About the Authors

Lesley Jones

Contributed equally to this work with: Lesley Jones, Peter A. Holmans

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Peter A. Holmans

Contributed equally to this work with: Lesley Jones, Peter A. Holmans

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Marian L. Hamshere

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Denise Harold

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Valentina Moskvina

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Dobril Ivanov

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Andrew Pocklington

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Richard Abraham

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Paul Hollingworth

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Rebecca Sims

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Amy Gerrish

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Jaspreet Singh Pahwa

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Nicola Jones

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Alexandra Stretton

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Angharad R. Morgan

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Simon Lovestone

Affiliation National Institute for Health Research Biomedical Research Centre for Mental Health at the South London and Maudsley National Health Service Foundation Trust and Institute of Psychiatry, Kings College, London, United Kingdom

John Powell

Affiliation Department of Neuroscience, Institute of Psychiatry, Kings College, London, United Kingdom

Petroula Proitsi

Affiliation Department of Neuroscience, Institute of Psychiatry, Kings College, London, United Kingdom

Michelle K. Lupton

Affiliation Department of Neuroscience, Institute of Psychiatry, Kings College, London, United Kingdom

Carol Brayne

Affiliation Institute of Public Health, University of Cambridge, Cambridge, United Kingdom

David C. Rubinsztein

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

Michael Gill

Affiliation Mercer's Institute for Research on Aging, St. James Hospital and Trinity College, Dublin, Ireland

Brian Lawlor

Affiliation Mercer's Institute for Research on Aging, St. James Hospital and Trinity College, Dublin, Ireland

Aoibhinn Lynch

Affiliation Mercer's Institute for Research on Aging, St. James Hospital and Trinity College, Dublin, Ireland

Kevin Morgan

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

Kristelle S. Brown

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

Peter A. Passmore

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

David Craig

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

Bernadette McGuinness

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

Stephen Todd

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

Clive Holmes

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

David Mann

Affiliation Clinical Neuroscience Research Group, Greater Manchester Neurosciences Centre, University of Manchester, Salford, United Kingdom

A. David Smith

Affiliation Oxford Project to Investigate Memory and Ageing, University of Oxford, John Radcliffe Hospital, Oxford, United Kingdom

Seth Love

Affiliation Dementia Research Group, University of Bristol Institute of Clinical Neurosciences, Frenchay Hospital, Bristol, United Kingdom

Patrick G. Kehoe

Affiliation Dementia Research Group, University of Bristol Institute of Clinical Neurosciences, Frenchay Hospital, Bristol, United Kingdom

Simon Mead

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

Nick Fox

Affiliation Dementia Research Centre, Department of Neurodegenerative Diseases, UCL Institute of Neurology, London, United Kingdom

Martin Rossor

Affiliation Dementia Research Centre, Department of Neurodegenerative Diseases, UCL Institute of Neurology, London, United Kingdom

John Collinge

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

Wolfgang Maier

Affiliation Department of Psychiatry, University of Bonn, Bonn, Germany

Frank Jessen

Affiliation Department of Psychiatry, University of Bonn, Bonn, Germany

Britta Schürmann

Affiliation Department of Psychiatry, University of Bonn, Bonn, Germany

Hendrik van den Bussche

Affiliation Institute of Primary Medical Care, University Medical Center Hamburg-Eppendorf, Germany

Isabella Heuser

Affiliation Institute of Primary Medical Care, University Medical Center Hamburg-Eppendorf, Germany

Oliver Peters

Affiliation Institute of Primary Medical Care, University Medical Center Hamburg-Eppendorf, Germany

Johannes Kornhuber

Affiliation Department of Psychiatry and Psychotherapy, University of Erlangen-Nuremberg, Erlangen, Germany

Jens Wiltfang

Affiliation Landschaftsverband Rheinland-Hospital Essen, Department of Psychiatry and Psychotherapy, University Duisburg-Essen, Essen, Germany

Martin Dichgans

Affiliations Institute for Stroke and Dementia Research, Klinikum der Universität München, Munich, Germany, Department of Neurology, Klinikum der Universität München, Munich, Germany

Lutz Frölich

Affiliation Department of Geriatric Psychiatry, Central Institute of Mental Health, Medical Faculty Mannheim, University of Heidelberg, Mannheim, Germany

Harald Hampel

Affiliations Discipline of Psychiatry, School of Medicine and Trinity College Institute of Neuroscience, Laboratory of Neuroimaging and Biomarker Research, Trinity College, University of Dublin, Dublin, Ireland, Alzheimer Memorial Center and Geriatric Psychiatry Branch, Department of Psychiatry, Ludwig-Maximilian University, Munich, Germany

Michael Hüll

Affiliation Centre for Geriatric Medicine and Section of Gerontopsychiatry and Neuropsychology, Medical School, University of Freiburg, Freiburg, Germany

Dan Rujescu

Affiliation Alzheimer Memorial Center and Geriatric Psychiatry Branch, Department of Psychiatry, Ludwig-Maximilian University, Munich, Germany

Alison M. Goate

Affiliation Departments of Psychiatry, Neurology and Genetics, Washington University School of Medicine, St. Louis, Missouri, United States of America

John S. K. Kauwe

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

Carlos Cruchaga

Affiliation Departments of Psychiatry, Neurology and Genetics, Washington University School of Medicine, St. Louis, Missouri, United States of America

Petra Nowotny

Affiliation Departments of Psychiatry, Neurology and Genetics, Washington University School of Medicine, St. Louis, Missouri, United States of America

John C. Morris

Affiliation Departments of Psychiatry, Neurology and Genetics, Washington University School of Medicine, St. Louis, Missouri, United States of America

Kevin Mayo

Affiliation Departments of Psychiatry, Neurology and Genetics, Washington University School of Medicine, St. Louis, Missouri, United States of America

Gill Livingston

Affiliation Department of Mental Health Sciences, UCL, London, United Kingdom

Nicholas J. Bass

Affiliation Department of Mental Health Sciences, UCL, London, United Kingdom

Hugh Gurling

Affiliation Department of Mental Health Sciences, UCL, London, United Kingdom

Andrew McQuillin

Affiliation Department of Mental Health Sciences, UCL, London, United Kingdom

Rhian Gwilliam

Affiliation The Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, United Kingdom

Panos Deloukas

Affiliation The Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, United Kingdom

Ammar Al-Chalabi

Affiliation MRC Centre for Neurodegeneration Research, Department of Clinical Neuroscience, King's College London, Institute of Psychiatry, London, United Kingdom

Christopher E. Shaw

Affiliation MRC Centre for Neurodegeneration Research, Department of Clinical Neuroscience, King's College London, Institute of Psychiatry, London, United Kingdom

Andrew B. Singleton

Affiliation Laboratory of Neurogenetics, National Institute on Aging, National Institutes of Health, Bethesda, Maryland, United States of America

Rita Guerreiro

Affiliation Laboratory of Neurogenetics, National Institute on Aging, National Institutes of Health, Bethesda, Maryland, United States of America

Thomas W. Mühleisen

Affiliations Department of Genomics, Life and Brain Center, University of Bonn, Bonn, Germany, Institute of Human Genetics, University of Bonn, Bonn, Germany

Markus M. Nöthen

Affiliations Department of Genomics, Life and Brain Center, University of Bonn, Bonn, Germany, Institute of Human Genetics, University of Bonn, Bonn, Germany

Susanne Moebus

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

Karl-Heinz Jöckel

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

Norman Klopp

Affiliation Institute of Epidemiology, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, Germany

H.-Erich Wichmann

Affiliations Institute of Epidemiology, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, Germany, Institute of Medical Informatics, Biometry and Epidemiology, Ludwig-Maximilians-Universität, Munich, Germany, Klinikum Grosshadern, Munich, Germany

Eckhard Rüther

Affiliation Department of Psychiatry, University of Gottingen, Gottingen, Germany

Minerva M. Carrasquillo

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

V. Shane Pankratz

Affiliation Division of Biomedical Statistics and Informatics, Mayo Clinic and Mayo Foundation, Rochester, Minnesota, United States of America

Steven G. Younkin

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

John Hardy

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

Michael C. O'Donovan

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Michael J. Owen

williamsj@cf.ac.uk (JW); owenmj@cf.ac.uk (MJO)

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Julie Williams

williamsj@cf.ac.uk (JW); owenmj@cf.ac.uk (MJO)

Affiliation Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, United Kingdom

Competing Interests

Profs Owen, J Williams and Dr Harold have a patent application in respect of genes identified in the GWAS Harold et al. Nature Genetics 2009;41(10):1088–93; this study provided data for this manuscript and was funded by the MRC and the Wellcome Trust. Dr Passmore has consulted for Pfizer and received compensation. Dr Hull has been funded by Wyeth and consulted for Pfizer, Wyeth and Merz; he has a patent pending for AD diagnostic tests. Dr Lynch has received travel expenses from Pfizer and Novartis. Prof Goate's work has been funded by NIH and AHAF. Prof Fox is a board member of the Alzheimer's Research Forum, has consulted for Abbott Laboratories and has received compensation for consulting for GE Healthcare. He has a patent for QA Box that did not arise from this work which contributes funds to his institution. Dr Morris is funded by NIA. Dr Livingston has received compensation from Lundbeck sa. Ms Stretton holds a CASE PhD studentship jointly funded by MRC and GSK. The remaining authors declare no potential conflict of interest. This does not alter the authors' adherence to all the PloS ONE policies on sharing data and materials.

Author Contributions

Conceived and designed the experiments: LJ PH MO MO JW. Performed the experiments: LJ PH DH RA PH RS AG NJ AS AM RG PD. Analyzed the data: LJ PH MH DH VM DI AP JSP. Contributed reagents/materials/analysis tools: PH SL JP PP MKL CB DCR MG BL AL KM KSB PP DC BM ST CH DM ADS SL PGK SM NCF MR JC WM FJ BS HvdB IH OP JK JW MD LF HH MH DR AMG JSKK CC PN JM KM GL NJB HMG AM AAC CES ABS RG TWM MN SM KHJ NK HEW ER MMC VSP SGY JH MO MO JW. Wrote the paper: LJ PH JH MO MO JW.