New insights into the genetic etiology of Alzheimer’s disease and related dementias

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Abstract

Characterization of the genetic landscape of Alzheimer’s disease (AD) and related dementias (ADD) provides a unique opportunity for a better understanding of the associated pathophysiological processes. We performed a two-stage genome-wide association study totaling 111,326 clinically diagnosed/‘proxy’ AD cases and 677,663 controls. We found 75 risk loci, of which 42 were new at the time of analysis. Pathway enrichment analyses confirmed the involvement of amyloid/tau pathways and highlighted microglia implication. Gene prioritization in the new loci identified 31 genes that were suggestive of new genetically associated processes, including the tumor necrosis factor alpha pathway through the linear ubiquitin chain assembly complex. We also built a new genetic risk score associated with the risk of future AD/dementia or progression from mild cognitive impairment to AD/dementia. The improvement in prediction led to a 1.6- to 1.9-fold increase in AD risk from the lowest to the highest decile, in addition to effects of age and the APOE ε4 allele.

Abstract

Meta-analysis of genome-wide association studies on Alzheimer’s disease and related dementias identifies new loci and enables generation of a new genetic risk score associated with the risk of future Alzheimer’s disease and dementia.

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STRING v11: protein–protein association networks with increased coverage, supporting functional discovery in genome-wide experimental datasets

Damian Szklarczyk, Annika L Gable, David Lyon … (2018)

Abstract Proteins and their functional interactions form the backbone of the cellular machinery. Their connectivity network needs to be considered for the full understanding of biological phenomena, but the available information on protein–protein associations is incomplete and exhibits varying levels of annotation granularity and reliability. The STRING database aims to collect, score and integrate all publicly available sources of protein–protein interaction information, and to complement these with computational predictions. Its goal is to achieve a comprehensive and objective global network, including direct (physical) as well as indirect (functional) interactions. The latest version of STRING (11.0) more than doubles the number of organisms it covers, to 5090. The most important new feature is an option to upload entire, genome-wide datasets as input, allowing users to visualize subsets as interaction networks and to perform gene-set enrichment analysis on the entire input. For the enrichment analysis, STRING implements well-known classification systems such as Gene Ontology and KEGG, but also offers additional, new classification systems based on high-throughput text-mining as well as on a hierarchical clustering of the association network itself. The STRING resource is available online at https://string-db.org/.

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Integrating single-cell transcriptomic data across different conditions, technologies, and species

Rahul Satija, Efthymia Papalexi, Peter Smibert … (2018)

Computational single-cell RNA-seq (scRNA-seq) methods have been successfully applied to experiments representing a single condition, technology, or species to discover and define cellular phenotypes. However, identifying subpopulations of cells that are present across multiple data sets remains challenging. Here, we introduce an analytical strategy for integrating scRNA-seq data sets based on common sources of variation, enabling the identification of shared populations across data sets and downstream comparative analysis. We apply this approach, implemented in our R toolkit Seurat (http://satijalab.org/seurat/), to align scRNA-seq data sets of peripheral blood mononuclear cells under resting and stimulated conditions, hematopoietic progenitors sequenced using two profiling technologies, and pancreatic cell 'atlases' generated from human and mouse islets. In each case, we learn distinct or transitional cell states jointly across data sets, while boosting statistical power through integrated analysis. Our approach facilitates general comparisons of scRNA-seq data sets, potentially deepening our understanding of how distinct cell states respond to perturbation, disease, and evolution.

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Conducting Meta-Analyses inRwith themetaforPackage

Wolfgang Viechtbauer (2010)

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Author and article information

Contributors

Céline Bellenguez:

ORCID: http://orcid.org/0000-0002-1240-7874

celine.bellenguez@pasteur-lille.fr

Jean-Charles Lambert:

ORCID: http://orcid.org/0000-0003-0829-7817

Jean-Charles.Lambert@pasteur-lille.fr

Journal

Journal ID (nlm-ta): Nat Genet

Journal ID (iso-abbrev): Nat Genet

Title: Nature Genetics

Publisher: Nature Publishing Group US (New York )

ISSN (Print): 1061-4036

ISSN (Electronic): 1546-1718

Publication date (Electronic): 4 April 2022

Publication date PMC-release: 4 April 2022

Publication date (Print): 2022

Volume: 54

Issue: 4

Pages: 412-436

Affiliations

[1 ]GRID grid.503422.2, ISNI 0000 0001 2242 6780, Université de Lille, INSERM, CHU Lille, Institut Pasteur Lille, U1167-RID-AGE, , Facteurs de risque et déterminants moléculaires des maladies liées au vieillissement, ; Lille, France

[2 ]GRID grid.11486.3a, ISNI 0000000104788040, Complex Genetics of Alzheimer’s Disease Group, , VIB Center for Molecular Neurology, VIB, ; Antwerp, Belgium

[3 ]GRID grid.5284.b, ISNI 0000 0001 0790 3681, Laboratory of Neurogenetics, , Institute Born - Bunge, ; Antwerp, Belgium

[4 ]GRID grid.5284.b, ISNI 0000 0001 0790 3681, Department of Biomedical Sciences, , University of Antwerp, ; Antwerp, Belgium

[5 ]GRID grid.12380.38, ISNI 0000 0004 1754 9227, Alzheimer Center Amsterdam, Department of Neurology, Amsterdam Neuroscience, , Vrije Universiteit Amsterdam, Amsterdam UMC, ; Amsterdam, the Netherlands

[6 ]GRID grid.12380.38, ISNI 0000 0004 1754 9227, Department of Complex Trait Genetics, Center for Neurogenomics and Cognitive Research, Amsterdam Neuroscience, , Vrije University, ; Amsterdam, the Netherlands

[7 ]GRID grid.15090.3d, ISNI 0000 0000 8786 803X, Department of Neurodegenerative Diseases and Geriatric Psychiatry, , University Hospital Bonn, ; Bonn, Germany

[8 ]GRID grid.6190.e, ISNI 0000 0000 8580 3777, Division of Neurogenetics and Molecular Psychiatry, Department of Psychiatry and Psychotherapy, , University of Cologne, Medical Faculty, ; Cologne, Germany

[9 ]GRID grid.424247.3, ISNI 0000 0004 0438 0426, German Center for Neurodegenerative Diseases (DZNE Bonn), ; Bonn, Germany

[10 ]GRID grid.410675.1, ISNI 0000 0001 2325 3084, Research Center and Memory Clinic Fundació ACE, , Institut Català de Neurociències Aplicades, Universitat Internacional de Catalunya, ; Barcelona, Spain

[11 ]GRID grid.413448.e, ISNI 0000 0000 9314 1427, CIBERNED, Network Center for Biomedical Research in Neurodegenerative Diseases, , National Institute of Health Carlos III, ; Madrid, Spain

[12 ]GRID grid.5645.2, ISNI 000000040459992X, Department of Epidemiology, , Erasmus MC, ; Rotterdam, the Netherlands

[13 ]GRID grid.4991.5, ISNI 0000 0004 1936 8948, Nuffield Department of Population Health, , Oxford University, ; Oxford, UK

[14 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Department of Biostatistics, Epidemiology, and Informatics, Penn Neurodegeneration Genomics Center, , University of Pennsylvania Perelman School of Medicine, ; Philadelphia, PA USA

[15 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Department of Pathology and Laboratory Medicine, , University of Pennsylvania Perelman School of Medicine, ; Philadelphia, PA USA

[16 ]GRID grid.5600.3, ISNI 0000 0001 0807 5670, MRC Centre for Neuropsychiatric Genetics and Genomics, Division of Psychological Medicine and Clinical Neuroscience, School of Medicine, , Cardiff University, ; Cardiff, UK

[17 ]GRID grid.460789.4, ISNI 0000 0004 4910 6535, CEA, Centre National de Recherche en Génomique Humaine, , Université Paris-Saclay, ; Evry, France

[18 ]GRID grid.12380.38, ISNI 0000 0004 1754 9227, Section Genomics of Neurodegenerative Diseases and Aging, Department of Human Genetics Amsterdam UMC, , Vrije Universiteit Amsterdam, Amsterdam UMC, ; Amsterdam, the Netherlands

[19 ]GRID grid.411233.6, ISNI 0000 0000 9687 399X, Brain Institute, , Federal University of Rio Grande do Norte, ; Natal, Brazil

[20 ]GRID grid.9668.1, ISNI 0000 0001 0726 2490, Institute of Biomedicine, , University of Eastern Finland, ; Kuopio, Finland

[21 ]GRID grid.189504.1, ISNI 0000 0004 1936 7558, Department of Biostatistics, , Boston University School of Public Health, ; Boston, MA USA

[22 ]GRID grid.510954.c, ISNI 0000 0004 0444 3861, Framingham Heart Study, ; Framingham, MA USA

[23 ]GRID grid.34477.33, ISNI 0000000122986657, Cardiovascular Health Research Unit, Department of Medicine, , University of Washington, ; Seattle, WA USA

[24 ]LACDR, Leiden, the Netherlands

[25 ]GRID grid.8993.b, ISNI 0000 0004 1936 9457, Department of Public Health and Carins Sciences/Geriatrics, , Uppsala University, ; Uppsala, Sweden

[26 ]GRID grid.412835.9, ISNI 0000 0004 0627 2891, Centre of Age-Related Medicine, , Stavanger University Hospital, ; Stavanger, Norway

[27 ]GRID grid.13097.3c, ISNI 0000 0001 2322 6764, Institute of Psychiatry, Psychology & Neuroscience, ; London, UK

[28 ]GRID grid.10215.37, ISNI 0000 0001 2298 7828, Department of Surgery, Biochemistry and Molecular Biology, School of Medicine, , University of Málaga, ; Málaga, Spain

[29 ]GRID grid.7080.f, ISNI 0000 0001 2296 0625, Department of Neurology, II B Sant Pau, Hospital de la Santa Creu i Sant Pau, , Universitat Autònoma de Barcelona, ; Barcelona, Spain

[30 ]Fundació Docència i Recerca MútuaTerrassa and Movement Disorders Unit, Department of Neurology, University Hospital MútuaTerrassa, Terrassa, Spain

[31 ]GRID grid.414875.b, ISNI 0000 0004 1794 4956, Memory Disorders Unit, Department of Neurology, , Hospital Universitari Mutua de Terrassa, ; Terrassa, Spain

[32 ]GRID grid.411052.3, ISNI 0000 0001 2176 9028, Laboratorio de Genética, , Hospital Universitario Central de Asturias, ; Oviedo, Spain

[33 ]GRID grid.411052.3, ISNI 0000 0001 2176 9028, Servicio de Neurología, , Hospital Universitario Central de Asturias- Oviedo and Instituto de Investigación Biosanitaria del Principado de Asturias, ; Oviedo, Spain

[34 ]GRID grid.1005.4, ISNI 0000 0004 4902 0432, Centre for Healthy Brain Ageing, School of Psychiatry, Faculty of Medicine, , University of New South Wales, ; Sydney, New South Wales Australia

[35 ]GRID grid.4793.9, ISNI 0000000109457005, First Department of Neurology, , Medical School, Aristotle University of Thessaloniki, ; Thessaloniki, Greece

[36 ]GRID grid.428867.7, Alzheimer Hellas, ; Thessaloniki, Greece

[37 ]GRID grid.411372.2, ISNI 0000 0001 0534 3000, Unidad de Demencias, , Hospital Clínico Universitario Virgen de la Arrixaca, ; Murcia, Spain

[38 ]GRID grid.7563.7, ISNI 0000 0001 2174 1754, School of Medicine and Surgery, , University of Milano-Bicocca, ; Milano, Italy

[39 ]GRID grid.415025.7, ISNI 0000 0004 1756 8604, Neurology Unit, , San Gerardo Hospital, ; Monza, Italy

[40 ]GRID grid.414818.0, ISNI 0000 0004 1757 8749, Fondazione IRCCS Ca’Granda, Ospedale Policlinico, ; Milan, Italy

[41 ]Department of Laboratory Diagnostics, III Laboratory of Analysis, Brescia Hospital, Brescia, Italy

[42 ]GRID grid.490181.5, Unitat Trastorns Cognitius, , Hospital Universitari Santa Maria de Lleida, ; Lleida, Spain

[43 ]GRID grid.420395.9, ISNI 0000 0004 0425 020X, Institut de Recerca Biomedica de Lleida (IRBLLeida), ; Lleida, Spain

[44 ]GRID grid.4708.b, ISNI 0000 0004 1757 2822, Department of Clinical Sciences and Community Health, , University of Milan, ; Milan, Italy

[45 ]GRID grid.414818.0, ISNI 0000 0004 1757 8749, Geriatic Unit, , Fondazione Cà Granda, IRCCS Ospedale Maggiore Policlinico, ; Milan, Italy

[46 ]GRID grid.5510.1, ISNI 0000 0004 1936 8921, NORMENT Centre, , University of Oslo, ; Oslo, Norway

[47 ]GRID grid.508487.6, ISNI 0000 0004 7885 7602, EA 4468, , Université de Paris, APHP, Hôpital Broca, ; Paris, France

[48 ]GRID grid.417778.a, ISNI 0000 0001 0692 3437, Laboratory of Neuropsychiatry, Department of Clinical and Behavioral Neurology, , IRCCS Santa Lucia Foundation, ; Rome, Italy

[49 ]GRID grid.84393.35, ISNI 0000 0001 0360 9602, Servei de Neurologia, , Hospital Universitari i Politècnic La Fe, ; Valencia, Spain

[50 ]GRID grid.21729.3f, ISNI 0000000419368729, Taub Institute on Alzheimer’s Disease and the Aging Brain, Department of Neurology, , Columbia University, ; New York, NY USA

[51 ]GRID grid.10383.39, ISNI 0000 0004 1758 0937, Unit of Neurology, , University of Parma and AOU, ; Parma, Italy

[52 ]GRID grid.410563.5, ISNI 0000 0004 0621 0092, Clinic of Neurology, , UH ‘Alexandrovska’, Medical University - Sofia, ; Sofia, Bulgaria

[53 ]GRID grid.510954.c, ISNI 0000 0004 0444 3861, Boston University and the NHLBI’s Framingham Heart Study, ; Boston, MA USA

[54 ]GRID grid.413448.e, ISNI 0000 0000 9314 1427, CIEN Foundation/Queen Sofia Foundation Alzheimer Center, ; Madrid, Spain

[55 ]GRID grid.152326.1, ISNI 0000 0001 2264 7217, Vanderbilt Brain Institute, , Vanderbilt University, ; Nashville, TN USA

[56 ]GRID grid.67105.35, ISNI 0000 0001 2164 3847, Cleveland Institute for Computational Biology, , Case Western Reserve University, ; Cleveland, OH USA

[57 ]GRID grid.67105.35, ISNI 0000 0001 2164 3847, Department of Population and Quantitative Health Sciences, , Case Western Reserve University, ; Cleveland, OH USA

[58 ]GRID grid.419422.8, Molecular Markers Laboratory, , IRCCS Istituto Centro San Giovanni di Dio Fatebenefratelli, ; Brescia, Italy

[59 ]GRID grid.121334.6, ISNI 0000 0001 2097 0141, Neuropsychiatry: Epidemiological and Clinical Research, PSNREC, , Université de Montpellier, INSERM U1061, ; Montpellier, France

[60 ]GRID grid.8404.8, ISNI 0000 0004 1757 2304, Department of Neuroscience, Psychology, Drug Research and Child Health, , University of Florence, ; Florence, Italy

[61 ]GRID grid.24704.35, ISNI 0000 0004 1759 9494, Azienda Ospedaliero-Universitaria Careggi, ; Florence, Italy

[62 ]GRID grid.419422.8, MAC - Memory Clinic, , IRCCS Istituto Centro San Giovanni di Dio Fatebenefratelli, ; Brescia, Italy

[63 ]GRID grid.414603.4, Geriatrics Unit, , Fondazione Policlinico A. Gemelli IRCCS, ; Rome, Italy

[64 ]GRID grid.267308.8, ISNI 0000 0000 9206 2401, Human Genetics Center, School of Public Health, , University of Texas Health Science Center at Houston, ; Houston, TX USA

[65 ]GRID grid.39382.33, ISNI 0000 0001 2160 926X, Human Genome Sequencing Center, , Baylor College of Medicine, ; Houston, TX USA

[66 ]GRID grid.7637.5, ISNI 0000000417571846, Centre for Neurodegenerative Disorders, Department of Clinical and Experimental Sciences, , University of Brescia, ; Brescia, Italy

[67 ]GRID grid.7605.4, ISNI 0000 0001 2336 6580, Department of Neuroscience “Rita Levi Montalcini”, , University of Torino, ; Torino, Italy

[68 ]GRID grid.417778.a, ISNI 0000 0001 0692 3437, Experimental Neuro-psychobiology Laboratory, Department of Clinical and Behavioral Neurology, , IRCCS Santa Lucia Foundation, ; Rome, Italy

[69 ]GRID grid.52522.32, ISNI 0000 0004 0627 3560, Department of Neurology and Clinical Neurophysiology, , University Hospital of Trondheim, ; Trondheim, Norway

[70 ]GRID grid.5947.f, ISNI 0000 0001 1516 2393, Department of Neuromedicine and Movement Science, , Norwegian University of Science and Technology, ; Trondheim, Norway

[71 ]GRID grid.267308.8, ISNI 0000 0000 9206 2401, School of Public Health, , University of Texas Health Science Center at Houston, ; Houston, TX USA

[72 ]GRID grid.1005.4, ISNI 0000 0004 4902 0432, Dementia Centre for Research Collaboration, School of Psychiatry, , University of New South Wales, ; Sydney, New South Wales Australia

[73 ]GRID grid.12361.37, ISNI 0000 0001 0727 0669, Biosciences, School of Science and Technology, , Nottingham Trent University, ; Nottingham, UK

[74 ]GRID grid.7345.5, ISNI 0000 0001 0056 1981, Centro de Neuropsiquiatría y Neurología de la Conducta (CENECON), Facultad de Medicina, , Universidad de Buenos Aires (UBA), C.A.B.A., ; Buenos Aires, Argentina

[75 ]GRID grid.7345.5, ISNI 0000 0001 0056 1981, Departamento Ciencias Fisiológicas UAII, Facultad de Medicina, UBA, C.A.B.A., ; Buenos Aires, Argentina

[76 ]Hospital Interzonal General de Agudos Eva Perón, San Martín, Buenos Aires, Argentina

[77 ]GRID grid.5645.2, ISNI 000000040459992X, Department of Neurology, , Erasmus MC, ; Rotterdam, the Netherlands

[78 ]GRID grid.5252.0, ISNI 0000 0004 1936 973X, Institute for Stroke and Dementia Research, , Klinikum der Universität München, Ludwig Maximilians Universität (LMU), ; Munich, Germany

[79 ]GRID grid.424247.3, ISNI 0000 0004 0438 0426, German Center for Neurodegenerative Diseases (DZNE, Munich), ; Munich, Germany

[80 ]GRID grid.9654.e, ISNI 0000 0004 0372 3343, Faculty of Medical & Health Sciences, , University of Auckland, ; Auckland, New Zealand

[81 ]Wales Centre for Ageing & Dementia Research, Swansea University, Wales, New Zealand

[82 ]GRID grid.67105.35, ISNI 0000 0001 2164 3847, Department of Population & Quantitative Health Sciences, , Case Western Reserve University, ; Cleveland, OH USA

[83 ]GRID grid.413448.e, ISNI 0000 0000 9314 1427, UFIEC, Instituto de Salud Carlos III, ; Madrid, Spain

[84 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Department of Pathology and Laboratory Medicine, , University of Pennsylvania, ; Philadelphia, PA USA

[85 ]GRID grid.412041.2, ISNI 0000 0001 2106 639X, INSERM, Bordeaux Population Health Research Center, UMR 1219, ISPED, CIC 1401-EC, , Université de Bordeaux, ; Bordeaux, France

[86 ]GRID grid.42399.35, ISNI 0000 0004 0593 7118, Pole Santé Publique, , CHU de Bordeaux, ; Bordeaux, France

[87 ]GRID grid.189504.1, ISNI 0000 0004 1936 7558, Medicine Biomedical Genetics Boston University School of Medicine, ; Boston, MA USA

[88 ]GRID grid.26790.3a, ISNI 0000 0004 1936 8606, Dr. John T. Macdonald Foundation Department of Human Genetics, , University of Miami, ; Miami, FL USA

[89 ]GRID grid.11794.3a, ISNI 0000000109410645, Grupo de Medicina Xenómica, Centro Nacional de Genotipado (CEGEN-PRB3-ISCIII), , Universidade de Santiago de Compostela, ; Santiago de Compostela, Spain

[90 ]GRID grid.11794.3a, ISNI 0000000109410645, Fundación Pública Galega de Medicina Xenómica- CIBERER-IDIS, , University of Santiago de Compostela, ; Santiago de Compostela, Spain

[91 ]GRID grid.9027.c, ISNI 0000 0004 1757 3630, Institute of Gerontology and Geriatrics, Department of Medicine and Surgery, , University of Perugia, ; Perugia, Italy

[92 ]GRID grid.41724.34, ISNI 0000 0001 2296 5231, Department of Genetics and CNR-MAJ, , Normandie University, UNIROUEN, INSERM U1245, CHU Rouen, ; Rouen, France

[93 ]GRID grid.152326.1, ISNI 0000 0001 2264 7217, Division of Genetic Medicine, , Vanderbilt University, ; Nashville, TN USA

[94 ]Unit of Clinical Pharmacology, University Hospital of Cagliari, Cagliari, Italy

[95 ]GRID grid.10417.33, ISNI 0000 0004 0444 9382, Radboudumc Alzheimer Center, Department of Geriatrics, , Radboud University Medical Center, ; Nijmegen, the Netherlands

[96 ]GRID grid.7400.3, ISNI 0000 0004 1937 0650, Institute for Regenerative Medicine, , University of Zürich, ; Schlieren, Switzerland

[97 ]GRID grid.412800.f, ISNI 0000 0004 1768 1690, Unidad Clínica de Enfermedades Infecciosas y Microbiología, , Hospital Universitario de Valme, ; Sevilla, Spain

[98 ]GRID grid.8142.f, ISNI 0000 0001 0941 3192, Department of Neuroscience, , Catholic University of Sacred Heart, Fondazione Policlinico Universitario A. Gemelli IRCCS, ; Rome, Italy

[99 ]GRID grid.7644.1, ISNI 0000 0001 0120 3326, University of Bari, “A. Moro”, ; Bary, Italy

[100 ]GRID grid.410558.d, ISNI 0000 0001 0035 6670, School of Medicine, , University of Thessaly, ; Larissa, Greece

[101 ]GRID grid.412041.2, ISNI 0000 0001 2106 639X, Bordeaux Population Health Research Center, , University Bordeaux, INSERM, ; Bordeaux, France

[102 ]GRID grid.4494.d, ISNI 0000 0000 9558 4598, Department of Neurology, , University Medical Center Groningen, ; Groningen, the Netherlands

[103 ]GRID grid.59734.3c, ISNI 0000 0001 0670 2351, Ronald M. Loeb Center for Alzheimer’s Disease, , Icahn School of Medicine at Mount Sinai, ; New York, NY USA

[104 ]GRID grid.59734.3c, ISNI 0000 0001 0670 2351, Department of Genetics and Genomic Sciences & Icahn Institute for Data Science and Genomic Technology, , Icahn School of Medicine at Mount Sinai, ; New York, NY USA

[105 ]GRID grid.59734.3c, ISNI 0000 0001 0670 2351, Estelle and Daniel Maggin Department of Neurology, , Icahn School of Medicine at Mount Sinai, ; New York, NY USA

[106 ]GRID grid.59734.3c, ISNI 0000 0001 0670 2351, Department of Psychiatry, , Icahn School of Medicine at Mount Sinai, ; New York, NY USA

[107 ]GRID grid.411760.5, ISNI 0000 0001 1378 7891, Department of Psychiatry, Psychosomatics and Psychotherapy, Center of Mental Health, , University Hospital, ; Wuerzburg, Germany

[108 ]GRID grid.5600.3, ISNI 0000 0001 0807 5670, UKDRI@ Cardiff, School of Medicine, , Cardiff University, ; Cardiff, UK

[109 ]GRID grid.189504.1, ISNI 0000 0004 1936 7558, Department of Neurology, , Boston University School of Medicine, ; Boston, MA USA

[110 ]GRID grid.452617.3, Munich Cluster for Systems Neurology (SyNergy), ; Munich, Germany

[111 ]GRID grid.6936.a, ISNI 0000000123222966, Klinikum rechts der Isar, Department of Psychiatry and Psychotherapy, , Technical University of Munich, School of Medicine, ; Munich, Germany

[112 ]GRID grid.5807.a, ISNI 0000 0001 1018 4307, Institute of Cognitive Neurology and Dementia Research (IKND), , Otto-Von-Guericke University, ; Magdeburg, Germany

[113 ]GRID grid.424247.3, ISNI 0000 0004 0438 0426, German Center for Neurodegenerative Diseases (DZNE), ; Magdeburg, Germany

[114 ]GRID grid.420802.c, ISNI 0000 0000 9458 5898, Icelandic Heart Association, ; Kopovagur, Iceland

[115 ]GRID grid.8767.e, ISNI 0000 0001 2290 8069, Center for Neurosciences, , Vrije Universiteit Brussel (VUB), ; Brussels, Belgium

[116 ]GRID grid.5284.b, ISNI 0000 0001 0790 3681, Reference Center for Biological Markers of Dementia (BIODEM), Institute Born-Bunge, , University of Antwerp, ; Antwerp, Belgium

[117 ]GRID grid.5284.b, ISNI 0000 0001 0790 3681, Institute Born-Bunge, , University of Antwerp, ; Antwerp, Belgium

[118 ]GRID grid.411326.3, ISNI 0000 0004 0626 3362, Department of Neurology, , UZ Brussel, ; Brussels, Belgium

[119 ]GRID grid.257413.6, ISNI 0000 0001 2287 3919, Department of Medical and Molecular Genetics, , Indiana University, ; Indianapolis, IN USA

[120 ]GRID grid.417894.7, ISNI 0000 0001 0707 5492, Fondazione IRCCS, Istituto Neurologico Carlo Besta, ; Milan, Italy

[121 ]GRID grid.512920.d, Department of Clinical Biochemistry, , Herlev and Gentofte Hospital, ; Herlev, Denmark

[122 ]GRID grid.266539.d, ISNI 0000 0004 1936 8438, Sanders-Brown Center on Aging, Department of Biostatistics, , University of Kentucky, ; Lexington, KY USA

[123 ]GRID grid.9027.c, ISNI 0000 0004 1757 3630, Centre for Memory Disturbances, Lab of Clinical Neurochemistry, Section of Neurology, , University of Perugia, ; Perugia, Italy

[124 ]GRID grid.4708.b, ISNI 0000 0004 1757 2822, University of Milan, ; Milan, Italy

[125 ]GRID grid.416992.1, ISNI 0000 0001 2179 3554, Laboratory of Neurogenetics, Department of Internal Medicine, , Texas Tech University Health Science Center, ; Lubbock, TX USA

[126 ]GRID grid.83440.3b, ISNI 0000000121901201, Reta Lila Weston Research Laboratories, Department of Molecular Neuroscience, , UCL Institute of Neurology, ; London, UK

[127 ]GRID grid.9983.b, ISNI 0000 0001 2181 4263, Faculty of Medicine, , University of Lisbon, ; Lisbon, Portugal

[128 ]GRID grid.482677.8, ISNI 0000 0000 9663 7831, Department of Psychiatry, , Social Medicine Center East- Donauspital, ; Vienna, Austria

[129 ]GRID grid.5510.1, ISNI 0000 0004 1936 8921, Institute of Clinical Medicine, , University of Oslo, ; Oslo, Norway

[130 ]GRID grid.267309.9, ISNI 0000 0001 0629 5880, Glenn Biggs Institute for Alzheimer’s & Neurodegenerative Diseases, , University of Texas Health Sciences Center, ; San Antonio, TX USA

[131 ]GRID grid.83440.3b, ISNI 0000000121901201, Dementia Research Centre, , UCL Queen Square Institute of Neurology, ; London, UK

[132 ]Unidad de Demencias, Servicio de Neurología y Neurofisiología. Instituto de Biomedicina de Sevilla (IBiS), Hospital Universitario Virgen del Rocío/CSIC/Universidad de Sevilla, Seville, Spain

[133 ]GRID grid.81821.32, ISNI 0000 0000 8970 9163, Instituto de Investigacion Sanitaria ‘Hospital la Paz’ (IdIPaz), ; Madrid, Spain

[134 ]GRID grid.465524.4, Centro de Biología Molecular Severo Ochoa (UAM-CSIC), ; Madrid, Spain

[135 ]GRID grid.81821.32, ISNI 0000 0000 8970 9163, Hospital Universitario la Paz, ; Madrid, Spain

[136 ]GRID grid.7700.0, ISNI 0000 0001 2190 4373, Department of Geriatric Psychiatry, Central Institute for Mental Health, Mannheim, , University of Heidelberg, ; Heidelberg, Germany

[137 ]GRID grid.59734.3c, ISNI 0000 0001 0670 2351, Department of Genetics and Genomic Sciences, , Ronald M. Loeb Center for Alzheimer’s Disease Icahn School of Medicine at Mount Sinai, ; New York, NY USA

[138 ]Alzheimer Research Center & Memory Clinic, Andalusian Institute for Neuroscience, Málaga, Spain

[139 ]GRID grid.411347.4, ISNI 0000 0000 9248 5770, Hospital Universitario Ramon y Cajal, IRYCIS, ; Madrid, Spain

[140 ]GRID grid.22937.3d, ISNI 0000 0000 9259 8492, Department of Psychiatry and Psychotherapy, , Medical University of Vienna, ; Vienna, Austria

[141 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Department of Biostatistics, Epidemiology, and Informatics Perelman School of Medicine, , University of Pennsylvania, ; Philadelphia, PA USA

[142 ]CAEBI, Centro Andaluz de Estudios Bioinformáticos, Sevilla, Spain

[143 ]GRID grid.4714.6, ISNI 0000 0004 1937 0626, Center for Alzheimer Research, Department NVS, Division of Neurogeriatrics, , Karolinska Institutet, ; Stockholm, Sweden

[144 ]GRID grid.24381.3c, ISNI 0000 0000 9241 5705, Unit for Hereditary Dementias, , Karolinska University Hospital-Solna, ; Stockholm, Sweden

[145 ]GRID grid.10548.38, ISNI 0000 0004 1936 9377, Aging Research Center, Department of Neurobiology, Care Sciences and Society, , Karolinska Institutet and Stockholm University, ; Stockholm, Sweden

[146 ]GRID grid.5335.0, ISNI 0000000121885934, Institute of Public Health, , University of Cambridge, ; Cambridge, UK

[147 ]GRID grid.7400.3, ISNI 0000 0004 1937 0650, Department of Child and Adolescent Psychiatry and Psychotherapy, , University Hospital of Psychiatry Zurich, University of Zurich, ; Zurich, Switzerland

[148 ]GRID grid.7400.3, ISNI 0000 0004 1937 0650, Neuroscience Center Zurich, , University of Zurich and ETH Zurich, ; Zurich, Switzerland

[149 ]GRID grid.7400.3, ISNI 0000 0004 1937 0650, Zurich Center for Integrative Human Physiology, , University of Zurich, ; Zurich, Switzerland

[150 ]GRID grid.14013.37, ISNI 0000 0004 0640 0021, Icelandic Heart Association, Faculty of Medicine, , University of Iceland, ; Reykjavik, Iceland

[151 ]GRID grid.4563.4, ISNI 0000 0004 1936 8868, Human Genetics, School of Life Sciences, Life Sciences Building, , University Park, University of Nottingham, ; Nottingham, UK

[152 ]GRID grid.9668.1, ISNI 0000 0001 0726 2490, AI Virtanen Institute for Molecular Sciences, , University of Eastern Finland, ; Kuopio, Finland

[153 ]GRID grid.6603.3, ISNI 0000000121167908, Department of Neurology, Medical School, , University of Cyprus, ; Nicosia, Cyprus

[154 ]GRID grid.26790.3a, ISNI 0000 0004 1936 8606, The John P. Hussman Institute for Human Genomics, , University of Miami, ; Miami, FL USA

[155 ]GRID grid.411439.a, ISNI 0000 0001 2150 9058, GRC 21, Alzheimer Precision Medicine Initiative (APMI), , Sorbonne University, AP-HP, Pitié-Salpêtrière Hospital, ; Paris, France

[156 ]GRID grid.10388.32, ISNI 0000 0001 2240 3300, Institute of Human Genetics, , University of Bonn, School of Medicine & University Hospital Bonn, ; Bonn, Germany

[157 ]GRID grid.9668.1, ISNI 0000 0001 0726 2490, Institute of Clinical Medicine, Neurology, , University of Eastern, ; Kuopio, Finland

[158 ]GRID grid.9668.1, ISNI 0000 0001 0726 2490, Institute of Clinical Medicine, Internal Medicine, , University of Eastern Finland, ; Kuopio, Finland

[159 ]GRID grid.5491.9, ISNI 0000 0004 1936 9297, Clinical and Experimental Science, Faculty of Medicine, , University of Southampton, ; Southampton, UK

[160 ]GRID grid.59734.3c, ISNI 0000 0001 0670 2351, Nash Family Department of Neuroscience & Friedman Brain Institute, , Icahn School of Medicine at Mount Sinai, ; New York, NY USA

[161 ]GRID grid.7692.a, ISNI 0000000090126352, Department of Neurology, , UMC Utrecht Brain Center, ; Utrecht, the Netherlands

[162 ]GRID grid.266539.d, ISNI 0000 0004 1936 8438, Biostatistics, , University of Kentucky College of Public Health, ; Lexington, KY USA

[163 ]GRID grid.253294.b, ISNI 0000 0004 1936 9115, Department of Biology, , Brigham Young University, ; Provo, UT USA

[164 ]GRID grid.5337.2, ISNI 0000 0004 1936 7603, Translational Health Sciences, Bristol Medical School, , University of Bristol, ; Bristol, UK

[165 ]GRID grid.4714.6, ISNI 0000 0004 1937 0626, Division of Clinical Geriatrics, Center for Alzheimer Research, Care Sciences and Society (NVS), , Karolinska Institutet, ; Stockholm, Sweden

[166 ]GRID grid.9668.1, ISNI 0000 0001 0726 2490, Institute of Public Health and Clinical Nutrition, , University of Eastern Finland, ; Kuopio, Finland

[167 ]GRID grid.7445.2, ISNI 0000 0001 2113 8111, Neuroepidemiology and Ageing Research Unit, , School of Public Health, Imperial College London, ; London, UK

[168 ]Research & Development, UnitStockholms Sjukhem, Stockholm, Sweden

[169 ]GRID grid.410705.7, ISNI 0000 0004 0628 207X, Department of Neurology, , Kuopio University Hospital, ; Kuopio, Finland

[170 ]GRID grid.15485.3d, ISNI 0000 0000 9950 5666, Department of Neurosciences, , University of Helsinki and Department of Geriatrics, Helsinki University Hospital, ; Helsinki, Finland

[171 ]GRID grid.5330.5, ISNI 0000 0001 2107 3311, Department of Psychiatry and Psychotherapy, , Universitätsklinikum Erlangen, and Friedrich-Alexander Universität Erlangen-Nürnberg, ; Erlangen, Germany

[172 ]GRID grid.4793.9, ISNI 0000000109457005, Laboratory of Cognitive Neuroscience, School of Psychology, , Aristotle University of Thessaloniki, ; Thessaloniki, Greece

[173 ]GRID grid.34477.33, ISNI 0000000122986657, Department of Epidemiology, , University of Washington, ; Seattle, WA USA

[174 ]GRID grid.411325.0, ISNI 0000 0001 0627 4262, Neurology Service, , Marqués de Valdecilla University Hospital (University of Cantabria and IDIVAL), ; Santander, Spain

[175 ]GRID grid.419683.1, ISNI 0000 0004 0513 0226, Stockholm Gerontology Research Center, ; Stockholm, Sweden

[176 ]GRID grid.419475.a, ISNI 0000 0000 9372 4913, Laboratory of Epidemiology, Demography, and Biometry, , National Institute of Aging, The National Institutes of Health, ; Bethesda, MD USA

[177 ]GRID grid.94365.3d, ISNI 0000 0001 2297 5165, Intramural Research Program/National Institute on Aging/National Institutes of Health, ; Bethesda, MD USA

[178 ]GRID grid.14758.3f, ISNI 0000 0001 1013 0499, Public Health Promotion Unit, Finnish Institute for Health and Welfare, ; Helsinki, Finland

[179 ]GRID grid.4491.8, ISNI 0000 0004 1937 116X, Memory Clinic, Department of Neurology, , Charles University, 2nd Faculty of Medicine and Motol University Hospital, ; Praha, Czechia

[180 ]GRID grid.483343.b, International Clinical Research Center, , St. Anne’s University Hospital Brno, ; Brno, Czechia

[181 ]GRID grid.34477.33, ISNI 0000000122986657, Departments of Neurology and Epidemiology, , University of Washington, ; Seattle, WA USA

[182 ]GRID grid.414651.3, ISNI 0000 0000 9920 5292, Department of Neurology, , Hospital Universitario Donostia, OSAKIDETZA-Servicio Vasco de Salud, ; San Sebastian, Spain

[183 ]GRID grid.21729.3f, ISNI 0000000419368729, Department of Neurology, , Columbia University, ; New York, NY USA

[184 ]GRID grid.7362.0, ISNI 0000000118820937, School of Health Sciences, , Bangor University, ; Bangor, UK

[185 ]GRID grid.8142.f, ISNI 0000 0001 0941 3192, Institute of Neurology, , Catholic University of the Sacred Heart, ; Rome, Italy

[186 ]GRID grid.21729.3f, ISNI 0000000419368729, Gertrude H. Sergievsky Center, , Columbia University, ; New York, NY USA

[187 ]GRID grid.83440.3b, ISNI 0000000121901201, MRC Prion Unit at UCL, , UCL Institute of Prion Diseases, ; London, UK

[188 ]Unidad de Trastornos del Movimiento, Servicio de Neurología y Neurofisiología. Instituto de Biomedicina de Sevilla (IBiS), Hospital Universitario Virgen del Rocío/CSIC/Universidad de Sevilla, Seville, Spain

[189 ]GRID grid.5718.b, ISNI 0000 0001 2187 5445, Institute for Urban Public Health, , University Hospital of University Duisburg-Essen, ; Essen, Germany

[190 ]GRID grid.10403.36, ISNI 0000000091771775, Neurological Tissue Bank of the Biobanc-Hospital Clinic-IDIBAPS, , Institut d’Investigacions Biomèdiques August Pi i Sunyer, ; Barcelona, Spain

[191 ]GRID grid.410458.c, ISNI 0000 0000 9635 9413, Alzheimer’s Disease and Other Cognitive Disorders Unit, Neurology Department, , Hospital Clinic, ; Barcelona, Spain

[192 ]Laboratory of Brain Aging and Neurodegeneration, FIL-CONICET, Buenos Aires, Argentina

[193 ]GRID grid.4563.4, ISNI 0000 0004 1936 8868, Human Genetics, School of Life Sciences, , University of Nottingham, ; Nottingham, UK

[194 ]GRID grid.410721.1, ISNI 0000 0004 1937 0407, Memory Impairment and Neurodegenerative Dementia (MIND) Center, , University of Mississippi Medical Center, ; Jackson, MS USA

[195 ]GRID grid.7345.5, ISNI 0000 0001 0056 1981, Laboratorio de Bioquímica Molecular, Facultad de Medicina, , Instituto de Investigaciones Médicas A. Lanari, UBA, C.A.B.A, ; Buenos Aires, Argentina

[196 ]GRID grid.34477.33, ISNI 0000000122986657, Department of Medicine, , University of Washington, ; Seattle, WA USA

[197 ]GRID grid.418563.d, ISNI 0000 0001 1090 9021, IRCCS Fondazione Don Carlo Gnocchi, ; Florence, Italy

[198 ]GRID grid.5254.6, ISNI 0000 0001 0674 042X, Department of Clinical Medicine, , University of Copenhagen, ; Copenhagen, Denmark

[199 ]GRID grid.10383.39, ISNI 0000 0004 1758 0937, DIMEC, , University of Parma, ; Parma, Italy

[200 ]GRID grid.410463.4, ISNI 0000 0004 0471 8845, Resources and Research Memory Center (MRRC) of Distalz, LicendUniversity of Lille, , INSERM, CHU Lille, UMR1172, ; Lille, France

[201 ]Institut de Biomedicina de València-CSIC CIBERNED, València, Spain

[202 ]Unitat Mixta de de Neurología y Genética, Institut d’Investigació Sanitària La Fe, València, Spain

[203 ]GRID grid.503422.2, ISNI 0000 0001 2242 6780, US 41-UMS 2014-PLBS, bilille, , Université de Lille, CNRS, INSERM, CHU Lille, Institut Pasteur de Lille, ; Lille, France

[204 ]Institute of Psychiatry and Psychotherapy, Charité-Universitätsmedizin Berlin, Freie Universität Berlin, Humboldt-Universität Zu Berlin, and Berlin Institute of Health, Berlin, Germany

[205 ]GRID grid.424247.3, ISNI 0000 0004 0438 0426, German Center for Neurodegenerative Diseases (DZNE), ; Berlin, Germany

[206 ]GRID grid.7763.5, ISNI 0000 0004 1755 3242, Department of Biomedical Sciences, , University of Cagliari, ; Cagliari, Italy

[207 ]GRID grid.8515.9, ISNI 0000 0001 0423 4662, CHUV, Old Age Psychiatry, , Department of Psychiatry, ; Lausanne, Switzerland

[208 ]GRID grid.8515.9, ISNI 0000 0001 0423 4662, Old Age Psychiatry, Department of Psychiatry, , Lausanne University Hospital, ; Lausanne, Switzerland

[209 ]GRID grid.412004.3, ISNI 0000 0004 0478 9977, Department of Geriatric Psychiatry, , University Hospital of Psychiatry Zürich, ; Zürich, Switzerland

[210 ]GRID grid.6363.0, ISNI 0000 0001 2218 4662, Department of Neuropsychiatry and Laboratory of Molecular Psychiatry, Charité, Charitéplatz 1, ; Berlin, Germany

[211 ]GRID grid.475435.4, Department of Clinical Biochemistry, Rigshospitalet, ; Copenhagen, Denmark

[212 ]GRID grid.5012.6, ISNI 0000 0001 0481 6099, Department of Psychiatry & Neuropsychologie, , Maastricht University, Alzheimer Center Limburg, ; Maastricht, the Netherlands

[213 ]GRID grid.10215.37, ISNI 0000 0001 2298 7828, Depatamento de Especialidades Quirúrgicas Bioquímica e Inmunología, Facultad de Medicina, , Universidad de Málaga, ; Málaga, Spain

[214 ]GRID grid.5292.c, ISNI 0000 0001 2097 4740, Delft Bioinformatics Lab, , Delft University of Technology, ; Delft, the Netherlands

[215 ]GRID grid.21729.3f, ISNI 0000000419368729, Taub Institute, , Columbia University, ; New York, NY USA

[216 ]GRID grid.253294.b, ISNI 0000 0004 1936 9115, Bioinformatics, College of Life Sciences, , Brigham Young University, ; Provo, UT USA

[217 ]GRID grid.9647.c, ISNI 0000 0004 7669 9786, Institute of Social Medicine, Occupational Health and Public Health, , University of Leipzig, ; Leipzig, Germany

[218 ]GRID grid.411760.5, ISNI 0000 0001 1378 7891, Center of Mental Health, Clinic and Policlinic of Psychiatry, Psychosomatics and Psychotherapy, , University Hospital of Würzburg, ; Wuerzburg, Germany

[219 ]GRID grid.413782.b, Department of Research and Innovation, , Helse Fonna, Haugesund Hospital, ; Haugesund, Norway

[220 ]GRID grid.7914.b, ISNI 0000 0004 1936 7443, Institute of Clinical Medicine (K1), , The University of Bergen, ; Bergen, Norway

[221 ]GRID grid.10215.37, ISNI 0000 0001 2298 7828, Departamento de Especialidades Quirúrgicas, Bioquímicas e Inmunología, School of Medicine, , University of Málaga, ; Málaga, Spain

[222 ]GRID grid.432329.d, ISNI 0000 0004 1789 4477, Department of Neuroscience and Mental Health, , AOU Città della Salute e della Scienza di Torino, ; Torino, Italy

[223 ]Athens Association of Alzheimer’s Disease and Related Disorders, Athens, Greece

[224 ]GRID grid.52522.32, ISNI 0000 0004 0627 3560, Department of Geriatrics, St. Olav’s Hospital, , Trondheim University Hospital, ; Trondheim, Norway

[225 ]Department of Immunology, Hospital Universitario Doctor Negrín, Las Palmas de Gran Canaria, Las Palmas, Spain

[226 ]GRID grid.5841.8, ISNI 0000 0004 1937 0247, Neurology Department-Hospital Clínic, IDIBAPS, , Universitat de Barcelona, ; Barcelona, Spain

[227 ]GRID grid.5216.0, ISNI 0000 0001 2155 0800, First Department of Neurology, , Aiginition Hospital, National and Kapodistrian University of Athens, Medical School, ; Athens, Greece

[228 ]GRID grid.5718.b, ISNI 0000 0001 2187 5445, LVR-Hospital Essen, Department of Psychiatry and Psychotherapy, Medical Faculty, , University of Duisburg-Essen, ; Essen, Germany

[229 ]GRID grid.13648.38, ISNI 0000 0001 2180 3484, Department of Primary Medical Care, , University Medical Centre Hamburg-Eppendorf, ; Hamburg, Germany

[230 ]GRID grid.15090.3d, ISNI 0000 0000 8786 803X, Institute of Medical Biometry, Informatics and Epidemiology, , University Hospital of Bonn, ; Bonn, Germany

[231 ]GRID grid.55325.34, ISNI 0000 0004 0389 8485, Department of Geriatric Medicine, , Oslo University Hospital, ; Oslo, Norway

[232 ]GRID grid.417011.2, ISNI 0000 0004 1769 6825, Laboratory for Advanced Hematological Diagnostics, Department of Hematology and Stem Cell Transplant, , Vito Fazzi Hospital, ; Lecce, Italy

[233 ]GRID grid.411068.a, ISNI 0000 0001 0671 5785, Centro de Investigación Biomédica en Red de Diabetes y Enfermedades Metabólicas Asociadas, , CIBERDEM, Hospital Clínico San Carlos, ; Madrid, Spain

[234 ]GRID grid.10388.32, ISNI 0000 0001 2240 3300, Department of Neurology, , University of Bonn, ; Bonn, Germany

[235 ]GRID grid.7763.5, ISNI 0000 0004 1755 3242, Department of Biomedical Sciences, Section of Neuroscience and Clinical Pharmacology, , University of Cagliari, ; Cagliari, Italy

[236 ]GRID grid.461096.c, ISNI 0000 0004 0627 3042, Department of Psychiatry, , Namsos Hospital, ; Namsos, Norway

[237 ]GRID grid.5645.2, ISNI 000000040459992X, Department of Internal Medicine and Biostatistics, , Erasmus MC, ; Rotterdam, the Netherlands

[238 ]GRID grid.11486.3a, ISNI 0000000104788040, Neurodegenerative Brain Diseases Group, , VIB Center for Molecular Neurology, VIB, ; Antwerp, Belgium

[239 ]GRID grid.5645.2, ISNI 000000040459992X, Department of Neurology, ErasmusMC, ; Rotterdam, the Netherlands

[240 ]GRID grid.5596.f, ISNI 0000 0001 0668 7884, Laboratory for Cognitive Neurology, Department of Neurosciences, , University of Leuven, ; Leuven, Belgium

[241 ]GRID grid.410569.f, ISNI 0000 0004 0626 3338, Neurology Department, , University Hospitals Leuven, ; Leuven, Belgium

[242 ]GRID grid.411984.1, ISNI 0000 0001 0482 5331, Department of Psychiatry and Psychotherapy, , University Medical Center Goettingen, ; Goettingen, Germany

[243 ]GRID grid.38142.3c, ISNI 000000041936754X, Department of Psychiatry, , Harvard Medical School, McLean Hospital, ; Belmont, MA USA

[244 ]GRID grid.41724.34, ISNI 0000 0001 2296 5231, Department of Neurology and CNR-MAJ, F 76000, Normandy Center for Genomic and Personalized Medicine, , Normandie University, UNIROUEN, INSERM U1245, CHU Rouen, ; Rouen, France

[245 ]GRID grid.424247.3, ISNI 0000 0004 0438 0426, German Center for Neurodegenerative Diseases (DZNE), ; Goettingen, Germany

[246 ]Medical Science Department, iBiMED, Aveiro, Portugal

[247 ]GRID grid.15823.3d, ISNI 0000 0004 0622 2843, Department of Nutrition and Diatetics, , Harokopio University, ; Athens, Greece

[248 ]GRID grid.189504.1, ISNI 0000 0004 1936 7558, Department of Medicine (Biomedical Genetics), , Boston University School of Medicine, ; Boston, MA USA

[249 ]GRID grid.432380.e, Neurosciences Area, , Instituto Biodonostia, ; San Sebastian, Spain

[250 ]GRID grid.34477.33, ISNI 0000000122986657, Department of Health Service, , University of Washington, ; Seattle, WA USA

[251 ]GRID grid.6190.e, ISNI 0000 0000 8580 3777, Excellence Cluster on Cellular Stress Responses in Aging-Associated Diseases (CECAD), , University of Cologne, ; Cologne, Germany

[252 ]GRID grid.414877.9, ISNI 0000 0004 0609 2583, Department of Clinical Biochemistry, , Hematology and Immunology, Na Homolce Hospital, ; Prague, Czechia

[253 ]GRID grid.9027.c, ISNI 0000 0004 1757 3630, Institute of Gerontology and Geriatrics, Department of Medicine, , University of Perugia, ; Perugia, Italy

[254 ]GRID grid.9668.1, ISNI 0000 0001 0726 2490, Insitute of Biomedicine, , University of Eastern Finland, ; Kuopio, Finland

[255 ]GRID grid.10858.34, ISNI 0000 0001 0941 4873, Center for Life Course Health Research, , University of Oulu, ; Oulu, Finland

[256 ]GRID grid.412326.0, ISNI 0000 0004 4685 4917, Medical Research Center Oulu, , Oulu University Hospital, ; Oulu, Finland

[257 ]GRID grid.7737.4, ISNI 0000 0004 0410 2071, University of Helsinki and Helsinki University Hospital, ; Helsinki, Finland

[258 ]GRID grid.5600.3, ISNI 0000 0001 0807 5670, Division of Psychological Medicine and Clinial Neurosciences, MRC Centre for Neuropsychiatric Genetics and Genomics, , Cardiff University, ; Cardiff, UK

[259 ]GRID grid.13097.3c, ISNI 0000 0001 2322 6764, Institute of Psychiatry, Psychology and Neuroscience, , Kings College London, ; London, UK

[260 ]GRID grid.83440.3b, ISNI 0000000121901201, Division of Psychiatry, , University College London, ; London, UK

[261 ]GRID grid.5335.0, ISNI 0000000121885934, Institute of Public Health, , University of Cambridge, ; Cambridge, UK

[262 ]GRID grid.4563.4, ISNI 0000 0004 1936 8868, Institute of Genetics, Queens Medical Centre, , University of Nottingham, ; Nottingham, UK

[263 ]GRID grid.4777.3, ISNI 0000 0004 0374 7521, Ageing Group, , Centre for Public Health, School of Medicine, Dentistry and Biomedical Sciences, Queen’s University Belfast, ; Belfast, UK

[264 ]GRID grid.10306.34, ISNI 0000 0004 0606 5382, The Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, ; Cambridge, UK

[265 ]GRID grid.83440.3b, ISNI 0000000121901201, Dementia Research Centre, Department of Neurodegenerative Disease, , UCL Institute of Neurology, ; London, UK

[266 ]GRID grid.416409.e, ISNI 0000 0004 0617 8280, Mercer’s Institute for Research on Ageing, , St James’ Hospital, ; Dublin, Ireland

[267 ]GRID grid.15596.3e, ISNI 0000000102380260, School of Biotechnology, , Dublin City University, ; Dublin, Ireland

[268 ]GRID grid.4777.3, ISNI 0000 0004 0374 7521, Centre for Public Health, School of Medicine, Dentistry and Biomedical Sciences, , Queens University, ; Belfast, UK

[269 ]GRID grid.4991.5, ISNI 0000 0004 1936 8948, Department of Psychiatry, , University of Oxford, ; Oxford, UK

[270 ]GRID grid.1049.c, ISNI 0000 0001 2294 1395, Genetic Epidemiology, , QIMR Berghofer Medical Research Institute, ; Herston, Queensland Australia

[271 ]GRID grid.13097.3c, ISNI 0000 0001 2322 6764, Department of Basic and Clinical Neuroscience, , Institute of Psychiatry, Psychology and Neuroscience, Kings College London, ; London, UK

[272 ]GRID grid.5379.8, ISNI 0000000121662407, Division of Neuroscience and Experimental Psychology, School of Biological Sciences, Faculty of Biology, Medicine and Health, , University of Manchester, Manchester Academic Health Science Centre, ; Manchester, UK

[273 ]GRID grid.8348.7, ISNI 0000 0001 2306 7492, Oxford Project to Investigate Memory and Ageing (OPTIMA), , University of Oxford, Level 4, John Radcliffe Hospital, ; Oxford, UK

[274 ]GRID grid.83440.3b, ISNI 0000000121901201, Department of Mental Health Sciences, , University College London, ; London, UK

[275 ]GRID grid.4777.3, ISNI 0000 0004 0374 7521, Ageing Group, Centre for Public Health, School of Medicine, Dentistry and Biomedical Sciences, , Queen’s University Belfast, ; Belfast, UK

[276 ]GRID grid.83440.3b, ISNI 0000000121901201, Dementia Research Centre, , UCL, ; London, UK

[277 ]GRID grid.5338.d, ISNI 0000 0001 2173 938X, Servei de Neurologia Hospital Clínic, , Universitari de València, ; Valencia, Spain

[278 ]GRID grid.15090.3d, ISNI 0000 0000 8786 803X, Department of Radiology, , University Hospital Bonn, ; Bonn, Germany

[279 ]GRID grid.424247.3, ISNI 0000 0004 0438 0426, German Center for Neurodegenerative Diseases (DZNE), ; Tübingen, Germany

[280 ]GRID grid.428620.a, Section for Dementia Research, Department of Psychiatry, , Hertie Institute for Clinical Brain Research, ; Tübingen, Germany

[281 ]GRID grid.5252.0, ISNI 0000 0004 1936 973X, Department of Psychiatry and Psychotherapy, , University Hospital, LMU Munich, ; Munich, Germany

[282 ]GRID grid.7821.c, ISNI 0000 0004 1770 272X, Service of Neurology, University Hospital Marqués de Valdecilla, IDIVAL, , University of Cantabria, ; Santander, Spain

[283 ]GRID grid.410563.5, ISNI 0000 0004 0621 0092, Molecular Medicine Center, Department of Medical chemistry and biochemistry, , Medical University of Sofia, ; Sofia, Bulgaria

[284 ]GRID grid.411760.5, ISNI 0000 0001 1378 7891, Department of Psychiatry, Psychosomatics and Psychotherapy, Center of Mental Health, , University Hospital of Würzburg, ; Würzburg, Germany

[285 ]GRID grid.490137.8, ISNI 0000 0004 0474 3784, ENYS (Estudio en Neurociencias y Sistemas Complejos) CONICET- Hospital El Cruce “Nestor Kirchner”- UNAJ, ; Buenos Aires, Argentina

[286 ]HIGA Eva Perón, Buenes Aires, Argentina

[287 ]Neurología Clinica, Buenes Aires, Argentina

[288 ]Dirección de Atención de Adultos Mayores del Min. Salud Desarrollo Social y Deportes de la Pcia. de Mendoza, Mendoza, Argentina

[289 ]Laboratorio de Genética Forense del Ministerio Público de la Pcia de La Pampa, La Pampa, Argentina

[290 ]Fundacion Sinapsis, Santa Rosa, Argentina

[291 ]Hospital Dr. Lucio Molas, Santa Rosa; Fundacion Ayuda Enfermo Renal y Alta Complejidad (FERNAC), Santa Rosa, Argentina

[292 ]Laboratory of Brain Aging and Neurodegeneration (FIL), Buneos Aires, Argentina

[293 ]GRID grid.7345.5, ISNI 0000 0001 0056 1981, Centro de Neuropsiquiatría y Neurología de la Conducta (CENECON), Facultad de Medicina, Universidad de Buenos Aires (UBA), C.A.B.A., ; Buenos Aires, Argentina

[294 ]GRID grid.5335.0, ISNI 0000000121885934, Cambridge Institute for Medical Research and UK Dementia Research Institute, , University of Cambridge, ; Cambridge, UK

[295 ]GRID grid.424247.3, ISNI 0000 0004 0438 0426, German Center for Neurodegenerative Diseases (DZNE), ; Rostock, Germany

[296 ]GRID grid.411068.a, ISNI 0000 0001 0671 5785, Centro de Investigación Biomédica en Red de Diabetes y Enfermedades Metabólicas Asociadas, , CIBERDEM, Hospital Clínico San Carlos, ; Madrid, Spain

[297 ]GRID grid.11794.3a, ISNI 0000000109410645, Grupo de Medicina Xen´omica, Centro Nacional de Genotipado (CEGEN-PRB3-ISCIII), , Universidad de Santiago de Compostela, ; Santiago de Compostela, Spain

[298 ]GRID grid.106023.6, ISNI 0000 0004 1770 977X, Servei de Neurologia, , Hospital Clínic Universitari de València, ; Valencia, Spain

[299 ]GRID grid.411164.7, ISNI 0000 0004 1796 5984, Department of Neurology, , Hospital Universitario Son Espases, ; Palma, Spain

[300 ]GRID grid.11480.3c, ISNI 0000000121671098, BIOMICs, País Vasco; Centro de Investigación Lascaray, , Universidad del País Vasco UPV/EHU, ; Vitoria-Gasteiz, Spain

[301 ]GRID grid.424841.f, Fundación para la Formación e Investigación Sanitarias de la Región de Murcia, ; Palma, Spain

[302 ]GRID grid.428824.0, Centro de Investigación y Terapias Avanzadas, Fundación CITA-Alzheimer, ; San Sebastian, Spain

[303 ]GRID grid.428855.6, Navarrabiomed, ; Pamplona, Spain

[304 ]GRID grid.411251.2, ISNI 0000 0004 1767 647X, Hospital Universitario La Princesa, ; Madrid, Spain

[305 ]UMR 7179 CNRS/MNHN, Brunoy, France

[306 ]GRID grid.50550.35, ISNI 0000 0001 2175 4109, Sorbonne Université, Paris Brain Institute, APHP, INSERM, CNRS, ; Paris, France

[307 ]GRID grid.462844.8, ISNI 0000 0001 2308 1657, Department of Neurology, , Institute of Memory and Alzheimer’s Disease (IM2A), Pitié-Salpêtrière Hospital, AP-HP, Boulevard de l’Hôpital, ; Paris, France

[308 ]GRID grid.4563.4, ISNI 0000 0004 1936 8868, Institute of Genetics, Queen’s Medical Centre, , University of Nottingham, ; Nottingham, UK

[309 ]GRID grid.1006.7, ISNI 0000 0001 0462 7212, Institute for Ageing and Health, , Newcastle University, Campus for Ageing and Vitality, ; Newcastle upon Tyne, UK

[310 ]GRID grid.83440.3b, ISNI 0000000121901201, Department of Neurodegenerative Disease, , UCL Institute of Neurology, ; London, UK

[311 ]GRID grid.13097.3c, ISNI 0000 0001 2322 6764, Department of Old Age Psychiatry, , Institute of Psychiatry, Psychology and Neuroscience, King’s College London, ; London, UK

[312 ]GRID grid.10388.32, ISNI 0000 0001 2240 3300, Department of Psychiatry and Psychotherapy, , University of Bonn, ; Bonn, Germany

[313 ]GRID grid.13097.3c, ISNI 0000 0001 2322 6764, Department of Basic and Clinical Neuroscience, , Institute of Psychiatry, Psychology and Neuroscience, King’s College London, ; London, UK

[314 ]GRID grid.10388.32, ISNI 0000 0001 2240 3300, Institute for Medical Biometry, Informatics and Epidemiology, , University of Bonn, ; Bonn, Germany

[315 ]GRID grid.5337.2, ISNI 0000 0004 1936 7603, Population Health Sciences, Bristol Medical School, , University of Bristol, ; Bristol, UK

[316 ]GRID grid.5491.9, ISNI 0000 0004 1936 9297, Division of Clinical Neurosciences, School of Medicine, , University of Southampton, ; Southampton, UK

[317 ]GRID grid.500936.9, ISNI 0000 0000 8621 4130, Somerset Partnership NHS Trust, ; Somerset, UK

[318 ]GRID grid.241103.5, ISNI 0000 0001 0169 7725, Institute of Primary Care and Public Health, , Cardiff University, University Hospital of Wales, ; Cardiff, UK

[319 ]GRID grid.6363.0, ISNI 0000 0001 2218 4662, Department of Psychiatry and Psychotherapy, , Charité University Medicine, ; Berlin, Germany

[320 ]GRID grid.6190.e, ISNI 0000 0000 8580 3777, Cologne Center for Genomics, , University of Cologne, ; Cologne, Germany

[321 ]GRID grid.4567.0, ISNI 0000 0004 0483 2525, Institute of Epidemiology, , Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, ; Munich, Germany

[322 ]GRID grid.492206.b, ISNI 0000 0004 0494 2070, Department of Psychiatry and Psychotherapy, , University Hospital, ; Saarland, Germany

[323 ]GRID grid.5963.9, Department of Psychiatry, , University of Freiburg, ; Freiburg, Germany

[324 ]GRID grid.7700.0, ISNI 0000 0001 2190 4373, Central Institute of Mental Health, Medical Faculty Mannheim, , University of Heidelberg, ; Heidelberg, Germany

[325 ]GRID grid.410718.b, ISNI 0000 0001 0262 7331, Institute for Medical Informatics, Biometry and Epidemiology, , University Hospital of Essen, University Duisburg-Essen, ; Essen, Germany

[326 ]GRID grid.55325.34, ISNI 0000 0004 0389 8485, Division of Mental Health and Addiction, , Oslo University Hospital, ; Oslo, Norway

[327 ]GRID grid.5947.f, ISNI 0000 0001 1516 2393, Department of Mental Health, Faculty of Medicine and Health Sciences, , Norwegian University of Science and Technology, ; Trondheim, Norway

[328 ]Department of Psychiatry, Hospital Namsos, Nord-Trøndelag Health Trust, Namsos, Norway

[329 ]GRID grid.411279.8, ISNI 0000 0000 9637 455X, Department of Neurology, , Akershus University Hospital, ; Lørenskog, Norway

[330 ]GRID grid.412929.5, ISNI 0000 0004 0627 386X, Centre for Old Age Psychiatry Research, , Innlandet Hospital Trust, ; Ottestad, Norway

[331 ]GRID grid.7737.4, ISNI 0000 0004 0410 2071, Institute for Molecular Medicine Finland, HiLIFE, , University of Helsinki, ; Helsinki, Finland

[332 ]GRID grid.431072.3, ISNI 0000 0004 0572 4227, AbbVie, ; Chicago, IL USA

[333 ]GRID grid.417815.e, ISNI 0000 0004 5929 4381, Astra Zeneca, ; Cambridge, UK

[334 ]GRID grid.417832.b, ISNI 0000 0004 0384 8146, Biogen, ; Cambridge, MA USA

[335 ]GRID grid.419971.3, ISNI 0000 0004 0374 8313, Celgene, ; Summit, NJ USA

[336 ]GRID grid.418158.1, ISNI 0000 0004 0534 4718, Genentech, ; San Francisco, CA USA

[337 ]GRID grid.418236.a, ISNI 0000 0001 2162 0389, GlaxoSmithKline, ; Brentford, UK

[338 ]GRID grid.417993.1, ISNI 0000 0001 2260 0793, Merck, ; Kenilworth, NJ USA

[339 ]GRID grid.410513.2, ISNI 0000 0000 8800 7493, Pfizer, ; New York, NY USA

[340 ]GRID grid.417924.d, Sanofi, ; Paris, France

[341 ]GRID grid.511646.1, ISNI 0000 0004 7480 276X, Maze Therapeutics, ; San Francisco, CA USA

[342 ]Janssen Biotech, Beerse, Belgium

[343 ]GRID grid.7737.4, ISNI 0000 0004 0410 2071, HiLIFE, , University of Helsinki, ; Helsinki, Finland

[344 ]GRID grid.1374.1, ISNI 0000 0001 2097 1371, Auria Biobank, , University of Turku, Hospital District of Southwest Finland, ; Turku, Finland

[345 ]GRID grid.14758.3f, ISNI 0000 0001 1013 0499, THL Biobank, , The National Institute of Health and Welfare Helsinki, ; Helsinki, Finland

[346 ]GRID grid.452433.7, ISNI 0000 0000 9387 9501, Finnish Red Cross Blood Service, , Finnish Hematology Registry and Clinical Biobank, ; Helsinki, Finland

[347 ]GRID grid.424664.6, ISNI 0000 0004 0410 2290, Helsinki Biobank, , Helsinki University and Hospital District of Helsinki and Uusimaa, ; Helsinki, Finland

[348 ]GRID grid.10858.34, ISNI 0000 0001 0941 4873, Northern Finland Biobank Borealis, , University of Oulu, Northern Ostrobothnia Hospital District, ; Oulu, Finland

[349 ]GRID grid.4991.5, ISNI 0000 0004 1936 8948, Oxford Healthy Aging Project, Clinical Trial Service Unit, , University of Oxford, ; Oxford, UK

[350 ]GRID grid.502801.e, ISNI 0000 0001 2314 6254, Finnish Clinical Biobank Tampere, , University of Tampere, Pirkanmaa Hospital District, ; Tampere, Finland

[351 ]GRID grid.9668.1, ISNI 0000 0001 0726 2490, Biobank of Eastern Finland, , University of Eastern Finland / Northern Savo Hospital District, ; Kuopio, Finland

[352 ]Central Finland Biobank, University of Jyväskylä, Central Finland Health Care District, Jyväskylä, Finland

[353 ]GRID grid.511030.6, Business Finland, ; Helsinki, Finland

[354 ]Northern Savo Hospital District, Kuopio, Finland

[355 ]GRID grid.415018.9, ISNI 0000 0004 0472 1956, Pirkanmaa Hospital District, ; Tampere, Finland

[356 ]GRID grid.424664.6, ISNI 0000 0004 0410 2290, Hospital District of Helsinki and Uusimaa, ; Helsinki, Finland

[357 ]GRID grid.426612.5, ISNI 0000 0004 0366 9623, Hospital District of Southwest Finland, ; Turku, Finland

[358 ]GRID grid.415018.9, ISNI 0000 0004 0472 1956, Pirkanmaa Hospital District, ; Tampere, Finland

[359 ]GRID grid.437577.5, ISNI 0000 0004 0450 6025, Northern Ostrobothnia Hospital District, ; Oulu, Finland

[360 ]Northern Savo Hospital District, Kuopio, Finland

[361 ]GRID grid.14758.3f, ISNI 0000 0001 1013 0499, The National Institute of Health and Welfare Helsinki, ; Helsinki, Finland

[362 ]GRID grid.419971.3, ISNI 0000 0004 0374 8313, Bristol Myers Squibb, ; New York, NY USA

[363 ]GRID grid.66859.34, ISNI 0000 0004 0546 1623, Broad Institute, ; Cambridge, MA USA

[364 ]GRID grid.168010.e, ISNI 0000000419368956, University of Stanford, ; Stanford, CA USA

[365 ]GRID grid.7737.4, ISNI 0000 0004 0410 2071, University of Helsinki, ; Helsinki, Finland

[366 ]GRID grid.502801.e, ISNI 0000 0001 2314 6254, University of Tampere, ; Tampere, Finland

[367 ]GRID grid.452433.7, ISNI 0000 0000 9387 9501, Finnish Red Cross Blood Service, ; Helsinki, Finland

[368 ]GRID grid.1374.1, ISNI 0000 0001 2097 1371, University of Turku, ; Turku, Finland

[369 ]GRID grid.266539.d, ISNI 0000 0004 1936 8438, Sanders-Brown Center on Aging, Department of Epidemiology, , College of Public Health, University of Kentucky, ; Lexington, KY USA

[370 ]GRID grid.267313.2, ISNI 0000 0000 9482 7121, Department of Psychiatry, , University of Texas Southwestern Medical Center, ; Dallas, TX USA

[371 ]GRID grid.89336.37, ISNI 0000 0004 1936 9924, Department of Neurology, Dell Medical School, , University of Texas at Austin, ; Austin, TX USA

[372 ]GRID grid.21107.35, ISNI 0000 0001 2171 9311, Department of Neurology, , Johns Hopkins University, ; Baltimore, MD USA

[373 ]GRID grid.214458.e, ISNI 0000000086837370, Department of Neurology, , University of Michigan, ; Ann Arbor, MI USA

[374 ]GRID grid.413800.e, ISNI 0000 0004 0419 7525, Geriatric Research, Education and Clinical Center (GRECC), , VA Ann Arbor Healthcare System (VAAAHS), ; Ann Arbor, MI USA

[375 ]GRID grid.214458.e, ISNI 0000000086837370, Michigan Alzheimer’s Disease Center, , University of Michigan, ; Ann Arbor, MI USA

[376 ]GRID grid.417467.7, ISNI 0000 0004 0443 9942, Department of Neuroscience, , Mayo Clinic, ; Jacksonville, FL USA

[377 ]GRID grid.266871.c, ISNI 0000 0000 9765 6057, Department of Pharmacology and Neuroscience, , University of North Texas Health Science Center, ; Fort Worth, TX USA

[378 ]GRID grid.257413.6, ISNI 0000 0001 2287 3919, Departments of Neurology, Radiology, and Medical and Molecular Genetics, , Indiana University School of Medicine, ; Indianapolis, IN USA

[379 ]GRID grid.257413.6, ISNI 0000 0001 2287 3919, Indiana Alzheimer’s Disease Research Center, , Indiana University School of Medicine, ; Indianapolis, IN USA

[380 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Department of Psychiatry, Perelman School of Medicine, , University of Pennsylvania, ; Philadelphia, PA USA

[381 ]GRID grid.28803.31, ISNI 0000 0001 0701 8607, Geriatric Research, Education and Clinical Center (GRECC), , University of Wisconsin, ; Madison, WI USA

[382 ]GRID grid.28803.31, ISNI 0000 0001 0701 8607, Department of Medicine, , University of Wisconsin, ; Madison, WI USA

[383 ]GRID grid.14003.36, ISNI 0000 0001 2167 3675, Wisconsin Alzheimer’s Disease Research Center, ; Madison, WI USA

[384 ]GRID grid.240684.c, ISNI 0000 0001 0705 3621, Department of Neurological Sciences, , Rush University Medical Center, ; Chicago, IL USA

[385 ]GRID grid.240684.c, ISNI 0000 0001 0705 3621, Department of Behavioral Sciences, , Rush University Medical Center, ; Chicago, IL USA

[386 ]GRID grid.240684.c, ISNI 0000 0001 0705 3621, Rush Alzheimer’s Disease Center, , Rush University Medical Center, ; Chicago, IL USA

[387 ]GRID grid.414208.b, ISNI 0000 0004 0619 8759, Civin Laboratory for Neuropathology, , Banner Sun Health Research Institute, ; Phoenix, AZ USA

[388 ]GRID grid.21925.3d, ISNI 0000 0004 1936 9000, Departments of Psychiatry, Neurology, and Psychology, , University of Pittsburgh School of Medicine, ; Pittsburgh, PA USA

[389 ]GRID grid.34477.33, ISNI 0000000122986657, National Alzheimer’s Coordinating Center, , University of Washington, ; Seattle, WA USA

[390 ]GRID grid.267308.8, ISNI 0000 0000 9206 2401, Human Genetics Center, Department of Epidemiology, Human Genetics, and Environmental Sciences, School of Public Health, , The University of Texas Health Science Center at Houston, ; Houston, TX USA

[391 ]GRID grid.4367.6, ISNI 0000 0001 2355 7002, Department of Psychiatry and Hope Center Program on Protein Aggregation and Neurodegeneration, , Washington University School of Medicine, ; St. Louis, MO USA

[392 ]GRID grid.89336.37, ISNI 0000 0004 1936 9924, Department of Psychiatry, , University of Texas at Austin/Dell Medical School, ; Austin, TX USA

[393 ]GRID grid.16753.36, ISNI 0000 0001 2299 3507, Department of Pathology, , Northwestern University Feinberg School of Medicine, ; Chicago, IL USA

[394 ]GRID grid.16753.36, ISNI 0000 0001 2299 3507, Cognitive Neurology and Alzheimer’s Disease Center, , Northwestern University Feinberg School of Medicine, ; Chicago, IL USA

[395 ]GRID grid.413919.7, ISNI 0000 0004 0420 6540, VA Puget Sound Health Care System/GRECC, ; Seattle, WA USA

[396 ]GRID grid.38142.3c, ISNI 000000041936754X, Department of Epidemiology, , Harvard School of Public Health, ; Boston, MA USA

[397 ]GRID grid.32224.35, ISNI 0000 0004 0386 9924, Department of Psychiatry, , Massachusetts General Hospital/Harvard Medical School, ; Boston, MA USA

[398 ]GRID grid.66875.3a, ISNI 0000 0004 0459 167X, Department of Neurology, , Mayo Clinic, ; Rochester, MN USA

[399 ]GRID grid.281044.b, ISNI 0000 0004 0463 5388, Swedish Medical Center, ; Seattle, WA USA

[400 ]GRID grid.266102.1, ISNI 0000 0001 2297 6811, Department of Neurology, , University of California San Francisco, ; San Francisco, CA USA

[401 ]GRID grid.266100.3, ISNI 0000 0001 2107 4242, Department of Neurosciences, , University of California San Diego, ; La Jolla, CA USA

[402 ]GRID grid.26009.3d, ISNI 0000 0004 1936 7961, Department of Medicine, , Duke University, ; Durham, NC USA

[403 ]GRID grid.412016.0, ISNI 0000 0001 2177 6375, University of Kansas Alzheimer’s Disease Center, , University of Kansas Medical Center, ; Kansas City, KS USA

[404 ]GRID grid.4367.6, ISNI 0000 0001 2355 7002, Department of Pathology and Immunology, , Washington University, ; St. Louis, MO USA

[405 ]GRID grid.170693.a, ISNI 0000 0001 2353 285X, USF Health Byrd Alzheimer’s Institute, , University of South Florida, ; Tampa, FL USA

[406 ]GRID grid.270240.3, ISNI 0000 0001 2180 1622, Fred Hutchinson Cancer Research Center, ; Seattle, WA USA

[407 ]GRID grid.484420.e, Mental Health and Behavioral Science Service, , Bruce W. Carter VA Medical Center, ; Miami, FL USA

[408 ]GRID grid.10698.36, ISNI 0000000122483208, Department of Genetics, , University of North Carolina at Chapel Hill, ; Chapel Hill, NC USA

[409 ]GRID grid.19006.3e, ISNI 0000 0000 9632 6718, Neurogenetics Program, , University of California, Los Angeles, ; Los Angeles, CA USA

[410 ]GRID grid.42505.36, ISNI 0000 0001 2156 6853, Department of Neurology, , University of Southern California, ; Los Angeles, CA USA

[411 ]GRID grid.241167.7, ISNI 0000 0001 2185 3318, Section of Gerontology and Geriatric Medicine Research, , Wake Forest School of Medicine, ; Winston-Salem, NC USA

[412 ]GRID grid.266093.8, ISNI 0000 0001 0668 7243, Department of Neurology, , University of California, Irvine, ; Irvine, CA USA

[413 ]GRID grid.26790.3a, ISNI 0000 0004 1936 8606, Department of Psychiatry and Behavioral Sciences, Miller School of Medicine, , University of Miami, ; Miami, FL USA

[414 ]GRID grid.4367.6, ISNI 0000 0001 2355 7002, NeuroGenomics and Informatics, , Washington University in St. Louis, ; St. Louis, MO USA

[415 ]GRID grid.4367.6, ISNI 0000 0001 2355 7002, Department of Psychiatry, , Washington University in St. Louis, ; St Louis, MO USA

[416 ]GRID grid.39382.33, ISNI 0000 0001 2160 926X, Alzheimer’s Disease and Memory Disorders Center, , Baylor College of Medicine, ; Houston, TX USA

[417 ]GRID grid.267313.2, ISNI 0000 0000 9482 7121, Department of Population and Data Sciences, , University of Texas Southwestern Medical Center, ; Dallas, Texas USA

[418 ]GRID grid.239585.0, ISNI 0000 0001 2285 2675, Center for Translational and Computational Neuroimmunology, Department of Neurology, , Columbia University Medical Center, ; New York, NY USA

[419 ]GRID grid.27860.3b, ISNI 0000 0004 1936 9684, Department of Neurology, , University of California, Davis, ; Sacramento, CA USA

[420 ]GRID grid.416992.1, ISNI 0000 0001 2179 3554, Departments of Neurology, Pharmacology and Neuroscience, , Texas Tech University Health Science Center, ; Lubbock, TX USA

[421 ]GRID grid.266093.8, ISNI 0000 0001 0668 7243, Institute for Memory Impairments and Neurological Disorders, , University of California, Irvine, ; Irvine, CA USA

[422 ]GRID grid.410396.9, ISNI 0000 0004 0430 4458, Wien Center for Alzheimer’s Disease and Memory Disorders, , Mount Sinai Medical Center, ; Miami Beach, FL USA

[423 ]GRID grid.417467.7, ISNI 0000 0004 0443 9942, Department of Neurology, , Mayo Clinic, ; Jacksonville, FL USA

[424 ]GRID grid.240684.c, ISNI 0000 0001 0705 3621, Rush Institute for Healthy Aging, Department of Internal Medicine, , Rush University Medical Center, ; Chicago, IL USA

[425 ]GRID grid.266871.c, ISNI 0000 0000 9765 6057, Office of Strategy and Measurement, , University of North Texas Health Science Center, ; Fort Worth, TX USA

[426 ]GRID grid.265892.2, ISNI 0000000106344187, Department of Pathology, , University of Alabama at Birmingham, ; Birmingham, AL USA

[427 ]GRID grid.257413.6, ISNI 0000 0001 2287 3919, Department of Neurology, , Indiana University, ; Indianapolis, IN USA

[428 ]GRID grid.137628.9, ISNI 0000 0004 1936 8753, Department of Psychiatry, , New York University, ; New York, NY USA

[429 ]GRID grid.32224.35, ISNI 0000 0004 0386 9924, C.S. Kubik Laboratory for Neuropathology, , Massachusetts General Hospital, ; Charlestown, MA USA

[430 ]GRID grid.59734.3c, ISNI 0000 0001 0670 2351, Department of Neuroscience, Ronald M. Loeb Center for Alzheimer’s Disease, , Icahn School of Medicine at Mount Sinai, ; New York, NY USA

[431 ]GRID grid.266871.c, ISNI 0000 0000 9765 6057, Department of Health Behavior and Health Systems, , University of North Texas Health Science Center, ; Fort Worth, TX USA

[432 ]GRID grid.266871.c, ISNI 0000 0000 9765 6057, Department of Health Management and Policy, School of Public Health, , University of North Texas Health Science Center, ; Fort Worth, TX USA

[433 ]GRID grid.189967.8, ISNI 0000 0001 0941 6502, Department of Pathology and Laboratory Medicine, , Emory University, ; Atlanta, GA USA

[434 ]GRID grid.189967.8, ISNI 0000 0001 0941 6502, Emory Alzheimer’s Disease Center, , Emory University, ; Atlanta, GA USA

[435 ]GRID grid.257413.6, ISNI 0000 0001 2287 3919, Department of Pathology and Laboratory Medicine, , Indiana University, ; Indianapolis, IN USA

[436 ]GRID grid.34477.33, ISNI 0000000122986657, Department of Radiology, , University of Washington, ; Seattle, WA USA

[437 ]GRID grid.239552.a, ISNI 0000 0001 0680 8770, Center for Spatial and Functional Genomics, Division of Human Genetics, , Children’s Hospital of Philadelphia, ; Philadelphia, PA USA

[438 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Department of Pediatrics, Perelman School of Medicine, , University of Pennsylvania, ; Philadelphia, PA USA

[439 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Department of Genetics, Perelman School of Medicine, , University of Pennsylvania, ; Philadelphia, PA USA

[440 ]GRID grid.62560.37, ISNI 0000 0004 0378 8294, Division of Genetics, Department of Medicine and Partners Center for Personalized Genetic Medicine, , Brigham and Women’s Hospital and Harvard Medical School, ; Boston, MA USA

[441 ]GRID grid.32224.35, ISNI 0000 0004 0386 9924, Department of Neurology, , Massachusetts General Hospital/Harvard Medical School, ; Boston, MA USA

[442 ]GRID grid.239552.a, ISNI 0000 0001 0680 8770, Center for Applied Genomics, , Children’s Hospital of Philadelphia, ; Philadelphia, PA USA

[443 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Division of Human Genetics, Department of Pediatrics, Perelman School of Medicine, , University of Pennsylvania, ; Philadelphia, PA USA

[444 ]GRID grid.21925.3d, ISNI 0000 0004 1936 9000, Department of Pathology (Neuropathology), , University of Pittsburgh, ; Pittsburgh, PA USA

[445 ]GRID grid.265892.2, ISNI 0000000106344187, Department of Neurology, , University of Alabama at Birmingham, ; Birmingham, AL USA

[446 ]GRID grid.266093.8, ISNI 0000 0001 0668 7243, Department of Pathology and Laboratory Medicine, , University of California Irvine, ; Irvine, CA USA

[447 ]GRID grid.168010.e, ISNI 0000000419368956, Department of Epidemiology and Population Health, , Stanford University, ; Stanford, CA USA

[448 ]GRID grid.168010.e, ISNI 0000000419368956, Department of Neurology and Neurological Sciences, , Stanford University, ; Stanford, CA USA

[449 ]GRID grid.412807.8, ISNI 0000 0004 1936 9916, Vanderbilt Memory and Alzheimer’s Center, Department of Neurology, , Vanderbilt University Medical Center, ; Nashville, TN USA

[450 ]GRID grid.267313.2, ISNI 0000 0000 9482 7121, Department of Surgery, , University of Texas Southwestern Medical Center, ; Dallas, TX USA

[451 ]GRID grid.250942.8, ISNI 0000 0004 0507 3225, Neurogenomics Division, , Translational Genomics Research Institute, ; Phoenix, AZ USA

[452 ]GRID grid.26009.3d, ISNI 0000 0004 1936 7961, Department of Pathology, , Duke University, ; Durham, NC USA

[453 ]GRID grid.267313.2, ISNI 0000 0000 9482 7121, Department of Neurology, , University of Texas Southwestern Medical Center, ; Dallas, TX USA

[454 ]GRID grid.267313.2, ISNI 0000 0000 9482 7121, Department of Neurological Surgery, , University of Texas Southwestern Medical Center, ; Dallas, TX USA

[455 ]GRID grid.4367.6, ISNI 0000 0001 2355 7002, Hope Center Program on Protein Aggregation and Neurodegeneration, , Washington University School of Medicine, ; St. Louis, MO USA

[456 ]GRID grid.34477.33, ISNI 0000000122986657, Department of Genome Sciences, , University of Washington, ; Seattle, WA USA

[457 ]GRID grid.27860.3b, ISNI 0000 0004 1936 9684, Department of Pathology and Laboratory Medicine, , University of California, Davis, ; Sacramento, CA USA

[458 ]GRID grid.21925.3d, ISNI 0000 0004 1936 9000, Department of Human Genetics, , University of Pittsburgh, ; Pittsburgh, PA USA

[459 ]GRID grid.21925.3d, ISNI 0000 0004 1936 9000, Alzheimer’s Disease Research Center, , University of Pittsburgh, ; Pittsburgh, PA USA

[460 ]GRID grid.251993.5, ISNI 0000000121791997, Department of Neurology, , Albert Einstein College of Medicine, ; New York, NY USA

[461 ]GRID grid.5288.7, ISNI 0000 0000 9758 5690, Department of Neurology, , Oregon Health and Science University, ; Portland, OR USA

[462 ]GRID grid.410404.5, ISNI 0000 0001 0165 2383, Department of Neurology, , Portland Veterans Affairs Medical Center, ; Portland, OR USA

[463 ]GRID grid.34477.33, ISNI 0000000122986657, Department of Laboratory Medicine and Pathology, , University of Washington, ; Seattle, WA USA

[464 ]GRID grid.189504.1, ISNI 0000 0004 1936 7558, Department of Pathology, , Boston University, ; Boston, MA USA

[465 ]GRID grid.266102.1, ISNI 0000 0001 2297 6811, Department of Neuropsychology, , University of California San Francisco, ; San Francisco, CA USA

[466 ]GRID grid.266093.8, ISNI 0000 0001 0668 7243, Department of Neurobiology and Behavior, , University of California Irvine, ; Irvine, CA USA

[467 ]GRID grid.189967.8, ISNI 0000 0001 0941 6502, Department of Neurology, , Emory University, ; Atlanta, GA USA

[468 ]GRID grid.488833.c, ISNI 0000 0004 0615 7519, Kaiser Permanente Washington Health Research Institute, ; Seattle, WA USA

[469 ]GRID grid.239578.2, ISNI 0000 0001 0675 4725, Cleveland Clinic Lou Ruvo Center for Brain Health, , Cleveland Clinic, ; Cleveland, OH USA

[470 ]GRID grid.214458.e, ISNI 0000000086837370, Department of Pathology, , University of Michigan, ; Ann Arbor, MI USA

[471 ]National Center for PTSD at Boston VA Healthcare System, Boston, MA USA

[472 ]GRID grid.21107.35, ISNI 0000 0001 2171 9311, Department of Psychiatry, , Johns Hopkins University, ; Baltimore, MD USA

[473 ]GRID grid.137628.9, ISNI 0000 0004 1936 8753, Department of Medicine (Pulmonary), , New York University, ; New York, NY USA

[474 ]GRID grid.26790.3a, ISNI 0000 0004 1936 8606, Department of Neurology, Miller School of Medicine, , University of Miami, ; Miami, FL USA

[475 ]GRID grid.266100.3, ISNI 0000 0001 2107 4242, Department of Pathology, , University of California San Diego, ; La Jolla, CA USA

[476 ]GRID grid.34477.33, ISNI 0000000122986657, School of Nursing Northwest Research Group on Aging, , University of Washington, ; Seattle, WA USA

[477 ]GRID grid.410513.2, ISNI 0000 0000 8800 7493, Pfizer Worldwide Research and Development, ; New York, NY USA

[478 ]GRID grid.266102.1, ISNI 0000 0001 2297 6811, Weill Institute for Neurosciences, Memory and Aging Center, , University of California, San Francisco, ; San Francisco, CA USA

[479 ]GRID grid.42505.36, ISNI 0000 0001 2156 6853, Department of Pathology, , University of Southern California, ; Los Angeles, CA USA

[480 ]GRID grid.168010.e, ISNI 0000000419368956, Department of Pathology, , Stanford University School of Medicine, ; Stanford, CA USA

[481 ]GRID grid.4367.6, ISNI 0000 0001 2355 7002, Department of Neurology, , Washington University at St. Louis, ; St. Louis, MO USA

[482 ]GRID grid.266871.c, ISNI 0000 0000 9765 6057, Institute for Translational Research, , University of North Texas Health Science Center, ; Fort Worth, TX USA

[483 ]GRID grid.27860.3b, ISNI 0000 0004 1936 9684, Center for Mind and Brain and Department of Neurology, , University of California, Davis, ; Sacramento, CA USA

[484 ]GRID grid.264756.4, ISNI 0000 0004 4687 2082, Center for Population Health and Aging, , Texas A&M University Health Science Center, ; Lubbock, TX USA

[485 ]GRID grid.267309.9, ISNI 0000 0001 0629 5880, Department of Family and Community Medicine, , University of Texas Health Science Center San Antonio, ; San Antonio, TX USA

[486 ]GRID grid.66875.3a, ISNI 0000 0004 0459 167X, Department of Laboratory Medicine and Pathology, , Mayo Clinic, ; Rochester, MN USA

[487 ]GRID grid.34477.33, ISNI 0000000122986657, Department of Psychiatry and Behavioral Sciences, , University of Washington School of Medicine, ; Seattle, WA USA

[488 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Department of Bioengineering, , University of Pennsylvania, ; Philadelphia, PA USA

[489 ]GRID grid.137628.9, ISNI 0000 0004 1936 8753, Alzheimer’s Disease Center, , New York University, ; New York, NY USA

[490 ]GRID grid.430503.1, ISNI 0000 0001 0703 675X, Department of Neurology, , University of Colorado School of Medicine, ; Aurora, CO USA

[491 ]GRID grid.266871.c, ISNI 0000 0000 9765 6057, Department of Internal Medicine and Geriatrics, , University of North Texas Health Science Center, ; Fort Worth, TX USA

[492 ]GRID grid.264766.7, ISNI 0000 0001 2289 1930, Department of Medical Education, , TCU/UNTHSC School of Medicine, ; Fort Worth, TX USA

[493 ]Arizona Alzheimer’s Consortium, Phoenix, AZ USA

[494 ]GRID grid.418204.b, ISNI 0000 0004 0406 4925, Banner Alzheimer’s Institute, ; Phoenix, AZ USA

[495 ]GRID grid.134563.6, ISNI 0000 0001 2168 186X, Department of Psychiatry, , University of Arizona, ; Phoenix, AZ USA

[496 ]GRID grid.17063.33, ISNI 0000 0001 2157 2938, Tanz Centre for Research in Neurodegenerative Disease, , University of Toronto, ; Toronto, Ontario Canada

[497 ]GRID grid.257413.6, ISNI 0000 0001 2287 3919, Department of Radiology and Imaging Sciences, , Indiana University, ; Indianapolis, IN USA

[498 ]GRID grid.240684.c, ISNI 0000 0001 0705 3621, Department of Pathology (Neuropathology), , Rush University Medical Center, ; Chicago, IL USA

[499 ]GRID grid.42505.36, ISNI 0000 0001 2156 6853, Department of Psychiatry, , University of Southern California, ; Los Angeles, CA USA

[500 ]GRID grid.5335.0, ISNI 0000000121885934, Cambridge Institute for Medical Research, , University of Cambridge, ; Cambridge, UK

[501 ]GRID grid.17063.33, ISNI 0000 0001 2157 2938, Faculty of Medicine, Department of Medicine (Neurology), , University of Toronto, ; Toronto, Ontario Canada

[502 ]GRID grid.486749.0, ISNI 0000 0004 4685 2620, Center for Applied Health Research, , Baylor Scott & White Health, ; Temple, TX USA

[503 ]GRID grid.412408.b, College of Medicine, , Texas A&M University Health Science Center, ; College Station, TX USA

[504 ]GRID grid.47100.32, ISNI 0000000419368710, Program in Cellular Neuroscience, Neurodegeneration and Repair, , Yale University School of Medicine, ; New Haven, CT USA

[505 ]GRID grid.266093.8, ISNI 0000 0001 0668 7243, Department of Psychiatry and Human Behavior, , University of California, Irvine, ; Irvine, CA USA

[506 ]GRID grid.10698.36, ISNI 0000000122483208, Renaissance Computing Institute, , University of North Carolina at Chapel Hill, ; Chapel Hill, NC USA

[507 ]GRID grid.21107.35, ISNI 0000 0001 2171 9311, Department of Pathology, , Johns Hopkins University, ; Baltimore, MD USA

[508 ]GRID grid.19006.3e, ISNI 0000 0000 9632 6718, Department of Pathology and Laboratory Medicine, , University of California, Los Angeles, ; Los Angeles, CA USA

[509 ]GRID grid.16753.36, ISNI 0000 0001 2299 3507, Department of Psychiatry and Behavioral Sciences, , Northwestern University Feinberg School of Medicine, ; Chicago, IL USA

[510 ]GRID grid.26009.3d, ISNI 0000 0004 1936 7961, Department of Psychiatry and Behavioral Sciences, , Duke University, ; Durham, NC USA

[511 ]GRID grid.34477.33, ISNI 0000000122986657, Department of Biostatistics, , University of Washington, ; Seattle, WA USA

[512 ]GRID grid.240324.3, ISNI 0000 0001 2109 4251, Department of Psychiatry, , New York University Grossman School of Medicine, ; New York, NY USA

[513 ]GRID grid.240324.3, ISNI 0000 0001 2109 4251, Center for Cognitive Neurology and Departments of Neurology and Pathology, , New York University Grossman School of Medicine, ; New York, NY USA

[514 ]GRID grid.5288.7, ISNI 0000 0000 9758 5690, Department of Pathology, , Oregon Health and Science University, ; Portland, OR USA

[515 ]GRID grid.26790.3a, ISNI 0000 0004 1936 8606, Evelyn F. McKnight Brain Institute, Department of Neurology, Miller School of Medicine, , University of Miami, ; Miami, FL USA

[516 ]GRID grid.67105.35, ISNI 0000 0001 2164 3847, Department of Pathology, , Case Western Reserve University, ; Cleveland, OH USA

[517 ]GRID grid.9582.6, ISNI 0000 0004 1794 5983, Centre for Genomic and Precision Medicine, , College of Medicine, UI, ; Ibadan, Nigeria

[518 ]GRID grid.4367.6, ISNI 0000 0001 2355 7002, Washington University at St. Louis, ; St. Louis, MO USA

[519 ]GRID grid.1032.0, ISNI 0000 0004 0375 4078, Mathematics and Statistics, , Curtin University, ; Perth, Western Australia Australia

[520 ]GRID grid.21925.3d, ISNI 0000 0004 1936 9000, Departments of Psychiatry, Neurology, and Psychology, , University of Pittsburgh, ; Pittsburgh, PA USA

[521 ]GRID grid.424247.3, ISNI 0000 0004 0438 0426, Population Health Sciences, , German Center for Neurodegenerative Diseases (DZNE), ; Bonn, Germany

[522 ]GRID grid.38142.3c, ISNI 000000041936754X, Brigham and Women’s Hospital, , Harvard University, ; Boston, MA USA

[523 ]GRID grid.412041.2, ISNI 0000 0001 2106 639X, INSERM U1219, , University of Bordeaux, ; Bordeaux, France

[524 ]GRID grid.4305.2, ISNI 0000 0004 1936 7988, University of Edinburgh, ; Edinburgh, UK

[525 ]GRID grid.4305.2, ISNI 0000 0004 1936 7988, Centre for Cognitive Ageing and Cognitive Epidemiology, , University of Edinburgh, ; Edinburgh, UK

[526 ]GRID grid.27860.3b, ISNI 0000 0004 1936 9684, Department of Neurology and Center for Neuroscience, , University of California, Davis, ; Davis, CA USA

[527 ]GRID grid.412041.2, ISNI 0000 0001 2106 639X, Bordeaux Population Health Research Center, Team VIN-TAGE, UMR 1219, , University of Bordeaux, INSERM, ; Bordeaux, France

[528 ]GRID grid.21925.3d, ISNI 0000 0004 1936 9000, University of Pittsburgh, ; Pittsburgh, PA USA

[529 ]GRID grid.34477.33, ISNI 0000000122986657, Department of Family Medicine, , University of Washington, ; Seattle, WA USA

[530 ]GRID grid.7692.a, ISNI 0000000090126352, University Medical Center Utrecht, ; Utrecht, the Netherlands

[531 ]GRID grid.411023.5, ISNI 0000 0000 9159 4457, Psychiatric Genetic Epidemiology & Neurobiology Laboratory (PsychGENe Lab), Department of Psychiatry and Behavioral Sciences, , SUNY Upstate Medical University, ; Syracuse, NY USA

[532 ]GRID grid.266100.3, ISNI 0000 0001 2107 4242, University of California, San Diego, ; San Diego, CA USA

[533 ]GRID grid.5603.0, Department of Psychiatry and Psychotherapy, , University Medicine Greifswald, ; Greifswald, Germany

[534 ]GRID grid.267309.9, ISNI 0000 0001 0629 5880, Department of Radiology, , University of Texas Health Science Center at San Antonio, ; San Antonio, TX USA

[535 ]GRID grid.267308.8, ISNI 0000 0000 9206 2401, School of Public Health, , University of Texas Health Science Center at Houston, ; Houston, TX USA

[536 ]GRID grid.11598.34, ISNI 0000 0000 8988 2476, Clinical Division of Neurogeriatrics, Department of Neurology, , Medical University of Graz, ; Graz, Austria

[537 ]GRID grid.164295.d, ISNI 0000 0001 0941 7177, University of Maryland, ; College Park, MD USA

[538 ]GRID grid.267309.9, ISNI 0000 0001 0629 5880, Department of General Medicine, , University of Texas Health Science Center, ; San Antonio, TX USA

[539 ]GRID grid.1002.3, ISNI 0000 0004 1936 7857, Monash University Clayton Campus, ; Mebourne, Victoria Australia

[540 ]GRID grid.7737.4, ISNI 0000 0004 0410 2071, University of Helsinki, ; Helsinki, Finland

[541 ]GRID grid.430503.1, ISNI 0000 0001 0703 675X, University of Colorado Anschutz Medical Center, ; Aurora, CO USA

[542 ]GRID grid.11598.34, ISNI 0000 0000 8988 2476, Medical University of Graz, ; Graz, Austria

[543 ]GRID grid.32224.35, ISNI 0000 0004 0386 9924, Massachusetts General Hospital, Harvard University, ; Cambridge, MA USA

[544 ]GRID grid.27860.3b, ISNI 0000 0004 1936 9684, Imaging of Dementia and Aging (IDeA) Laboratory, Department of Neurology, , University of California, Davis, ; Davis, CA USA

[545 ]University of Staffmail, Edinburgh, UK

[546 ]GRID grid.462010.1, University of Bordeaux, IMN, ; Bordeaux, France

[547 ]GRID grid.410721.1, ISNI 0000 0004 1937 0407, University of Mississippi Medical Center, ; Jackson, MS USA

[548 ]GRID grid.21107.35, ISNI 0000 0001 2171 9311, GeneSTAR Research Program, Department of Neurology, , Johns Hopkins University School of Medicine, ; Baltimore, MD USA

[549 ]GRID grid.17063.33, ISNI 0000 0001 2157 2938, University of Toronto, ; Toronto, Ontario Canada

[550 ]GRID grid.14848.31, ISNI 0000 0001 2292 3357, Departments of Psychiatry & Neuroscience, , Centre Hospitalier Universitaire Saint-Justine, University of Montreal, ; Montreal, Quebec Canada

[551 ]GRID grid.17063.33, ISNI 0000 0001 2157 2938, Hospital for Sick Children, , University of Toronto, ; Toronto, Ontario Canada

[552 ]GRID grid.279946.7, ISNI 0000 0004 0521 0744, Institute for Translational Genomics and Population Sciences, , Los Angeles Biomedical Research Institute and Pediatrics at Harbor-UCLA Medical Center, ; Torrance, CA USA

[553 ]GRID grid.239424.a, ISNI 0000 0001 2183 6745, Boston Medical Center, ; Boston, MA USA

[554 ]GRID grid.11598.34, ISNI 0000 0000 8988 2476, Gottfried Schatz Research Center, Department of Molecular Biology and Biochemistry, , Medical University of Graz, ; Graz, Austria

[555 ]GRID grid.11598.34, ISNI 0000 0000 8988 2476, Clinical Division of Neurogeriatrics, Department of Neurology, , Medical University of Graz, ; Graz, Austria

[556 ]GRID grid.39382.33, ISNI 0000 0001 2160 926X, Departments of Neurology, Molecular & Human Genetics, and Neuroscience and Program in Developmental Biology, , Baylor College of Medicine, ; Houston, TX USA

[557 ]GRID grid.214458.e, ISNI 0000000086837370, Department of Epidemiology, School of Public Health, , University of Michigan, ; Ann Arbor, MI USA

[558 ]GRID grid.17063.33, ISNI 0000 0001 2157 2938, Hospital for Sick Children, , University of Toronto, ; Toronto, Ontario Canada

[559 ]GRID grid.410721.1, ISNI 0000 0004 1937 0407, University of Mississippi Medical Center, ; Jackson, MS USA

[560 ]GRID grid.5603.0, Institute for Community Medicine, , University Medicine Greifswald, ; Greifswald, Germany

[561 ]GRID grid.410711.2, ISNI 0000 0001 1034 1720, University of North Carolina, ; Chapel Hill, NC USA

[562 ]GRID grid.55460.32, ISNI 0000000121548364, University of Texas, ; Austin, TX USA

[563 ]GRID grid.11598.34, ISNI 0000 0000 8988 2476, Clinical Division of Neurogeriatrics, Department of Neurology, , Medical University of Graz, ; Graz, Austria

[564 ]GRID grid.4714.6, ISNI 0000 0004 1937 0626, Karolinska Institute, ; Stockholm, Sweden

[565 ]GRID grid.424247.3, ISNI 0000 0004 0438 0426, German Center for Neurodegenerative Diseases (DZNE), Site Rostock/Greifswald, ; Greifswald, Germany

[566 ]GRID grid.267308.8, ISNI 0000 0000 9206 2401, Institute of Molecular Medicine, , University of Texas Health Science Center at Houston McGovern Medical School, ; Houston, TX USA

Author information

Céline Bellenguez http://orcid.org/0000-0002-1240-7874

Fahri Küçükali http://orcid.org/0000-0002-3835-9639

Najaf Amin http://orcid.org/0000-0002-8944-1771

Peter A. Holmans http://orcid.org/0000-0003-0870-9412

Sven J. van der Lee http://orcid.org/0000-0003-1606-8643

Marcos R. Costa http://orcid.org/0000-0002-4928-2163

Teemu Kuulasmaa http://orcid.org/0000-0002-1795-7314

Qiong Yang http://orcid.org/0000-0002-3658-1375

Itziar de Rojas http://orcid.org/0000-0002-2148-381X

Joshua C. Bis http://orcid.org/0000-0002-3409-1110

Amber Yaqub http://orcid.org/0000-0002-3579-8054

Ivana Prokic http://orcid.org/0000-0002-0370-1473

Julien Chapuis http://orcid.org/0000-0002-5802-2857

Shahzad Ahmad http://orcid.org/0000-0002-8658-3790

Vilmantas Giedraitis http://orcid.org/0000-0003-3423-2021

Pablo Garcia-Gonzalez http://orcid.org/0000-0003-0125-5403

Daniel Alcolea http://orcid.org/0000-0002-3819-3245

Ignacio Alvarez http://orcid.org/0000-0002-8537-3935

Anthoula Tsolaki http://orcid.org/0000-0002-5563-7776

Miquel Baquero http://orcid.org/0000-0002-6861-1831

Ana Belén Pastor http://orcid.org/0000-0001-9637-4688

Claudine Berr http://orcid.org/0000-0001-5254-7655

Valentina Bessi http://orcid.org/0000-0002-6176-3584

Mercè Boada http://orcid.org/0000-0003-2617-3009

Paola Bossù http://orcid.org/0000-0002-1432-0078

Geir Bråthen http://orcid.org/0000-0003-3224-7983

Jan Bressler http://orcid.org/0000-0001-6578-4772

Catherine Bresner http://orcid.org/0000-0003-2673-9762

Henry Brodaty http://orcid.org/0000-0001-9487-6617

Keeley J. Brookes http://orcid.org/0000-0003-2427-2513

Vanessa Burholt http://orcid.org/0000-0002-6789-127X

William S. Bush http://orcid.org/0000-0002-9729-6519

Miguel Calero http://orcid.org/0000-0001-5366-3324

Jaeyoon Chung http://orcid.org/0000-0002-6431-9454

Laura Cervera-Carles http://orcid.org/0000-0003-2286-200X

Emanuele Costantini http://orcid.org/0000-0002-1096-8221

Maria Carolina Dalmasso http://orcid.org/0000-0002-4901-9955

Katia de Paiva Lopes http://orcid.org/0000-0002-0240-0126

Lot D. de Witte http://orcid.org/0000-0002-7235-9958

Stéphanie Debette http://orcid.org/0000-0001-8675-7968

Teodoro del Ser http://orcid.org/0000-0001-9806-7083

Martin Dichgans http://orcid.org/0000-0002-0654-387X

Janine Diehl-Schmid http://orcid.org/0000-0002-7745-1382

Mónica Diez-Fairen http://orcid.org/0000-0003-1882-0309

Srdjan Djurovic http://orcid.org/0000-0002-8140-8061

Carole Dufouil http://orcid.org/0000-0003-2442-4476

Valentina Escott-Price http://orcid.org/0000-0003-1784-5483

Michael Ewers http://orcid.org/0000-0001-5231-1714

Tagliavini Fabrizio http://orcid.org/0000-0003-1039-7315

Tormod Fladby http://orcid.org/0000-0002-9984-9797

Myriam Fornage http://orcid.org/0000-0003-0677-8158

Nick C. Fox http://orcid.org/0000-0002-6660-657X

María J. Bullido http://orcid.org/0000-0002-6477-1117

Lutz Froelich http://orcid.org/0000-0003-1494-0813

Daniela Galimberti http://orcid.org/0000-0002-9284-5953

Jose Maria García-Alberca http://orcid.org/0000-0003-2951-6644

Alison M. Goate http://orcid.org/0000-0002-0576-2472

Antonio González-Pérez http://orcid.org/0000-0001-9771-5982

Emma Green http://orcid.org/0000-0002-8687-5590

Edna Grünblatt http://orcid.org/0000-0001-8505-7265

Vilmundur Gudnason http://orcid.org/0000-0001-5696-0084

Annakaisa Haapasalo http://orcid.org/0000-0003-0959-2957

Janet Harwood http://orcid.org/0000-0002-3225-0069

Stefanie Heilmann-Heimbach http://orcid.org/0000-0003-1057-465X

Martin J. Herrmann http://orcid.org/0000-0001-9970-2122

Henne Holstege http://orcid.org/0000-0002-7688-3087

Geert Jan Biessels http://orcid.org/0000-0001-6862-2496

Xueqiu Jian http://orcid.org/0000-0002-0313-6494

Charlotte Johansson http://orcid.org/0000-0002-5351-1950

Gyungah R. Jun http://orcid.org/0000-0002-3230-8697

Yuriko Kastumata http://orcid.org/0000-0002-0188-8094

Patrick G. Kehoe http://orcid.org/0000-0002-7542-1139

Johannes Kornhuber http://orcid.org/0000-0002-8096-3987

Mary H. Kosmidis http://orcid.org/0000-0001-8790-1220

Carmen Lage http://orcid.org/0000-0003-1703-121X

Lenore Launer http://orcid.org/0000-0002-3238-7612

Chien-Yueh Lee http://orcid.org/0000-0002-4304-974X

Alberto Lleó http://orcid.org/0000-0002-2568-5478

Oscar Lopez http://orcid.org/0000-0002-8546-8256

Adolfo Lopez de Munain http://orcid.org/0000-0002-9509-4032

Kathryn L. Lunetta http://orcid.org/0000-0002-9268-810X

Yiyi Ma http://orcid.org/0000-0002-3609-8877

Catherine A. MacLeod http://orcid.org/0000-0002-9314-7380

Marta Marquié http://orcid.org/0000-0002-0660-0950

Angel Martín Montes http://orcid.org/0000-0002-1694-786X

Simon Mead http://orcid.org/0000-0002-4326-1468

Miguel Medina http://orcid.org/0000-0002-7016-5340

Manuel Menéndez-González http://orcid.org/0000-0002-5218-0774

Merel Mol http://orcid.org/0000-0003-2533-2530

Kevin Morgan http://orcid.org/0000-0002-8217-2396

Markus M. Nöthen http://orcid.org/0000-0002-8770-2464

Carolina Muchnik http://orcid.org/0000-0002-1542-3706

Benedetta Nacmias http://orcid.org/0000-0001-9338-9040

Gael Nicolas http://orcid.org/0000-0001-9391-7800

Børge G. Nordestgaard http://orcid.org/0000-0002-1954-7220

Florence Pasquier http://orcid.org/0000-0001-9880-9788

Pau Pastor http://orcid.org/0000-0002-7493-8777

Gina Peloso http://orcid.org/0000-0002-5355-8636

Alba Pérez-Cordón http://orcid.org/0000-0002-6028-0791

Jordi Pérez-Tur http://orcid.org/0000-0002-9111-1712

Pierre Pericard http://orcid.org/0000-0001-8167-6448

Juan A. Pineda http://orcid.org/0000-0002-3751-0296

Claudia Pisanu http://orcid.org/0000-0002-9151-4319

Danielle Posthuma http://orcid.org/0000-0001-7582-2365

Raquel Puerta http://orcid.org/0000-0002-1191-5893

Olivier Quenez http://orcid.org/0000-0002-8273-8505

Jesper Qvist Thomassen http://orcid.org/0000-0003-3484-9531

Luis M. Real http://orcid.org/0000-0003-4932-7429

Marcel J. T. Reinders http://orcid.org/0000-0002-1148-1562

Christiane Reitz http://orcid.org/0000-0001-8757-7889

Steffi Riedel-Heller http://orcid.org/0000-0003-4321-6090

Eloy Rodriguez-Rodriguez http://orcid.org/0000-0001-7742-677X

Arvid Rongve http://orcid.org/0000-0002-0476-4134

María Eugenia Sáez http://orcid.org/0000-0001-9299-2534

Ingvild Saltvedt http://orcid.org/0000-0002-7897-9808

Pascual Sánchez Juan http://orcid.org/0000-0002-6081-8037

Chloé Sarnowski http://orcid.org/0000-0002-6090-7099

Claudia L. Satizabal http://orcid.org/0000-0002-1115-4430

Jonathan M. Schott http://orcid.org/0000-0003-2059-024X

Geir Selbæk http://orcid.org/0000-0001-6511-8219

Alexey A. Shadrin http://orcid.org/0000-0002-7467-250X

Hilkka Soininen http://orcid.org/0000-0002-2785-9937

Vincenzo Solfrizzi http://orcid.org/0000-0002-8524-0315

Yeunjoo Song http://orcid.org/0000-0002-7452-3731

Oscar Sotolongo-Grau http://orcid.org/0000-0002-9679-0670

Gianfranco Spalletta http://orcid.org/0000-0002-7432-4249

Alessio Squassina http://orcid.org/0000-0001-7415-7607

Eystein Stordal http://orcid.org/0000-0002-2443-7923

Giuseppe Tosto http://orcid.org/0000-0001-7075-8245

Andre Uitterlinden http://orcid.org/0000-0002-7276-3387

Otto Valladares http://orcid.org/0000-0001-8055-2187

Christine Van Broeckhoven http://orcid.org/0000-0003-0183-7665

Jean-Sébastien Vidal http://orcid.org/0000-0001-6770-0720

Jonathan Vogelgsang http://orcid.org/0000-0001-9326-8193

Michael Wagner http://orcid.org/0000-0003-2589-6440

David Wallon http://orcid.org/0000-0002-2634-7198

Jens Wiltfang http://orcid.org/0000-0003-1492-5330

Bob Woods http://orcid.org/0000-0002-6781-651X

Mary Yannakoulia http://orcid.org/0000-0003-2171-7337

Habil Zare http://orcid.org/0000-0001-5902-6238

Xiaoling Zhang http://orcid.org/0000-0001-8237-1857

Lindsay A. Farrer http://orcid.org/0000-0001-5533-4225

Bruce M. Psaty http://orcid.org/0000-0002-7278-2190

Mohsen Ghanbari http://orcid.org/0000-0002-9476-7143

Towfique Raj http://orcid.org/0000-0002-9355-5704

Perminder Sachdev http://orcid.org/0000-0002-9595-3220

Karen Mather http://orcid.org/0000-0003-4143-8941

M. Arfan Ikram http://orcid.org/0000-0003-0372-8585

Magda Tsolaki http://orcid.org/0000-0002-2072-8010

Margaret A. Pericak-Vance http://orcid.org/0000-0001-7283-8804

Philippe Amouyel http://orcid.org/0000-0001-9088-234X

Julie Williams http://orcid.org/0000-0002-4069-0259

Ruth Frikke-Schmidt http://orcid.org/0000-0003-4084-5027

Sudha Seshadri http://orcid.org/0000-0001-6135-2622

Ole A. Andreassen http://orcid.org/0000-0002-4461-3568

Kristel Sleegers http://orcid.org/0000-0002-0283-2332

Cornelia M. van Duijn http://orcid.org/0000-0002-2374-9204

Rebecca Sims http://orcid.org/0000-0002-3885-1199

Wiesje M. van der Flier http://orcid.org/0000-0001-8766-6224

Alfredo Ramirez http://orcid.org/0000-0003-4991-763X

Jean-Charles Lambert http://orcid.org/0000-0003-0829-7817

Article

Publisher ID: 1024

DOI: 10.1038/s41588-022-01024-z

PMC ID: 9005347

PubMed ID: 35379992

SO-VID: 3e789c27-0444-4051-8a2f-332f4fd963d4

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Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

History

Date received : 16 June 2021

Date accepted : 27 January 2022

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ScienceOpen disciplines: Genetics

Keywords: alzheimer's disease,genome-wide association studies

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ScienceOpen disciplines: Genetics

Keywords: alzheimer's disease, genome-wide association studies

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New insights into the genetic etiology of Alzheimer’s disease and related dementias

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