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Abstract
General cognitive function is a prominent and relatively stable human trait that is associated with many important life outcomes. We combine cognitive and genetic data from the CHARGE and COGENT consortia, and UK Biobank (total N = 300,486; age 16–102) and find 148 genome-wide significant independent loci (P < 5 × 10−8) associated with general cognitive function. Within the novel genetic loci are variants associated with neurodegenerative and neurodevelopmental disorders, physical and psychiatric illnesses, and brain structure. Gene-based analyses find 709 genes associated with general cognitive function. Expression levels across the cortex are associated with general cognitive function. Using polygenic scores, up to 4.3% of variance in general cognitive function is predicted in independent samples. We detect significant genetic overlap between general cognitive function, reaction time, and many health variables including eyesight, hypertension, and longevity. In conclusion we identify novel genetic loci and pathways contributing to the heritability of general cognitive function.
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1 Centre for Cognitive Ageing and Cognitive Epidemiology, Department of Psychology, School of Philosophy, Psychology and Language Sciences, The University of Edinburgh, Edinburgh, UK
2 Institute of Mental Health, Singapore, Singapore
3 Centre for Cognitive Ageing and Cognitive Epidemiology, Department of Psychology, School of Philosophy, Psychology and Language Sciences, The University of Edinburgh, Edinburgh, UK; Medical Genetics Section, Centre for Genomic & Experimental Medicine, MRC Institute of Genetics & Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, UK
4 BrainWorkup, LLC, Los Angeles, CA, USA; Department of Psychiatry and Behavioral Sciences, Keck School of Medicine, University of Southern California, Los Angeles, CA, USA
5 Centre for Cognitive Ageing and Cognitive Epidemiology, Department of Psychology, School of Philosophy, Psychology and Language Sciences, The University of Edinburgh, Edinburgh, UK; Social, Genetic and Developmental Psychiatry Centre, Institute of Psychiatry, Psychology & Neuroscience, King’s College London, London, UK
6 Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku, Finland; Department of Internal Medicine, Satakunta Central Hospital, Pori, Finland
7 Centre for Global Health Research, Usher Institute of Population Health Sciences and Informatics, University of Edinburgh, Teviot Place, Edinburgh, Scotland
8 Max Planck Institute for Molecular Genetics, Berlin, Germany
9 Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, Washington, USA
10 Department of Psychiatry, Icahn School of Medicine at Mount Sinai, New York, NY, USA; Mental Illness Research, Education, and Clinical Center (VISN 3), James J. Peters VA Medical Center, Bronx, NY, USA; Department of Psychiatry, Brigham and Women’s Hospital, Harvard Medical School, Boston, MA, USA
11 NORMENT, K.G. Jebsen Centre for Psychosis Research, Department of Clinical Science, University of Bergen, Bergen, Norway; Dr. Einar Martens Research Group for Biological Psychiatry, Center for Medical Genetics and Molecular Medicine, Haukeland University Hospital, Bergen, Norway
12 Institute of Mental Health, Singapore, Singapore; Center for Psychiatric Neuroscience, Feinstein Institute for Medical Research, Manhasset, NY, USA
13 NORMENT, K.G. Jebsen Centre for Psychosis Research, Department of Clinical Science, University of Bergen, Bergen, Norway; Department of Medical Genetics, Oslo University Hospital, University of Bergen, Oslo, Norway
14 Department of Psychology, University of Oslo, Oslo, Norway; Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway
15 Department of Psychology, University of Crete, Crete, Greece
16 Department of Psychiatry, University of California, San Diego, CA, USA; Center for Behavior Genetics of Aging, University of California, San Diego, CA, USA
17 Medical Research Council Human Genetics Unit, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, UK; Generation Scotland, Centre for Genomic and Experimental Medicine, University of Edinburgh, Edinburgh, UK
18 Department of Neurology, Clinical Division of Neurogeriatrics, Medical University of Graz, Graz, Austria; Institute of Medical Informatics Statistics and Documentation, Medical University of Graz, Graz, Austria
19 Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands; Department of Radiology and Nuclear Medicine, Erasmus University Medical Center, Rotterdam, The Netherlands; Department of Neurology, Erasmus University Medical Center, Rotterdam, The Netherlands
20 Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden
21 Department of Psychiatry, Yale University School of Medicine, New Haven, CT, USA
22 Department of Psychology and Logopedics, Faculty of Medicine, University of Helsinki, Helsinki, Finland; Helsinki Collegium for Advanced Studies, University of Helsinki, Helsinki, Finland
23 Department of Psychiatry and Psychotherapy, University of Cologne, Cologne, Germany
24 Department of Biostatistics, Boston University School of Public Health, Boston, MA, USA
25 Centre for Healthy Brain Ageing, School of Psychiatry, University of New South Wales, Sydney, Australia
26 NORMENT, K.G. Jebsen Centre for Psychosis Research, Department of Clinical Science, University of Bergen, Bergen, Norway; Department of Psychology, University of Oslo, Oslo, Norway
27 Neuroimaging, Cognition & Genomics (NICOG) Centre, School of Psychology and Discipline of Biochemistry, National University of Ireland Galway, Galway, Ireland
28 Medical Research Institute and Faculty of Health, University of Newcastle, New South Wa0les, Australia
29 Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland
30 Centre for EpidemiologyDivision of Population Health, Health Services Research & Primary Care, The University of Manchester, Manchester, UK
31 Department of Epidemiology and Biostatistics, School of Public Health, Imperial College London, London, UK
32 Department of Neurology, Clinical Division of Neurogeriatrics, Medical University of Graz, Graz, Austria
33 Department of Psychology, University of California Riverside, Riverside, CA, USA
34 University of Bordeaux, Bordeaux Population Health Research Center, Bordeaux, France
35 Institute for Medical Informatics, Statistics and Epidemiology, University of Leipzig, Leipzig, Germany; LIFE—Leipzig Research Center for Civilization Diseases, University of Leipzig, Leipzig, Germany
36 Department of Epidemiology, School of Public Health, University of Michigan, Ann Arbor, MI, USA; Survey Research Center, Institute for Social Research, University of Michigan, Ann Arbor, MI, USA
37 Icelandic Heart Association, Kopavogur, Iceland; University of Iceland, Reykjavik, Iceland
38 Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands; Department of Respiratory Medicine, Ghent University Hospital, Ghent, Belgium
39 Section of Gerontology and Geriatrics, Department of Internal Medicine, Leiden University Medical Center, Leiden, The Netherlands
40 Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands
41 Survey Research Center, Institute for Social Research, University of Michigan, Ann Arbor, MI, USA
42 Department of Medicine, Division of Geriatrics, University of Mississippi Medical Center, Jackson, MS, USA
43 Queensland Brain Institute, University of Queensland, Brisbane, Australia; Centre for Advanced Imaging, University of Queensland, Brisbane, Australia
44 Rush Alzheimer’s Disease Center, Rush University Medical Center, Chicago, IL, USA; Department of Neurological Sciences, Rush University Medical Center, Chicago, IL, USA
45 Center for Psychiatric Neuroscience, Feinstein Institute for Medical Research, Manhasset, NY, USA
46 National Ageing Research Institute, Royal Melbourne Hospital, Victoria, Australia; Academic Unit for Psychiatry of Old Age, University of Melbourne, St George’s Hospital, Kew, Australia
47 Univ. Lille, Inserm, CHU Lille, Institut Pasteur de Lille, U1167-LabEx DISTALZ, Lille, France
48 Department of Psychology, University of Oslo, Oslo, Norway; Norwegian Centre for Mental Disorders Research (NORMENT), Institute of Clinical Medicine, University of Oslo, Oslo, Norway
49 Mathematics and Statistics, Murdoch University, Perth, Australia
50 Hunter Medical Research Institute and Faculty of Health, University of Newcastle, New South Wales, Australia
51 Department of NeurologyBryan Alzheimer’s Disease Research Center, and Center for Genomic and Computational Biology, Duke University Medical Center, Durham, NC, USA; Psychiatry and Behavioral Sciences, Division of Medical Psychology, and Department of Neurology, Duke University Medical Center, Durham, NC, USA
52 Department of Psychiatry, Johns Hopkins University School of Medicine, Baltimore, USA; McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University School of Medicine, Baltimore, USA
53 Institute of Human Genetics, University of Bonn, Bonn, Germany; Department of Genomics, Life and Brain Center, University of Bonn, Bonn, Germany
54 Rush Alzheimer’s Disease Center, Rush University Medical Center, Chicago, IL, USA; Departments of Behavioral Sciences, Rush University Medical Center, Chicago, IL, USA
55 Centre for Healthy Brain Ageing, School of Psychiatry, University of New South Wales, Sydney, Australia; Dementia Centre for Research Collaboration, University of New South Wales, Sydney, NSW, Australia
56 Department of Psychology, Yale University, New Haven, CT, USA
57 Human Longevity Inc, Durham, NC, USA
58 UCLA Semel Institute for Neuroscience and Human Behavior, Los Angeles, CA, USA
59 23andMe, Inc., Mountain View, CA, USA
60 Department of Psychiatry, University of California, San Diego, CA, USA; Department of Cognitive Science, University of California, San Diego, La Jolla, CA, USA; Department of Neurosciences, University of California, San Diego, La Jolla, CA, USA; Department of Radiology, University of California, San Diego, La Jolla, CA, USA
61 Department of Epidemiology and Biostatistics, School of Public Health, Imperial College London, London, UK; MRC-PHE Centre for Environment, School of Public Health, Imperial College London, London, UK
62 Department of Psychology, Virginia Commonwealth University, Richmond, VA, USA
63 Clinical and Translational Neuroscience Branch, Intramural Research Program, National Institute of Mental Health, National Institute of Health, Bethesda, MD, USA
64 National Institute for Health and Welfare, Helsinki, Finland; Department of General Practice and Primary Health Care, University of Helsinki, Helsinki, Finland; Helsinki University Central Hospital, Unit of General Practice, Helsinki, Finland; Folkhälsan Research Centre, Helsinki, Finland
65 Department of Epidemiology and Biostatistics, School of Public Health, Imperial College London, London, UK; National Institute for Health and Welfare, Helsinki, Finland
66 Robertson Centre for Biostatistics, University of Glasgow, Glasgow, United Kingdom
67 Department of Psychiatry, Martin Luther University of Halle-Wittenberg, Halle, Germany
68 Virginia Institute for Psychiatric and Behavioral Genetics, Virginia Commonwealth University, Richmond, VA, USA
69 QIMR Berghofer Medical Research Institute, Brisbane, Australia
70 Department of Neurology, Johns Hopkins University School of Medicine, Baltimore, MD, USA; Department of Epidemiology, Johns Hopkins Bloomberg School of Public Health, Baltimore, MD, USA
71 Department of Data Science, University of Mississippi Medical Center, Jackson, MS, USA
72 Intramural Research Program National Institutes on Aging, National Institutes of Health, Bethesda, MD, USA
73 Department of Psychiatry, National and Kapodistrian University of Athens Medical School, Eginition Hospital, Athens, Greece; University Mental Health Research Institute, Athens, Greece; Neurobiology Research Institute, Theodor-Theohari Cozzika Foundation, Athens, Greece
74 Department of Epidemiology, University of North Carolina Gillings School of Global Public Health, Chapel Hill, NC, USA
75 Department of Clinical Physiology, Tampere University Hospital, and Finnish Cardiovascular Research Center, Tampere, Finland; Faculty of Medicine and Life Sciences, University of Tampere, Tampere, Finland; Department of Clinical Physiology, Finnish Cardiovascular Research Center—Tampere, Faculty of Medicine and Life Sciences, University of Tampere, Tampere, Finland
76 Department of Epidemiology, School of Public Health, University of Michigan, Ann Arbor, MI, USA
77 Department of Psychiatry Medical Faculty, University of Cologne, Cologne, Germany; Department of Psychiatry and Psychotherapy, University of Bonn, Bonn, Germany; German Center for Neurodegenerative Diseases (DZNE), Bonn, Germany; Department for Neurodegenerative Diseases and Geriatric Psychiatry, University of Bonn, Bonn, Germany
78 Department of Neurology, Mayo Clinic, Rochester, MN, USA
79 Centre for Healthy Brain Ageing, School of Psychiatry, University of New South Wales, Sydney, Australia; Neuropsychiatric Institute, Prince of Wales Hospital, Sydney, Australia
80 Brain and Mind Centre—The University of Sydney, Camperdown, NSW, Australia; School of Medical Sciences, University of New South Wales, Sydney, Australia
81 Department of Clinical Chemistry, Fimlab Laboratories, Finnish Cardiovascular Research Center—Tampere, Faculty of Medicine and Life Sciences, University of Tampere, Tampere, Finland; Department of Clinical Chemistry, Finnish Cardiovascular Research Center—Tampere, Faculty of Medicine and Life Sciences, University of Tampere, Tampere, Finland
82 Max Planck Institute for Human Development, Berlin, Germany; Technische Universität Dresden, Dresden, Germany
83 Genetic and Molecular Epidemiology Group, Lübeck Interdisciplinary Platform for Genome Analytics, Institutes of Neurogenetics & Cardiogenetics, University of Lübeck, Lübeck, Germany
84 Max Planck Institute for Molecular Genetics, Berlin, Germany; Max Planck Institute for Human Development, Berlin, Germany
85 Department of Neurology, School of Medicine, University of Washington, Seattle, WA, USA; Department of Epidemiology, University of Washington, Seattle, WA, USA
86 Department of Neurology and Psychiatry, University of Pittsburgh, Pittsburgh, PA, USA
87 LIFE—Leipzig Research Center for Civilization Diseases, University of Leipzig, Leipzig, Germany; Institute of Social Medicine, Occupational Health and Public Health (ISAP), University of Leipzig, Leipzig, Germany
88 Department of Biological and Medical Psychology, University of Bergen, Bergen, Norway; K. G. Jebsen Center for Neuropsychiatry, University of Bergen, Bergen, Norway
89 Umeå Center for Functional Brain Imaging (UFBI), Umeå University, Umeå, Sweden; Department of Statistics, USBE Umeå University, Umeå, Sweden
90 QIMR Berghofer Medical Research Institute, Brisbane, Australia; Institute for Molecular Bioscience, University of Queensland, Brisbane, Australia
91 Generation Scotland, Centre for Genomic and Experimental Medicine, University of Edinburgh, Edinburgh, UK; The Institute of Medical Sciences, Aberdeen Biomedical Imaging Centre, University of Aberdeen, Aberdeen, UK
92 Division of Brain Sciences, Department of Medicine, Imperial College, London, UK
93 Umeå Center for Functional Brain Imaging (UFBI), Umeå University, Umeå, Sweden; Department of Radiation Sciences, Umeå University, Umeå, Sweden; Department of Integrative Medical Biology, Umeå University, Umeå, Sweden
94 Centre for Integrated Genomic Medical Research, Institute of Population Health, University of Manchester, Manchester, UK
95 Max Planck Institute for Human Development, Berlin, Germany; Karolinska Institutet, Aging Research Center, Stockholm University, Stockholm, Sweden
96 Department of Psychology, School of Philosophy, Psychology and Language Sciences, The University of Edinburgh, Edinburgh, UK
97 Gen-Info LLC, Zagreb, Croatia; Faculty of Medicine, University of Split, Split, Croatia
98 Department of Psychology, Stanford University, Palo Alto, CA, USA
99 Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, Washington, USA; Deparment of Health Services, University of Washington, Seattle, WA, USA; Kaiser Permanente Washington Health Research Institute, Seattle, WA, USA
100 Centre for Healthy Brain Ageing, School of Psychiatry, University of New South Wales, Sydney, Australia; Department of Developmental Disability Neuropsychiatry, School of Psychiatry, University of New South Wales, Sydney, Australia
101 Institute of Social Medicine, Occupational Health and Public Health (ISAP), University of Leipzig, Leipzig, Germany
102 Department of Psychological and Brain Sciences, Indiana University, Bloomington, IN, USA
103 Institute for Translational Genomics and Population Sciences, Los Angeles Biomedical Research Institute at Harbor-UCLA Medical Center, Torrance, CA, USA; Department of Pediatrics, Harbor-UCLA Medical Center, Torrance, CA, USA
104 Department of Psychiatry, Icahn School of Medicine at Mount Sinai, New York, NY, USA; Department of Genetics and Genomic Science and Institute for Multiscale Biology, Icahn School of Medicine at Mount Sinai, New York, NY, USA; Mental Illness Research, Education, and Clinical Center (VISN 2), James J. Peters VA Medical Center, Bronx, NY, USA
105 Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku, Finland
106 Institute of Molecular Biology and Biochemistry, Centre for Molecular Medicine, Medical University of Graz, Graz, Austria
107 Robert and Beverly Lewis Center for Neuroimaging, University of Oregon, Eugene, OR, USA
108 Department of Neurology, Boston University School of Medicine, Boston, MA, USA; The National Heart, Lung, and Blood Institute’s Framingham Heart Study, Framingham, MA, USA
109 Department of Medical Biometry, Informatics and Epidemiology, University Hospital, Bonn, Germany
110 Laboratory of NeuroGenetics, Department of Psychology & Neuroscience, Duke University, Durham, NC, USA
111 Department of Molecular Epidemiology, Leiden University Medical Center, Leiden, The Netherlands
112 Department of Psychiatry, National and Kapodistrian University of Athens Medical School, Eginition Hospital, Athens, Greece; University Mental Health Research Institute, Athens, Greece
113 Institute of Cardiovascular and Medical Sciences, University of Glasgow, Glasgow, United Kingdom
114 Lieber Institute for Brain Development, Johns Hopkins University Medical Campus, Baltimore, MD, USA
115 Department of Epidemiology and Biostatistics, School of Public Health, Imperial College London, London, UK; MRC-PHE Centre for Environment, School of Public Health, Imperial College London, London, UK; Department of Hygiene and Epidemiology, University of Ioannina Medical School, Ioannina, Greece
116 University of Bordeaux, Bordeaux Population Health Research Center, Bordeaux, France; Department of Public Health, University Hospital of Bordeaux, Bordeaux, France
117 Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands; Department of Internal Medicine, Erasmus Medical Center, Rotterdam, The Netherlands
118 Medical Research Council Human Genetics Unit, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, UK
119 Campbell Family Mental Health Institute, Centre for Addiction and Mental Health, University of Toronto, Toronto, Canada
120 Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland; National Institute for Health and Welfare, Helsinki, Finland; Department of Public Health, University of Helsinki, Helsinki, Finland
121 German Center for Neurodegenerative Diseases (DZNE), Bonn, Germany; Department for Neurodegenerative Diseases and Geriatric Psychiatry, University of Bonn, Bonn, Germany
122 Department of Psychiatry and Psychotherapy, University of Bonn, Bonn, Germany
123 Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands; Department of Radiology, Erasmus MC, Rotterdam, The Netherlands
124 McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University School of Medicine, Baltimore, USA
125 Department of Psychiatry and Behavioral Sciences, Faculty of Medicine, University of Crete, Heraklion, Greece
126 Human Genetics Center, School of Public Health, University of Texas Health Science Center at Houston, Houston, TX, USA; Human Genome Sequencing Center, Baylor College of Medicine, Houston, TX, USA
127 Department of NeurologyBryan Alzheimer’s Disease Research Center, and Center for Genomic and Computational Biology, Duke University Medical Center, Durham, NC, USA
128 Neuropsychiatric Genetics Research Group, Department of Psychiatry and Trinity College Institute of Neuroscience, Trinity College Dublin, Dublin, Ireland
129 Center for Translational and Systems Neuroimmunology, Department of Neurology, Columbia University Medical Center, New York, NY, USA; Program in Medical and Population Genetics, Broad Institute, Cambridge, MA, USA
130 University of Bordeaux, Bordeaux Population Health Research Center, Bordeaux, France; Department of Neurology, University Hospital of Bordeaux, Bordeaux, France
131 Department of Epidemiology, University of Washington, Seattle, WA, USA; Department of Global Health, University of Washington, Seattle, WA, USA
132 Queensland Brain Institute, University of Queensland, Brisbane, Australia
133 Department of Epidemiology, Erasmus University Medical Center, Rotterdam, The Netherlands; Department of Neurology, Erasmus University Medical Center, Rotterdam, The Netherlands
134 Department of Cardiology, Leiden University Medical Center, Leiden, The Netherlands
135 Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland; Department of Public Health, University of Helsinki, Helsinki, Finland
136 Institute for Behavioral Genetics, University of Colorado, Boulder, CO, USA
137 Max Planck Institute for Human Development, Berlin, Germany
138 Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland; Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, UK; Department of Medical Genetics, University of Helsinki and University Central Hospital, Helsinki, Finland
139 Division of Neuroscience and Experimental Psychology, School of Biological Sciences, Manchester Academic Health Science Centre, and Manchester Medical School, Institute of Brain, Behaviour, and Mental Health, University of Manchester, Manchester, UK
140 Centre for Cognitive Ageing and Cognitive Epidemiology, Department of Psychology, School of Philosophy, Psychology and Language Sciences, The University of Edinburgh, Edinburgh, UK; Medical Genetics Section, Centre for Genomic & Experimental Medicine, MRC Institute of Genetics & Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, UK; Generation Scotland, Centre for Genomic and Experimental Medicine, University of Edinburgh, Edinburgh, UK
141 Department of Psychology and Logopedics, Faculty of Medicine, University of Helsinki, Helsinki, Finland
142 Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku, Finland; Department of Clinical Physiology and Nuclear Medicine, Turku University Hospital, Turku, Finland
143 Department of Psychiatry and Psychotherapy, University of Cologne, Cologne, Germany; Institute of Human Genetics, University of Bonn, Bonn, Germany; Department of Psychiatry and Psychotherapy, University of Bonn, Bonn, Germany
144 Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway
145 School of Medicine and Public Health, University of Newcastle, New South Wales, Australia
146 Neuroscience Research Australia, Sydney, Australia; Faculty of Medicine, University of New South Wales, Sydney, Australia
147 Centre for Cognitive Ageing and Cognitive Epidemiology, Department of Psychology, School of Philosophy, Psychology and Language Sciences, The University of Edinburgh, Edinburgh, UK; Alzheimer Scotland Dementia Research Centre, University of Edinburgh, Edinburgh, UK
148 Division of Nephrology and Hypertension, Department of Internal Medicine, Mayo Clinic, Rochester, MN, USA
149 Max Planck Institute for Human Cognitive and Brain Sciences, Leipzig, Germany; Day Clinic for Cognitive Neurology, University Hospital Leipzig, Leipzig, Germany
150 Centre for Global Health Research, Usher Institute of Population Health Sciences and Informatics, University of Edinburgh, Teviot Place, Edinburgh, Scotland; Medical Research Council Human Genetics Unit, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, UK
151 Center for Psychiatric Neuroscience, Feinstein Institute for Medical Research, Manhasset, NY, USA; Division of Psychiatry Research, Zucker Hillside Hospital, Glen Oaks, NY, USA; Department of Psychiatry, Hofstra Northwell School of Medicine, Hempstead, NY, USA
152 Centre for Cognitive Ageing and Cognitive Epidemiology, Department of Psychology, School of Philosophy, Psychology and Language Sciences, The University of Edinburgh, Edinburgh, UK; Division of Psychiatry, University of Edinburgh, Edinburgh, UK
153 Centre for Cognitive Ageing and Cognitive Epidemiology, Department of Psychology, School of Philosophy, Psychology and Language Sciences, The University of Edinburgh, Edinburgh, UK; MRC Lifecourse Epidemiology Unit, University of Southampton, Southampton, UK
154 Robert and Beverly Lewis Center for Neuroimaging, University of Oregon, Eugene, OR, USA; Department of Neurology, Boston University School of Medicine, Boston, MA, USA; Glenn Biggs Institute for Alzheimer’s and Neurodegenerative Diseases, University of Texas Health Sciences Center, San Antonio, TX, USA
155 Human Genetics Center, School of Public Health, University of Texas Health Science Center at Houston, Houston, TX, USA
156 Center for Psychiatric Neuroscience, Feinstein Institute for Medical Research, Manhasset, NY, USA; Division of Psychiatry, University of Edinburgh, Edinburgh, UK