Detailed stratified GWAS analysis for severe COVID-19 in four European populations.

Address: Institute of Clinical Molecular Biology, Christian-Albrechts-University, Kiel, Germany.; Novo Nordisk Foundation Center for Protein Research, Disease Systems Biology, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark.; Life Science Centre, Institute of Biotechnology, Vilnius University, Vilnius, Lithuania.; Institut de Biotecnologia i de Biomedicina, Universitat Autònoma de Barcelona, Barcelona, Spain.; Hospital Universitario Virgen del Rocío de Sevilla, Sevilla, Spain.; Centro de Investigación Biomédica en Red en Enfermedades Hepáticas y Digestivas (CIBEREHD), Instituto de Salud Carlos III (ISCIII), Madrid, Spain.; Instituto de Biomedicina de Sevilla (IBIS), Sevilla, Spain.; Institute for Medical Microbiology and Hospital Epidemiology, Hannover Medical School, Hannover, Germany.; Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA.; Division of Rheumatology, Inflammation and Immunity, Brigham and Women's Hospital and Harvard Medical School, Boston, MA, USA.; Division of Genetics, Department of Medicine, Brigham and Women's Hospital, Boston, MA, USA.; Department of Biomedical Informatics, Harvard Medical School, Boston, MA, USA.; Center for Data Sciences, Brigham and Women's Hospital, Boston, MA, USA.; Research Group for Evolutionary Immunogenomics, Max Planck Institute for Evolutionary Biology, Plön, Germany.; Research Unit for Evolutionary Immunogenomics, Department of Biology, University of Hamburg, Hamburg, Germany.; Research Department, Stavanger University Hospital, Stavanger, Norway.; Randaberg Municipality, Stavanger, Norway.; Division of Health Data and Digitalization, Department of Genetics and Bioinformatics (HDGB), Norwegian Institute of Public Health, Oslo, Norway.; Department of Quality and Health Technology, Faculty of Health Sciences, University of Stavanger, Stavanger, Norway.; Department of Intensive Care, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of Alcalá, Madrid, Spain.; Ibs.Granada Instituto de Investigación Biosanitaria, Granada, Spain.; Microbiology Unit. Hospital Universitario Clinico San Cecilio, Granada, Spain.; Clinical Biochemistry Department, Osakidetza Basque Health Service, Donostialdea Integrated Health Organisation, San Sebastian, Spain.; Vall d'Hebron Institut de Recerca (VHIR), Vall d'Hebron Hospital Universitari, Barcelona, Spain.; Liver Unit, Department of Internal Medicine, Hospital Universitari Vall d'Hebron, Vall d'Hebron Barcelona Hospital Campus, Barcelona, Spain.; Research Center and Memory Clinic, Ace Alzheimer Center Barcelona, Universitat Internacional de Catalunya, Barcelona, Spain.; CIBERNED, Network Center for Biomedical Research in Neurodegenerative Diseases, National Institute of Health Carlos III, Madrid, Spain.; Department of Biochemistry, University Hospital Vall d'Hebron, Barcelona, Spain.; IRCCS Humanitas Research Hospital, Milan, Italy.; Department of Biomedical Sciences, Humanitas University, Milan, Italy.; University of Milan, Milan, Italy.; Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico, Milan, Italy.; Department of Acute Medicine, Oslo University Hospital, Oslo, Norway.; Institute of Clinical Medicine, University of Oslo, Oslo, Norway.; Charite Universitätsmedizin Berlin, Berlin, Germany.; European Reference Network on Hepatological Diseases (ERN RARE-LIVER), San Gerardo Hospital, Monza, Italy.; Division of Gastroenterology, School of Medicine and Surgery, Center for Autoimmune Liver Diseases, University of Milano-Bicocca, Milan, Italy.; Division of Neurogenetics and Molecular Psychiatry, Department of Psychiatry and Psychotherapy, Faculty of Medicine and University Hospital Cologne, University of Cologne, Cologne, Germany.; Department of Neurodegenerative Diseases and Geriatric Psychiatry, University Hospital Bonn, Medical Faculty, Bonn, Germany.; Cologne Excellence Cluster on Cellular Stress Responses in Aging-Associated Diseases (CECAD), Faculty of Medicine and University Hospital Cologne, University of Cologne, Cologne, Germany.; German Center for Neurodegenerative Diseases (DZNE), Bonn, Germany.; Department of Psychiatry, Glenn Biggs Institute for Alzheimer's and Neurodegenerative Diseases, San Antonio, TX, USA.; Department of Anesthesiology and Intensive Care, University Hospital of North Norway, Tromsø, Norway.; Pediatric Departement, Centro Tettamanti-European Reference Network (ERN) PaedCan, EuroBloodNet, MetabERN-University of Milano-Bicocca-Fondazione MBBM/Ospedale, San Gerardo, Italy.; Biochemistry Department, Echevarne Laboratory, Barcelona, Spain.; Institute for Molecular Medicine Finland, University of Helsinki, Helsinki, Finland.; Centre for Multidisciplinary Research in Health Science (MACH), University of Milan, Milan, Italy.; Klinik für Innere Medizin I, Universitätsklinikum Schleswig-Holstein, Kiel, Germany.; Department of Microbiology, Oslo University Hospital, Oslo, Norway.; Department of Child and Adolescent Psychiatry, University Hospital Essen, University of Duisburg-Essen, Essen, Germany.; Genomes for Life-GCAT Lab, Germans Trias i Pujol Research Institute (IGTP), Badalona, Spain.; Laboratory of Microbiology, San Gerardo Hospital, Monza, Italy.; Department of Infectious diseases, Oslo University Hospital, Oslo, Norway.; Humanitas Gavazzeni-Castelli, Bergamo, Italy.; Microbiology Department, Hospital Universitari Vall d'Hebron, Barcelona, Spain.; Universitat Autònoma de Barcelona, Bellatera, Spain.; Department of Respiratory Diseases, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of Alcalá, Madrid, Spain.; Centro de Investigación Biomédica en Red en Enfermedades Respiratorias (CIBERES), University of Alcalá, Madrid, Spain.; Institute of Human Genetics, University of Bonn School of Medicine & University Hospital Bonn, Bonn, Germany.; Department of Gastroenterology, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of Alcalá, Madrid, Spain.; Department of Medicine I, Gastroenterology, Hepatology and Endocrinology, Medical University of Innsbruck, Innsbruck, Austria.; Christian Doppler Laboratory of Iron and Phosphate Biology at the Department of Medicine I, Medical University of Innsbruck, Innsbruck, Austria.; Department of Gastroenterology, Hepatology and Infectious Diseases, University Hospital Duesseldorf, Medical Faculty Heinrich Heine University, Duesseldorf, Germany.; Department of Respiratory Medicine and Allergology, University Hospital, Goethe University, Frankfurt am Main, Germany.; Institute of Genomic Statistics and Bioinformatics, University Hospital Bonn, Medical Faculty University of Bonn, Bonn, Germany.; Centro de Investigación Biomédica en Red de Enfermedades Respiratorias (CIBERES), Madrid, Spain.; Hospital Clinic, University of Barcelona, and IDIBAPS, Barcelona, Spain.; University of Sevilla, Sevilla, Spain.; Centro de Investigación Biomédica en Red de Epidemiología y Salud Pública (CIBERESP), Madrid, Spain.; Consejo Superior de Investigaciones Científicas, Sevilla, Spain.; Department of Infectious Diseases, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of Alcalá, Madrid, Spain.; Department of Transfusion Medicine and Haematology Laboratory, San Gerardo Hospital, Monza, Italy.; Department of Internal Medicine V-Pneumology, Allergology and Intensive Care Medicine, University Hospital Saarland, Homburg, Germany.; Department of Internal Medicine II, School of Medicine, University Hospital rechts der Isar, Technical University of Munich, Munich, Germany.; Institute of Translational Medicine, Private University in the Principality of Liechtenstein (UFL), Triesen, Liechtenstein.; Respiratory Medicine and International Health, University of Lübeck, Lübeck, Germany.; Division of Clinical Infectious Diseases, Research Center Borstel, Borstel, Germany.; Clinical Tuberculosis Unit, German Center for Infection Research (DZIF), Borstel, Germany.; Department I of Internal Medicine, Faculty of Medicine and University Hospital Cologne, University of Cologne, Cologne, Germany.; Center for Molecular Medicine Cologne (CMMC), University of Cologne, Cologne, Germany.; German Center for Infection Research (DZIF), Cologne, Germany.; Humanitas Clinical and Research Center, IRCCS, Milan, Italy.; Osakidetza Basque Health Service, Basurto University Hospital, Respiratory Service, Bilbao, Spain.; Department of Clinical and Molecular Medicine, Faculty of Medicine and Health Sciences, Norwegian University of Science and Technology, Trondheim, Norway.; Department of Medicine, Møre and Romsdal Hospital Trust, Ålesund, Norway.; Department of Internal Medicine II, Medical University of Innsbruck, Innsbruck, Austria.; Department of Anesthesiology and Critical Care, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of Alcalá, Madrid, Spain.; Immunohematology Department, Banc de Sang i Teixits, Autonomous University of Barcelona, Barcelona, Spain.; Department of Medical Sciences, Università degli Studi di Torino, Turin, Italy.; Osakidetza Basque Health Service, Galdakao Hospital, Respiratory Service, Galdakao, Spain.; Biocruces Bizkaia Health Research Institute, Barakaldo, Spain.; Department of Infectious Diseases, E.O. Ospedali Galliera, Genova, Italy.; Geminicenter for Sepsis Research, Institute of Circulation and Medical Imaging (ISB), NTNU, Trondheim, Norway.; Clinic of Anesthesia and Intensive Care, St. Olavs Hospital, Trondheim University Hospital, Trondheim, Norway.; Accident and Emergency and Emergency Medicine Unit, San Gerardo Hospital, Monza, Italy.; Centro de Investigación Biomédica en Red en Enfermedades Infecciosas (CIBERINFEC), Instituto de Salud Carlos III (ISCIII), Madrid, Spain.; Histocompatibilidad y Biologia Molecular, Centro de Transfusion de Madrid, Madrid, Spain.; Department of Pathophysiology and Transplantation, Dino Ferrari Center, University of Milan, Milan, Italy.; Neurology Unit, IRCCS Fondazione Ca' Granda Ospedale Maggiore Policlinico, Milan, Italy.; Department of Pathophysiology and Transplantation, University of Milan, Milan, Italy.; Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico, Angelo Bianchi Bonomi Hemophilia and Thrombosis Center, Milan, Italy.; Department of Tropical Medicine, Bernhard Nocht Institute for Tropical Medicine, Hamburg, Germany.; Department of Medicine I, University Medical Centre Hamburg-Eppendorf, Hamburg, Germany.; Respiratory Unit, Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico, Milan, Italy.; Department of Pathophysiology and Transplantation, Università degli Studi di Milano, Milan, Italy.; Internal Medicine Department, Virgen del Rocio University Hospital, Sevilla, Spain.; Biochemistry Department, University Hospital Vall d'Hebron, Barcelona, Spain.; Emergency Department, University Hospital Regensburg, Regensburg, Germany.; Department for Infectious Diseases and Infection Control, University Hospital Regensburg, Regensburg, Germany.; Department Emergency, Anesthesia and Intensive Care, San Gerardo Hospital, Monza, Italy.; School of Medicine and Surgery, University of Milano-Bicocca, Milan, Italy.; Fondazione Grigioni per il Morbo di Parkinson and Parkinson Institute, ASST Gaetano Pini-CTO, Milan, Italy.; Acute Geriatric Unit, San Gerardo Hospital, Monza, Italy.; Neurointensive Care Unit, San Gerardo Hospital, Monza, Italy.; Department of Anesthesiology, Intensive Care Medicine and Pain Therapy, University Hospital Frankfurt, Frankfurt am Main, Germany.; Institute of Human Genetics, Medical Faculty, RWTH Aachen University, Aachen, Germany.; Department of Pneumology, Hannover Medical School, Hannover, Germany.; Department of Medical Microbiology, Clinic of Laboratory Medicine, St. Olavs hospital, Trondheim, Norway.; Department of Infectious Diseases, St. Olavs Hospital, Trondheim University Hospital, Trondheim, Norway.; Department of Clinical and Molecular Medicine, NTNU, Trondheim, Norway.; German Centre for Infection Research (DZIF), Cologne, Germany.; Cologne Center for Genomics (CCG), University of Cologne, Cologne, Germany.; Digestive Diseases Unit, Virgen del Rocio University Hospital, Institute of Biomedicine of Seville, University of Seville, Seville, Spain.; Institute of Parasitology and Biomedicine Lopez-Neyra, CSIC, Granada, Spain.; Institute for Cardiogenetics, University of Lübeck, Lübeck, Germany.; German Research Center for Cardiovascular Research, Lübeck, Germany.; University Heart Center Lübeck, Lübeck, Germany.; Department of Liver and Gastrointestinal Diseases, Biodonostia Health Research Institute-Donostia University Hospital, University of the Basque Country (UPV/EHU), CIBERehd, Ikerbasque, San Sebastian, Spain.; Ikerbasque, Basque Foundation for Science, Bilbao, Spain.; Respiratory ICU, Institut Clínic Respiratory, Hospital Clinic, University of Barcelona, and IDIBAPS, Barcelona, Spain.; Bioinformatics Area, Fundación Progreso y Salud, and Institute of Biomedicine of Sevilla (IBIS), Sevilla, Spain.; Department of Research, Ostfold Hospital Trust, Gralum, Norway.; Germans Trias i Pujol Research Institute (IGTP), Badalona, Spain.; Department of Genetics & Epigenetics, Saarland University, Saarbrücken, Germany.; Department of Infectious Diseases, Hospital Universitario Clinico San Cecilio, Granada, Spain.; Infectious Diseases Service, Osakidetza, Biocruces Bizkaia Health Research Institute, Barakaldo, Spain.; Department of Research, St. Olav Hospital, Trondheim University Hospital, Trondheim, Norway.; Medical Department, Drammen Hospital, Vestre Viken Hospital Trust, Drammen, Norway.; Research Center Borstel, BioMaterialBank Nord, Borstel, Germany.; German Center for Lung Research (DZL), Airway Research Center North (ARCN), Borstel, Germany.; Popgen 2.0 Network (P2N), Kiel, Germany.; Department of Clinical Science, University of Bergen, Bergen, Norway.; Biodonostia Health Research Institute, Donostia University Hospital, San Sebastian, Spain.; Analytic and Translational Genetics Unit, Massachusetts General Hospital, Boston, MA, USA.; Stanley Center for Psychiatric Research & Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA.; Clinic of Medicine and Rehabilitation, Levanger Hospital, Nord-Trondelag Hospital Trust, Levanger, Norway.; HLA Laboratory, E.O. Ospedali Galliera, Genova, Italy.; ISGlobal, Barcelona, Spain.; Universitat Pompeu Fabra (UPF), Barcelona, Spain.; IMIM (Hospital del Mar Medical Research Institute), Barcelona, Spain.; Department of Anesthesiology and Intensive Care Medicine, University Hospital Essen, University Duisburg-Essen, Essen, Germany.; Stefan-Morsch-Stiftung, Birkenfeld, Germany.; Osakidetza, OSI Donostialdea, Altza Primary Care, Biodonostia Health Research Institute, San Sebastián, Spain.; Internal Medicine Department, Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico, Milan, Italy.; Center of Bioinformatics, Biostatistics and Bioimaging, School of Medicine and Surgery, University of Milano-Bicocca, Milan, Italy.; Department of Internal Medicine, Infectious Diseases, University Hospital Frankfurt & Goethe University Frankfurt, Frankfurt am Main, Germany.; Phase 1 Research Centre, ASST Monza, School of Medicine and Surgery, University of Milano-Bicocca, Monza, Italy.; Division of Surgery, Inflammatory Diseases and Transplantation, Research Institute for Internal Medicine, Oslo University Hospital Rikshospitalet and University of Oslo, Oslo, Norway.; Division of Surgery, Inflammatory Diseases and Transplantation, Department of Transplantation Medicine, Norwegian PSC Research Center, Oslo University Hospital Rikshospitalet, Oslo, Norway.; Department of Gastroenterology, Hepatology and Endocrinology, Hannover Medical School, Hannover, Germany.; Gastrointestinal Genetics Lab, CIC bioGUNE-BRTA, Derio, Spain.; Department of Medicine and Surgery, LUM University, Casamassima, Italy.; Department of Pneumology and Intensive Care Medicine, University Hospital Aachen, Aachen, Germany.; Munich Clinic Schwabing, Academic Teaching Hospital, Ludwig-Maximilians-University (LMU), Munich, Germany.; Division of Intensive Care and Emergency Medicine, Department of Internal Medicine, Medical University Innsbruck, Innsbruck, Austria.; University Hospital Schleswig-Holstein, Kiel, Germany.; Department of Human Genetics, Center of Human and Molecular Biology, University Hospital Saarland, Homburg/Saar, Germany.; Department of Internal Medicine I, RWTH Aachen University Hospital, Aachen, Germany.; Department of Anesthesiology, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), Madrid, Spain.; Faculty of Medicine and University Hospital Cologne, Clinical Trials Centre Cologne (ZKS Köln), University of Cologne, Cologne, Germany.; Department of Infectious Diseases, University Hospital Essen, University Duisburg-Essen, Essen, Germany.; Gastroenterology Unit, Fondazione IRCCS Casa Sollievo della Sofferenza, San Giovanni Rotondo, Italy.; Pulmonary Unit, San Gerardo Hospital, Monza, Italy.; Infectious Diseases Unit, San Gerardo Hospital, Monza, Italy.; Intensive Care Department, Vall d'Hebron University Hospital, SODIR-VHIR research group, Barcelona, Spain.; Department of Medicine, Møre & Romsdal Hospital Trust, Molde, Norway.; Institute of Medical Virology, University Hospital Frankfurt, Goethe University, Frankfurt am Main, Germany.; German Centre for Infection Research (DZIF), Frankfurt am Main, Germany.; Zentrum für Humangenetik Regensburg, Regensburg, Germany.; Department of Neurology, Bezirksklinikum Regensburg, University of Regensburg, Regensburg, Germany.; Hannover Unified Biobank, Hannover Medical School, Hannover, Germany.; Centre for Genetics and Genomics Versus Arthritis, Centre for Musculoskeletal Research, Manchester Academic Health Science Centre, The University of Manchester, Manchester, UK.; School of Biological Sciences, Monash University, Clayton, VIC, Australia.; Department of Microbiology, University Hospital Vall d'Hebron, Barcelona, Spain.; Autonoma University of Barcelona, Barcelona, Spain.; Division for Cancer Medicine, Surgery and Transplantation, Section for Gastroenterology, Department of Transplantation Medicine, Oslo University Hospital Rikshospitalet, Oslo, Norway.; Biochemistry Unit, Hospital Universitario Clinico San Cecilio, Granada, Spain.; Berlin Institute of Health, Charite Universitätsmedizin Berlin, Berlin Germany.; Institute of Virology, Technical University Munich/Helmholtz Zentrum München, Munich, Germany.; German Center for Infection Research (DZIF), Munich, Germany.; Department of Infectious Diseases, University Hospital of North Norway, Tromsø, Norway.; Faculty of Health Sciences, UIT The Arctic University of Norway, Norway.; Catalan Institute of Oncology (ICO), Barcelona, Spain.; Bellvitge Biomedical Research Institute (IDIBELL), Barcelona, Spain.; Universitat de Barcelona (UB), Barcelona, Spain.; Faculty of Medicine and University Hospital Cologne, Institute for Transfusion Medicine, University of Cologne, Cologne, Germany.; Institute of Transfusionsmedicine, University Hospital Schleswig-Holstein (UKSH), Kiel, Germany.; Institute of Immunology, Christian-Albrechts-University of Kiel and UKSH Schleswig-Holstein, Kiel, Germany.; Institute of Psychiatric Phenomics and Genomics, University Medical Center, University of Munich, Munich, Germany.; Department of Psychiatry, University Medical Center, University of Munich, Munich, Germany.; European Foundation for the Study of Chronic Liver Failure (EF-CLIF), Barcelona, Spain.; ICREA, Barcelona, Spain.; University Hospital Schleswig-Holstein (UKSH), Kiel, Germany.