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.

Details

Title
Study of 300,486 individuals identifies 148 independent genetic loci influencing general cognitive function
Author
Davies, Gail 1 ; Lam, Max 2 ; Harris, Sarah E 3 ; Trampush, Joey W 4   VIAFID ORCID Logo  ; Luciano, Michelle 1   VIAFID ORCID Logo  ; Hill, W David 1 ; Hagenaars, Saskia P 5 ; Ritchie, Stuart J 1 ; Marioni, Riccardo E 3 ; Fawns-Ritchie, Chloe 1 ; Liewald, David C M 1   VIAFID ORCID Logo  ; Okely, Judith A 1 ; Ahola-Olli, Ari V 6 ; Barnes, Catriona L K 7 ; Bertram, Lars 8 ; Bis, Joshua C 9 ; Burdick, Katherine E 10 ; Christoforou, Andrea 11 ; DeRosse, Pamela 12 ; Djurovic, Srdjan 13 ; Espeseth, Thomas 14   VIAFID ORCID Logo  ; Giakoumaki, Stella 15 ; Giddaluru, Sudheer 11 ; Gustavson, Daniel E 16 ; Hayward, Caroline 17   VIAFID ORCID Logo  ; Hofer, Edith 18 ; Ikram, M Arfan 19   VIAFID ORCID Logo  ; Karlsson, Robert 20   VIAFID ORCID Logo  ; Knowles, Emma 21 ; Lahti, Jari 22   VIAFID ORCID Logo  ; Leber, Markus 23 ; Li, Shuo 24   VIAFID ORCID Logo  ; Mather, Karen A 25 ; Melle, Ingrid 26 ; Morris, Derek 27   VIAFID ORCID Logo  ; Oldmeadow, Christopher 28 ; Palviainen, Teemu 29   VIAFID ORCID Logo  ; Payton, Antony 30 ; Raha Pazoki 31 ; Petrovic, Katja 32 ; Reynolds, Chandra A 33 ; Muralidharan Sargurupremraj 34 ; Scholz, Markus 35   VIAFID ORCID Logo  ; Smith, Jennifer A 36   VIAFID ORCID Logo  ; Smith, Albert V 37 ; Terzikhan, Natalie 38 ; Thalamuthu, Anbupalam 25 ; Trompet, Stella 39 ; Sven J van der Lee 40   VIAFID ORCID Logo  ; Ware, Erin B 41 ; Windham, B Gwen 42 ; Wright, Margaret J 43   VIAFID ORCID Logo  ; Yang, Jingyun 44 ; Yu, Jin 45   VIAFID ORCID Logo  ; Ames, David 46 ; Amin, Najaf 40 ; Amouyel, Philippe 47   VIAFID ORCID Logo  ; Andreassen, Ole A 48   VIAFID ORCID Logo  ; Armstrong, Nicola J 49 ; Assareh, Amelia A 25 ; Attia, John R 50   VIAFID ORCID Logo  ; Attix, Deborah 51 ; Avramopoulos, Dimitrios 52 ; Bennett, David A 44 ; Böhmer, Anne C 53   VIAFID ORCID Logo  ; Boyle, Patricia A 54 ; Brodaty, Henry 55   VIAFID ORCID Logo  ; Campbell, Harry 7 ; Cannon, Tyrone D 56 ; Cirulli, Elizabeth T 57 ; Congdon, Eliza 58 ; Emily Drabant Conley 59 ; Corley, Janie 1 ; Cox, Simon R 1   VIAFID ORCID Logo  ; Dale, Anders M 60 ; Dehghan, Abbas 61 ; Dick, Danielle 62 ; Dickinson, Dwight 63 ; Eriksson, Johan G 64 ; Evangelou, Evangelos 65   VIAFID ORCID Logo  ; Faul, Jessica D 41 ; Ford, Ian 66 ; Freimer, Nelson A 58   VIAFID ORCID Logo  ; Gao, He 31 ; Giegling, Ina 67 ; Gillespie, Nathan A 68 ; Gordon, Scott D 69 ; Gottesman, Rebecca F 70 ; Griswold, Michael E 71 ; Gudnason, Vilmundur 37 ; Harris, Tamara B 72 ; Hartmann, Annette M 67 ; Hatzimanolis, Alex 73 ; Heiss, Gerardo 74 ; Holliday, Elizabeth G 50 ; Joshi, Peter K 7   VIAFID ORCID Logo  ; Kähönen, Mika 75 ; Kardia, Sharon L R 76 ; Karlsson, Ida 20 ; Kleineidam, Luca 77 ; Knopman, David S 78 ; Kochan, Nicole A 79 ; Konte, Bettina 67   VIAFID ORCID Logo  ; Kwok, John B 80 ; Stephanie Le Hellard 11 ; Lee, Teresa 79 ; Lehtimäki, Terho 81 ; Shu-Chen, Li 82 ; Lill, Christina M 83 ; Liu, Tian 84 ; Koini, Marisa 32 ; London, Edythe 58 ; Longstreth, Will T, Jr 85 ; Lopez, Oscar L 86 ; Loukola, Anu 29 ; Luck, Tobias 87 ; Lundervold, Astri J 88   VIAFID ORCID Logo  ; Lundquist, Anders 89 ; Leo-Pekka Lyytikäinen 81   VIAFID ORCID Logo  ; Martin, Nicholas G 69 ; Montgomery, Grant W 90   VIAFID ORCID Logo  ; Murray, Alison D 91   VIAFID ORCID Logo  ; Need, Anna C 92   VIAFID ORCID Logo  ; Noordam, Raymond 39 ; Nyberg, Lars 93 ; Ollier, William 94 ; Papenberg, Goran 95 ; Pattie, Alison 96 ; Polasek, Ozren 97 ; Poldrack, Russell A 98   VIAFID ORCID Logo  ; Psaty, Bruce M 99 ; Reppermund, Simone 100 ; Riedel-Heller, Steffi G 101 ; Rose, Richard J 102 ; Rotter, Jerome I 103 ; Roussos, Panos 104 ; Rovio, Suvi P 105 ; Saba, Yasaman 106 ; Sabb, Fred W 107 ; Sachdev, Perminder S 79   VIAFID ORCID Logo  ; Satizabal, Claudia L 108   VIAFID ORCID Logo  ; Schmid, Matthias 109 ; Scott, Rodney J 50 ; Scult, Matthew A 110   VIAFID ORCID Logo  ; Simino, Jeannette 71 ; P Eline Slagboom 111 ; Smyrnis, Nikolaos 112 ; Soumaré, Aïcha 34 ; Stefanis, Nikos C 73 ; Stott, David J 113 ; Straub, Richard E 114 ; Sundet, Kjetil 14 ; Taylor, Adele M 96 ; Taylor, Kent D 103 ; Tzoulaki, Ioanna 115   VIAFID ORCID Logo  ; Tzourio, Christophe 116 ; Uitterlinden, André 117 ; Vitart, Veronique 118 ; Voineskos, Aristotle N 119 ; Kaprio, Jaakko 120   VIAFID ORCID Logo  ; Wagner, Michael 121 ; Wagner, Holger 122 ; Weinhold, Leonie 109 ; Wen, K Hoyan 40 ; Widen, Elisabeth 29 ; Yang, Qiong 24   VIAFID ORCID Logo  ; Zhao, Wei 76 ; Adams, Hieab H H 123   VIAFID ORCID Logo  ; Arking, Dan E 124 ; Bilder, Robert M 58 ; Bitsios, Panos 125 ; Boerwinkle, Eric 126 ; Chiba-Falek, Ornit 127   VIAFID ORCID Logo  ; Corvin, Aiden 128 ; De Jager, Philip L 129   VIAFID ORCID Logo  ; Debette, Stéphanie 130 ; Donohoe, Gary 27 ; Elliott, Paul 61   VIAFID ORCID Logo  ; Fitzpatrick, Annette L 131 ; Gill, Michael 128   VIAFID ORCID Logo  ; Glahn, David C 21 ; Hägg, Sara 20 ; Hansell, Narelle K 132   VIAFID ORCID Logo  ; Hariri, Ahmad R 110 ; M Kamran Ikram 133 ; Jukema, J Wouter 134   VIAFID ORCID Logo  ; Vuoksimaa, Eero 135 ; Keller, Matthew C 136 ; Kremen, William S 16 ; Launer, Lenore 72 ; Ulman Lindenberger 137 ; Palotie, Aarno 138 ; Pedersen, Nancy L 20 ; Pendleton, Neil 139   VIAFID ORCID Logo  ; Porteous, David J 140 ; Räikkönen, Katri 141 ; Raitakari, Olli T 142 ; Ramirez, Alfredo 143 ; Reinvang, Ivar 144 ; Rudan, Igor 7   VIAFID ORCID Logo  ; Rujescu, Dan 67 ; Schmidt, Reinhold 32 ; Schmidt, Helena 106 ; Schofield, Peter W 145 ; Schofield, Peter R 146   VIAFID ORCID Logo  ; Starr, John M 147 ; Steen, Vidar M 11 ; Trollor, Julian N 100 ; Turner, Steven T 148 ; Van Duijn, Cornelia M 40 ; Villringer, Arno 149 ; Weinberger, Daniel R 114 ; Weir, David R 41 ; Wilson, James F 150   VIAFID ORCID Logo  ; Malhotra, Anil 151 ; McIntosh, Andrew M 152   VIAFID ORCID Logo  ; Gale, Catharine R 153 ; Seshadri, Sudha 154 ; Mosley, Thomas H, Jr 42 ; Bressler, Jan 155 ; Lencz, Todd 156 ; Deary, Ian J 1 

 Centre for Cognitive Ageing and Cognitive Epidemiology, Department of Psychology, School of Philosophy, Psychology and Language Sciences, The University of Edinburgh, Edinburgh, UK 
 Institute of Mental Health, Singapore, Singapore 
 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 
 BrainWorkup, LLC, Los Angeles, CA, USA; Department of Psychiatry and Behavioral Sciences, Keck School of Medicine, University of Southern California, Los Angeles, CA, USA 
 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 
 Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku, Finland; Department of Internal Medicine, Satakunta Central Hospital, Pori, Finland 
 Centre for Global Health Research, Usher Institute of Population Health Sciences and Informatics, University of Edinburgh, Teviot Place, Edinburgh, Scotland 
 Max Planck Institute for Molecular Genetics, Berlin, Germany 
 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 
Pages
1-16
Publication year
2018
Publication date
May 2018
Publisher
Nature Publishing Group
e-ISSN
20411723
Source type
Scholarly Journal
Language of publication
English
ProQuest document ID
2046590727
Copyright
© 2018. This work is published under http://creativecommons.org/licenses/by/4.0/ (the “License”). Notwithstanding the ProQuest Terms and Conditions, you may use this content in accordance with the terms of the License.