About the Authors

Minerva M. Carrasquillo

Contributed equally to this work with: Minerva M. Carrasquillo, Olivia Belbin, Fanggeng Zou, Mariet Allen

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

Olivia Belbin

Contributed equally to this work with: Minerva M. Carrasquillo, Olivia Belbin, Fanggeng Zou, Mariet Allen

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

Fanggeng Zou

Contributed equally to this work with: Minerva M. Carrasquillo, Olivia Belbin, Fanggeng Zou, Mariet Allen

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

Mariet Allen

Contributed equally to this work with: Minerva M. Carrasquillo, Olivia Belbin, Fanggeng Zou, Mariet Allen

Affiliations Department of Neuroscience, Mayo Clinic College of Medicine, Jacksonville, Florida, United States of America, Medical Research Council (MRC) Human Genetics Unit, The Institute of Genetics and Molecular Medicine, Western General Hospital, Edinburgh, United Kingdom

Nilufer Ertekin-Taner

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

Morad Ansari

Affiliation Medical Research Council (MRC) Human Genetics Unit, The Institute of Genetics and Molecular Medicine, Western General Hospital, Edinburgh, United Kingdom

Samantha L. Wilcox

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

Mariah R. Kashino

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

Li Ma

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

Linda H. Younkin

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

Samuel G. Younkin

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

Curtis S. Younkin

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

Toros A. Dincman

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

Melissa E. Howard

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

Chanley C. Howell

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

Chloe M. Stanton

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

Christopher M. Watson

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

Michael Crump

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

Veronique Vitart

Affiliation Medical Research Council (MRC) Human Genetics Unit, The Institute of Genetics and Molecular Medicine, Western General Hospital, Edinburgh, United Kingdom

Caroline Hayward

Affiliation Medical Research Council (MRC) Human Genetics Unit, The Institute of Genetics and Molecular Medicine, Western General Hospital, Edinburgh, United Kingdom

Nicholas D. Hastie

Affiliation Medical Research Council (MRC) Human Genetics Unit, The Institute of Genetics and Molecular Medicine, Western General Hospital, Edinburgh, United Kingdom

Igor Rudan

Current address: Gen-info Ltd., Zagreb, Croatia

Affiliations Department of Public Health Sciences, University of Edinburgh Medical School, Edinburgh, Scotland, United Kingdom, Croatian Centre for Global Health, University of Split Medical School, Split, Croatia, Centre for Clinical Medical Research, University Hospital “Sestre Milosrdnice”, Zagreb, Croatia

Harry Campbell

Affiliation Department of Public Health Sciences, University of Edinburgh Medical School, Edinburgh, Scotland, United Kingdom

Ozren Polasek

Current address: Gen-info Ltd., Zagreb, Croatia

Affiliations Department of Public Health Sciences, University of Edinburgh Medical School, Edinburgh, Scotland, United Kingdom, Centre for Clinical Medical Research, University Hospital “Sestre Milosrdnice”, Zagreb, Croatia

Kristelle Brown

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

Peter Passmore

Affiliation Division of Psychiatry and Neuroscience, School of Medicine and Dentistry, Queen's University Belfast, Belfast, Northern Ireland, United Kingdom

David Craig

Affiliation Division of Psychiatry and Neuroscience, School of Medicine and Dentistry, Queen's University Belfast, Belfast, Northern Ireland, United Kingdom

Bernadette McGuinness

Affiliation Division of Psychiatry and Neuroscience, School of Medicine and Dentistry, Queen's University Belfast, Belfast, Northern Ireland, United Kingdom

Stephen Todd

Affiliation Division of Psychiatry and Neuroscience, School of Medicine and Dentistry, Queen's University Belfast, Belfast, Northern Ireland, United Kingdom

Patrick G. Kehoe

Affiliation Department of Clinical Science at North Bristol, University of Bristol, Frenchay Hospital, Bristol, United Kingdom

David M. Mann

Affiliation Greater Manchester Neurosciences Centre, University of Manchester, Manchester, United Kingdom

A. David Smith

Affiliation Oxford Project to Investigate Memory and Ageing (OPTIMA), University Department of Physiology, Anatomy and Genetics, Oxford, United Kingdom

Helen Beaumont

Affiliation Oxford Project to Investigate Memory and Ageing (OPTIMA), University Department of Physiology, Anatomy and Genetics, Oxford, United Kingdom

Donald Warden

Affiliation Oxford Project to Investigate Memory and Ageing (OPTIMA), University Department of Physiology, Anatomy and Genetics, Oxford, United Kingdom

Clive Holmes

Affiliation Memory Assessment and Research Centre, University of Southampton, Southampton, United Kingdom

Reinhard Heun

Affiliation Division of Neuroscience, University of Birmingham, Birmingham, United Kingdom

Heike Kölsch

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

Noor Kalsheker

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

V. Shane Pankratz

Affiliation Department of Biostatistics, Mayo Clinic and Mayo Foundation, Rochester, Minnesota, United States of America

Dennis W. Dickson

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

Neill R. Graff-Radford

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

Ronald C. Petersen

Affiliation Department of Neurology and the Mayo Alzheimer Disease Research Center, Mayo Clinic College of Medicine, Rochester, Minnesota, United States of America

Alan F. Wright

Affiliation Medical Research Council (MRC) Human Genetics Unit, The Institute of Genetics and Molecular Medicine, Western General Hospital, Edinburgh, United Kingdom

Steven G. Younkin

younkin.steven@mayo.edu

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

Kevin Morgan

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

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: MMC OB FZ MA LHY AW SGY KM. Performed the experiments: MMC OB FZ MA MA SLW MRK LM LHY TAD MEH CCH CMS CMW MC VV CH KB. Analyzed the data: MMC OB FZ MA NET MA SLW MRK SGY CSY VV CH VSP SGY KM. Contributed reagents/materials/analysis tools: MMC NH IR HC OP PP DC BM ST PGK DM ADS HB DW CH RH HK NK DWD NRGR RCP SGY KM. Wrote the paper: MMC OB FZ MA NET SGY KM. Principal investigator: SGY KM.