affiliations Department of Ophthalmology, Boston University School of Medicine, Boston, Massachusetts, United States of America, Fondation Jean Dausset- CEPH, Paris, France, McGill University and Génome Québec Innovation Centre, Montreal, Canada

Margaret A. Pericak-Vance

Affiliations The John P. Hussman Institute for Human Genomics, University of Miami, Miami, Florida, United States of America, Dr. John T. Macdonald Foundation Department of Human Genetics, University of Miami, Miami, Florida, United States of America

Lenore J. Launer

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

Christine Van Broeckhoven

Affiliations Neurodegenerative Brain Diseases Group, Department of Molecular Genetics, VIB, Antwerp, Belgium, Laboratory of Neurogenetics, Institute Born-Bunge, University of Antwerp, Antwerp, Belgium

Lindsay A. Farrer

Affiliations Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts, United States of America, Department of Medicine (Biomedical Genetics), Boston University School of Medicine, Boston, Massachusetts, United States of America, Department of Ophthalmology, Boston University School of Medicine, Boston, Massachusetts, United States of America, Department of Epidemiology, Boston University School of Public Health, Boston, Massachusetts, United States of America, Department of Neurology, Boston University School of Medicine, Boston, Massachusetts, United States of America

Cornelia M. van Duijn

Affiliations Netherlands Consortium for Healthy Aging, Leiden, The Netherlands, Department of Epidemiology, Erasmus MC University Medical Center, Rotterdam, the Netherlands, Center for Medical Systems Biology, Leiden, The Netherlands

Alfredo Ramirez

Affiliation Department of Psychiatry and Psychotherapy and Institute of Human Genetics, University of Bonn, Bonn, Germany

Sudha Seshadri

Contributed equally to this work with: Valentina Escott-Price, Céline Bellenguez, Li-San Wang, Seung-Hoan Choi, Sudha Seshadri, Gerard D. Schellenberg, Philippe Amouyel, Julie Williams

Affiliations Department of Neurology, Boston University School of Medicine, Boston, Massachusetts, United States of America, The Framingham Heart Study, Framingham, Massachusetts, United States of America

Gerard D. Schellenberg

Contributed equally to this work with: Valentina Escott-Price, Céline Bellenguez, Li-San Wang, Seung-Hoan Choi, Sudha Seshadri, Gerard D. Schellenberg, Philippe Amouyel, Julie Williams

WilliamsJ@cf.ac.uk (JW); philippe.amouyel@pasteur-lille.fr (PA); gerardsc@mail.med.upenn.edu (GDS)

Affiliation Department of Pathology and Laboratory Medicine, University of Pennsylvania Perelman School of Medicine, Philadelphia, Pennsylvania, United States of America

Philippe Amouyel

Contributed equally to this work with: Valentina Escott-Price, Céline Bellenguez, Li-San Wang, Seung-Hoan Choi, Sudha Seshadri, Gerard D. Schellenberg, Philippe Amouyel, Julie Williams

WilliamsJ@cf.ac.uk (JW); philippe.amouyel@pasteur-lille.fr (PA); gerardsc@mail.med.upenn.edu (GDS)

Affiliations Inserm U744, Lille, France, Université Lille 2, Lille, France, Institut Pasteur de Lille, Lille, France, CNR-MAJ, Centre Hospitalier Régional Universitaire de Lille, Lille, France, Centre Hospitalier Régional Universitaire de Lille, Lille, France

Julie Williams

Contributed equally to this work with: Valentina Escott-Price, Céline Bellenguez, Li-San Wang, Seung-Hoan Choi, Sudha Seshadri, Gerard D. Schellenberg, Philippe Amouyel, Julie Williams

WilliamsJ@cf.ac.uk (JW); philippe.amouyel@pasteur-lille.fr (PA); gerardsc@mail.med.upenn.edu (GDS)

Affiliation Institute of Psychological Medicine and Clinical Neurosciences, MRC Centre for Neuropsychiatric Genetics & Genomics, Cardiff University, Cardiff, United Kingdom

Competing Interests

Bruce M. Psaty serves on the DSMB for a clinical trial of a device funded by the manufacturer (Zoll LifeCor) and on the Steering Committee of the Yale Open Data Access Project funded by Johnson & Johnson. Data used in preparation of this article were obtained from the Alzheimer's Disease Neuroimaging Initiative (ADNI) database (adni.loni.usc.edu). As such, the investigators within the ADNI contributed to the design and implementation of ADNI and/or provided data but did not participate in analysis or writing of this report. A complete listing of ADNI investigators can be found at: http://adni.loni.usc.edu/wp-content/uploads/how_to_apply/ADNI_Acknowledgement_List.pdf. This does not alter the authors' adherence to PLOS ONE policies on sharing data and materials.

Author Contributions

Conceived and designed the experiments: VEP D. Harold P. Holmans S. Seshadri GDS PA JW. Analyzed the data: VEP JCL C. Bellengues LSW SHC D. Harold P. Holmans A. Richards AJ AV GR MV VC. Contributed reagents/materials/analysis tools: VEP C. Bellengues LSW SHC LJ P. Holmans D. Harold AG AV A. Richards ALdS JCL CAIV ACN RS GJ JCB GWB BGB GR TATW ND AVS VC C. Thomas MAI DZ BNV YK CFL HS BK MLD MV A. Ruiz MTB C. Reitz F. Panza P. Hollingworth OH ALF JDB D. Campion PKC C. Baldwin TB VG CC D. Craig NA C. Berr) OLL PLdJ VD JAJ DE S. Love LL IH DCR GE KS AMG NF VS AFG MJH MG K. Brown MIK LK PBG BMcG EBL AFG AJM CD ST DW S. Lovestone ER JG PStGH JC AL A. Bayer DWT LY MT P. Bosco GS P. Proitsi JC S. Sorbi FSG NCF JH MCDN P. Bossù RC C. Brayne DG ES UB M. Mancuso GS S. Moebus PM MdZ WM HH AP M. Boada F. Pasquier PC BN WP M. Mayhaus LL HH SP MMC MI D. Beekly VA FZ OV SGY EC KLHN WG C. Razquin P. Pastor IM MJO KMF PVJ OC MCOD LBC HS D. Blacker S. Mead THM DAB TBH LF CH RFAGbB P. P Passmore TJM K. Bettens JIR A. Brice KM TMF WAK D. Hannequin JFP MAN KR KLL JSKL EB MR MH ERM RS DR JFD RM C. Tzourio AH MMN CG BMP JLH ML MAPV LJL CvB LAF CMvD A. Ramirez UKBEC S. Seshadri GDS PA JW. Wrote the paper: VEP C. Bellengues LSW SHC D. Harold LJ P. Holmans AJ LAF S. Seshadri GDS PA JW.