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

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  • Frauke Degenhardt
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • David Ellinghaus
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Simonas Juzenas
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Jon Lerga-Jaso
    Institut de Biotecnologia i de Biomedicina, Universitat Autònoma de Barcelona , Barcelona , Spain
  • Mareike Wendorff
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Douglas Maya-Miles
    Hospital Universitario Virgen del Rocío de Sevilla , Sevilla , Spain
  • Florian Uellendahl-Werth
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Hesham ElAbd
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Malte C Rühlemann
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Jatin Arora
    Program in Medical and Population Genetics, Broad Institute of MIT and Harvard , Cambridge, MA , USA
  • Onur Özer
    Research Group for Evolutionary Immunogenomics, Max Planck Institute for Evolutionary Biology , Plön , Germany
  • Ole Bernt Lenning
    Research Department, Stavanger University Hospital , Stavanger , Norway
  • Ronny Myhre
    Division of Health Data and Digitalization , Department of Genetics and Bioinformatics (HDGB), , Oslo , Norway
  • May Sissel Vadla
    Randaberg Municipality , Stavanger , Norway
  • Eike M Wacker
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Lars Wienbrandt
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Aaron Blandino Ortiz
    Department of Intensive Care, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of Alcalá , Madrid , Spain
  • Adolfo de Salazar
    Ibs.Granada Instituto de Investigación Biosanitaria , Granada , Spain
  • Adolfo Garrido Chercoles
    Clinical Biochemistry Department, Osakidetza Basque Health Service, Donostialdea Integrated Health Organisation , San Sebastian , Spain
  • Adriana Palom
    Vall d’Hebron Institut de Recerca (VHIR), Vall d’Hebron Hospital Universitari , Barcelona , Spain
  • Agustín Ruiz
    Research Center and Memory Clinic, Ace Alzheimer Center Barcelona, Universitat Internacional de Catalunya , Barcelona , Spain
  • Alba-Estela Garcia-Fernandez
    Department of Biochemistry, University Hospital Vall d’Hebron , Barcelona , Spain
  • Albert Blanco-Grau
    Department of Biochemistry, University Hospital Vall d’Hebron , Barcelona , Spain
  • Alberto Mantovani
    IRCCS Humanitas Research Hospital , Milan , Italy
  • Alberto Zanella
    University of Milan , Milan , Italy
  • Aleksander Rygh Holten
    Department of Acute Medicine, Oslo University Hospital , Oslo , Norway
  • Alena Mayer
    Charite Universitätsmedizin Berlin , Berlin , Germany
  • Alessandra Bandera
    University of Milan , Milan , Italy
  • Alessandro Cherubini
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Alessandro Protti
    IRCCS Humanitas Research Hospital , Milan , Italy
  • Alessio Aghemo
    IRCCS Humanitas Research Hospital , Milan , Italy
  • Alessio Gerussi
    European Reference Network on Hepatological Diseases (ERN RARE-LIVER), San Gerardo Hospital , Monza , Italy
  • Alfredo Ramirez
    Division of Neurogenetics and Molecular Psychiatry , Department of Psychiatry and Psychotherapy, , Cologne , Germany
  • Alice Braun
    Charite Universitätsmedizin Berlin , Berlin , Germany
  • Almut Nebel
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Ana Barreira
    Liver Unit , Department of Internal Medicine, , Barcelona , Spain
  • Ana Lleo
    IRCCS Humanitas Research Hospital , Milan , Italy
  • Ana Teles
    Research Group for Evolutionary Immunogenomics, Max Planck Institute for Evolutionary Biology , Plön , Germany
  • Anders Benjamin Kildal
    Department of Anesthesiology and Intensive Care, University Hospital of North Norway , Tromsø , Norway
  • Andrea Biondi
    Pediatric Departement, Centro Tettamanti-European Reference Network (ERN) PaedCan, EuroBloodNet, MetabERN-University of Milano-Bicocca-Fondazione MBBM/Ospedale , San Gerardo , Italy
  • Andrea Caballero-Garralda
    Biochemistry Department, Echevarne Laboratory , Barcelona , Spain
  • Andrea Ganna
    Institute for Molecular Medicine Finland, University of Helsinki , Helsinki , Finland
  • Andrea Gori
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Andreas Glück
    Klinik für Innere Medizin I, Universitätsklinikum Schleswig-Holstein , Kiel , Germany
  • Andreas Lind
    Department of Microbiology, Oslo University Hospital , Oslo , Norway
  • Anja Tanck
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Anke Hinney
    Department of Child and Adolescent Psychiatry, University Hospital Essen, University of Duisburg-Essen , Essen , Germany
  • Anna Carreras Nolla
    Genomes for Life-GCAT Lab, Germans Trias i Pujol Research Institute (IGTP) , Badalona , Spain
  • Anna Ludovica Fracanzani
    University of Milan , Milan , Italy
  • Anna Peschuck
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Annalisa Cavallero
    Laboratory of Microbiology, San Gerardo Hospital , Monza , Italy
  • Anne Ma Dyrhol-Riise
    Institute of Clinical Medicine, University of Oslo , Oslo , Norway
  • Antonella Ruello
    Humanitas Gavazzeni-Castelli , Bergamo , Italy
  • Antonio Julià
    Vall d’Hebron Institut de Recerca (VHIR), Vall d’Hebron Hospital Universitari , Barcelona , Spain
  • Antonio Muscatello
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Antonio Pesenti
    University of Milan , Milan , Italy
  • Antonio Voza
    IRCCS Humanitas Research Hospital , Milan , Italy
  • Ariadna Rando-Segura
    Microbiology Department, Hospital Universitari Vall d'Hebron , Barcelona , Spain
  • Aurora Solier
    Department of Respiratory Diseases, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of Alcalá , Madrid , Spain
  • Axel Schmidt
    Institute of Human Genetics, University of Bonn School of Medicine & University Hospital Bonn , Bonn , Germany
  • Beatriz Cortes
    Genomes for Life-GCAT Lab, Germans Trias i Pujol Research Institute (IGTP) , Badalona , Spain
  • Beatriz Mateos
    Centro de Investigación Biomédica en Red en Enfermedades Hepáticas y Digestivas (CIBEREHD), Instituto de Salud Carlos III (ISCIII) , Madrid , Spain
  • Beatriz Nafria-Jimenez
    Clinical Biochemistry Department, Osakidetza Basque Health Service, Donostialdea Integrated Health Organisation , San Sebastian , Spain
  • Benedikt Schaefer
    Department of Medicine I , Gastroenterology, Hepatology and Endocrinology, , Innsbruck , Austria
  • Björn Jensen
    Department of Gastroenterology , Hepatology and Infectious Diseases, , Duesseldorf , Germany
  • Carla Bellinghausen
    Department of Respiratory Medicine and Allergology, University Hospital, Goethe University , Frankfurt am Main , Germany
  • Carlo Maj
    Institute of Genomic Statistics and Bioinformatics, University Hospital Bonn, Medical Faculty University of Bonn , Bonn , Germany
  • Carlos Ferrando
    Centro de Investigación Biomédica en Red de Enfermedades Respiratorias (CIBERES) , Madrid , Spain
  • Carmen de la Horra
    Hospital Universitario Virgen del Rocío de Sevilla , Sevilla , Spain
  • Carmen Quereda
    Department of Infectious Diseases, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of Alcalá , Madrid , Spain
  • Carsten Skurk
    Charite Universitätsmedizin Berlin , Berlin , Germany
  • Charlotte Thibeault
    Charite Universitätsmedizin Berlin , Berlin , Germany
  • Chiara Scollo
    Department of Transfusion Medicine and Haematology Laboratory, San Gerardo Hospital , Monza , Italy
  • Christian Herr
    Department of Internal Medicine V—Pneumology , Allergology and Intensive Care Medicine, , Homburg , Germany
  • Christoph D Spinner
    Department of Internal Medicine II, School of Medicine, University Hospital rechts der Isar, Technical University of Munich , Munich , Germany
  • Christoph Gassner
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Christoph Lange
    Respiratory Medicine and International Health, University of Lübeck , Lübeck , Germany
  • Cinzia Hu
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Cinzia Paccapelo
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Clara Lehmann
    Department I of Internal Medicine, Faculty of Medicine and University Hospital Cologne, University of Cologne , Cologne , Germany
  • Claudio Angelini
    Humanitas Clinical and Research Center, IRCCS , Milan , Italy
  • Claudio Cappadona
    Department of Biomedical Sciences, Humanitas University , Milan , Italy
  • Clinton Azuure
    Research Group for Evolutionary Immunogenomics, Max Planck Institute for Evolutionary Biology , Plön , Germany
  • Cristiana Bianco
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Cristina Cea
    Department of Biochemistry, University Hospital Vall d’Hebron , Barcelona , Spain
  • Cristina Sancho
    Osakidetza Basque Health Service, Basurto University Hospital, Respiratory Service , Bilbao , Spain
  • Dag Arne Lihaug Hoff
    Department of Clinical and Molecular Medicine, Faculty of Medicine and Health Sciences, Norwegian University of Science and Technology , Trondheim , Norway
  • Daniela Galimberti
    University of Milan , Milan , Italy
  • Daniele Prati
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • David Haschka
    Department of Internal Medicine II, Medical University of Innsbruck , Innsbruck , Austria
  • David Jiménez
    Department of Respiratory Diseases, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of Alcalá , Madrid , Spain
  • David Pestaña
    Department of Anesthesiology and Critical Care, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of Alcalá , Madrid , Spain
  • David Toapanta
    Hospital Clinic, University of Barcelona, and IDIBAPS , Barcelona , Spain
  • Eduardo Muñiz-Diaz
    Immunohematology Department, Banc de Sang i Teixits, Autonomous University of Barcelona , Barcelona , Spain
  • Elena Azzolini
    IRCCS Humanitas Research Hospital , Milan , Italy
  • Elena Sandoval
    Hospital Clinic, University of Barcelona, and IDIBAPS , Barcelona , Spain
  • Eleonora Binatti
    European Reference Network on Hepatological Diseases (ERN RARE-LIVER), San Gerardo Hospital , Monza , Italy
  • Elio Scarpini
    University of Milan , Milan , Italy
  • Elisa T Helbig
    Charite Universitätsmedizin Berlin , Berlin , Germany
  • Elisabetta Casalone
    Department of Medical Sciences, Università degli Studi di Torino , Turin , Italy
  • Eloisa Urrechaga
    Osakidetza Basque Health Service, Galdakao Hospital, Respiratory Service , Galdakao , Spain
  • Elvezia Maria Paraboschi
    IRCCS Humanitas Research Hospital , Milan , Italy
  • Emanuele Pontali
    Department of Infectious Diseases, E.O. Ospedali Galliera , Genova , Italy
  • Enric Reverter
    Hospital Clinic, University of Barcelona, and IDIBAPS , Barcelona , Spain
  • Enrique J Calderón
    Hospital Universitario Virgen del Rocío de Sevilla , Sevilla , Spain
  • Enrique Navas
    Department of Infectious Diseases, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of Alcalá , Madrid , Spain
  • Erik Solligård
    Geminicenter for Sepsis Research, Institute of Circulation and Medical Imaging (ISB), NTNU , Trondheim , Norway
  • Ernesto Contro
    Accident and Emergency and Emergency Medicine Unit, San Gerardo Hospital , Monza , Italy
  • Eunate Arana-Arri
    Biocruces Bizkaia Health Research Institute , Barakaldo , Spain
  • Fátima Aziz
    Hospital Clinic, University of Barcelona, and IDIBAPS , Barcelona , Spain
  • Federico Garcia
    Ibs.Granada Instituto de Investigación Biosanitaria , Granada , Spain
  • Félix García Sánchez
    Histocompatibilidad y Biologia Molecular, Centro de Transfusion de Madrid , Madrid , Spain
  • Ferruccio Ceriotti
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Filippo Martinelli-Boneschi
    Department of Pathophysiology and Transplantation, Dino Ferrari Center, University of Milan , Milan , Italy
  • Flora Peyvandi
    Department of Pathophysiology and Transplantation, University of Milan , Milan , Italy
  • Florian Kurth
    Charite Universitätsmedizin Berlin , Berlin , Germany
  • Francesco Blasi
    Respiratory Unit, Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Francesco Malvestiti
    University of Milan , Milan , Italy
  • Francisco J Medrano
    Instituto de Biomedicina de Sevilla (IBIS) , Sevilla , Spain
  • Francisco Mesonero
    Centro de Investigación Biomédica en Red en Enfermedades Hepáticas y Digestivas (CIBEREHD), Instituto de Salud Carlos III (ISCIII) , Madrid , Spain
  • Francisco Rodriguez-Frias
    Centro de Investigación Biomédica en Red en Enfermedades Hepáticas y Digestivas (CIBEREHD), Instituto de Salud Carlos III (ISCIII) , Madrid , Spain
  • Frank Hanses
    Emergency Department, University Hospital Regensburg , Regensburg , Germany
  • Fredrik Müller
    Institute of Clinical Medicine, University of Oslo , Oslo , Norway
  • Georg Hemmrich-Stanisak
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Giacomo Bellani
    Department Emergency , Anesthesia and Intensive Care, , Monza , Italy
  • Giacomo Grasselli
    University of Milan , Milan , Italy
  • Gianni Pezzoli
    Fondazione Grigioni per il Morbo di Parkinson and Parkinson Institute, ASST Gaetano Pini-CTO , Milan , Italy
  • Giorgio Costantino
    University of Milan , Milan , Italy
  • Giovanni Albano
    Humanitas Gavazzeni-Castelli , Bergamo , Italy
  • Giulia Cardamone
    Department of Biomedical Sciences, Humanitas University , Milan , Italy
  • Giuseppe Bellelli
    School of Medicine and Surgery, University of Milano-Bicocca , Milan , Italy
  • Giuseppe Citerio
    School of Medicine and Surgery, University of Milano-Bicocca , Milan , Italy
  • Giuseppe Foti
    Department Emergency , Anesthesia and Intensive Care, , Monza , Italy
  • Giuseppe Lamorte
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Giuseppe Matullo
    Department of Medical Sciences, Università degli Studi di Torino , Turin , Italy
  • Guido Baselli
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Hayato Kurihara
    Humanitas Clinical and Research Center, IRCCS , Milan , Italy
  • Holger Neb
    Department of Anesthesiology , Intensive Care Medicine and Pain Therapy, , Frankfurt am Main , Germany
  • Ilaria My
    IRCCS Humanitas Research Hospital , Milan , Italy
  • Ingo Kurth
    Institute of Human Genetics, Medical Faculty, RWTH Aachen University , Aachen , Germany
  • Isabel Hernández
    Research Center and Memory Clinic, Ace Alzheimer Center Barcelona, Universitat Internacional de Catalunya , Barcelona , Spain
  • Isabell Pink
    Department of Pneumology, Hannover Medical School , Hannover , Germany
  • Itziar de Rojas
    Research Center and Memory Clinic, Ace Alzheimer Center Barcelona, Universitat Internacional de Catalunya , Barcelona , Spain
  • Iván Galván-Femenia
    Genomes for Life-GCAT Lab, Germans Trias i Pujol Research Institute (IGTP) , Badalona , Spain
  • Jan Cato Holter
    Institute of Clinical Medicine, University of Oslo , Oslo , Norway
  • Jan Egil Afset
    Osakidetza Basque Health Service, Basurto University Hospital, Respiratory Service , Bilbao , Spain
  • Jan Heyckendorf
    Respiratory Medicine and International Health, University of Lübeck , Lübeck , Germany
  • Jan Kässens
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Jan Kristian Damås
    Department of Infectious Diseases, St. Olavs Hospital, Trondheim University Hospital , Trondheim , Norway
  • Jan Rybniker
    Department I of Internal Medicine, Faculty of Medicine and University Hospital Cologne, University of Cologne , Cologne , Germany
  • Janine Altmüller
    Cologne Center for Genomics (CCG), University of Cologne , Cologne , Germany
  • Javier Ampuero
    Hospital Universitario Virgen del Rocío de Sevilla , Sevilla , Spain
  • Javier Martín
    Institute of Parasitology and Biomedicine Lopez-Neyra , CSIC, Granada , Spain
  • Jeanette Erdmann
    Institute for Cardiogenetics, University of Lübeck , Lübeck , Germany
  • Jesus M Banales
    Centro de Investigación Biomédica en Red en Enfermedades Hepáticas y Digestivas (CIBEREHD), Instituto de Salud Carlos III (ISCIII) , Madrid , Spain
  • Joan Ramon Badia
    Respiratory ICU, Institut Clínic Respiratory, Hospital Clinic, University of Barcelona, and IDIBAPS , Barcelona , Spain
  • Joaquin Dopazo
    Instituto de Biomedicina de Sevilla (IBIS) , Sevilla , Spain
  • Jochen Schneider
    Department of Internal Medicine V—Pneumology , Allergology and Intensive Care Medicine, , Homburg , Germany
  • Jonas Bergan
    Department of Research, Ostfold Hospital Trust , Gralum , Norway
  • Jordi Barretina
    Germans Trias i Pujol Research Institute (IGTP) , Badalona , Spain
  • Jörn Walter
    Department of Genetics & Epigenetics, Saarland University , Saarbrücken , Germany
  • Jose Hernández Quero
    Ibs.Granada Instituto de Investigación Biosanitaria , Granada , Spain
  • Josune Goikoetxea
    Infectious Diseases Service, Osakidetza, Biocruces Bizkaia Health Research Institute , Barakaldo , Spain
  • Juan Delgado
    Hospital Universitario Virgen del Rocío de Sevilla , Sevilla , Spain
  • Juan M Guerrero
    Hospital Universitario Virgen del Rocío de Sevilla , Sevilla , Spain
  • Julia Fazaal
    Institute of Human Genetics, University of Bonn School of Medicine & University Hospital Bonn , Bonn , Germany
  • Julia Kraft
    Charite Universitätsmedizin Berlin , Berlin , Germany
  • Julia Schröder
    Institute of Human Genetics, University of Bonn School of Medicine & University Hospital Bonn , Bonn , Germany
  • Kari Risnes
    Department of Clinical and Molecular Medicine, NTNU , Trondheim , Norway
  • Karina Banasik
    Novo Nordisk Foundation Center for Protein Research, Disease Systems Biology, Faculty of Health and Medical Sciences, University of Copenhagen , Copenhagen , Denmark
  • Karl Erik Müller
    Medical Department, Drammen Hospital, Vestre Viken Hospital Trust , Drammen , Norway
  • Karoline I Gaede
    Research Center Borstel, BioMaterialBank Nord , Borstel , Germany
  • Koldo Garcia-Etxebarria
    Centro de Investigación Biomédica en Red en Enfermedades Hepáticas y Digestivas (CIBEREHD), Instituto de Salud Carlos III (ISCIII) , Madrid , Spain
  • Kristian Tonby
    Institute of Clinical Medicine, University of Oslo , Oslo , Norway
  • Lars Heggelund
    Medical Department, Drammen Hospital, Vestre Viken Hospital Trust , Drammen , Norway
  • Laura Izquierdo-Sanchez
    Centro de Investigación Biomédica en Red en Enfermedades Hepáticas y Digestivas (CIBEREHD), Instituto de Salud Carlos III (ISCIII) , Madrid , Spain
  • Laura Rachele Bettini
    Pediatric Departement, Centro Tettamanti-European Reference Network (ERN) PaedCan, EuroBloodNet, MetabERN-University of Milano-Bicocca-Fondazione MBBM/Ospedale , San Gerardo , Italy
  • Lauro Sumoy
    Germans Trias i Pujol Research Institute (IGTP) , Badalona , Spain
  • Leif Erik Sander
    Charite Universitätsmedizin Berlin , Berlin , Germany
  • Lena J Lippert
    Charite Universitätsmedizin Berlin , Berlin , Germany
  • Leonardo Terranova
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Lindokuhle Nkambule
    Analytic and Translational Genetics Unit, Massachusetts General Hospital , Boston, MA , USA
  • Lisa Knopp
    Department of Gastroenterology , Hepatology and Infectious Diseases, , Duesseldorf , Germany
  • Lise Tuset Gustad
    Geminicenter for Sepsis Research, Institute of Circulation and Medical Imaging (ISB), NTNU , Trondheim , Norway
  • Lucia Garbarino
    HLA Laboratory, E.O. Ospedali Galliera , Genova , Italy
  • Luigi Santoro
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Luis Téllez
    Centro de Investigación Biomédica en Red en Enfermedades Hepáticas y Digestivas (CIBEREHD), Instituto de Salud Carlos III (ISCIII) , Madrid , Spain
  • Luisa Roade
    Centro de Investigación Biomédica en Red en Enfermedades Hepáticas y Digestivas (CIBEREHD), Instituto de Salud Carlos III (ISCIII) , Madrid , Spain
  • Mahnoosh Ostadreza
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Maider Intxausti
    Osakidetza Basque Health Service, Basurto University Hospital, Respiratory Service , Bilbao , Spain
  • Manolis Kogevinas
    Centro de Investigación Biomédica en Red de Epidemiología y Salud Pública (CIBERESP) , Madrid , Spain
  • Mar Riveiro-Barciela
    Centro de Investigación Biomédica en Red en Enfermedades Hepáticas y Digestivas (CIBEREHD), Instituto de Salud Carlos III (ISCIII) , Madrid , Spain
  • Marc M Berger
    Department of Anesthesiology and Intensive Care Medicine, University Hospital Essen, University Duisburg-Essen , Essen , Germany
  • Marco Schaefer
    Stefan-Morsch-Stiftung , Birkenfeld , Germany
  • Mari E K Niemi
    Institute for Molecular Medicine Finland, University of Helsinki , Helsinki , Finland
  • María A Gutiérrez-Stampa
    Osakidetza , OSI Donostialdea, Altza Primary Care, , San Sebastián , Spain
  • Maria Carrabba
    Internal Medicine Department, Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Maria E Figuera Basso
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Maria Grazia Valsecchi
    Center of Bioinformatics, Biostatistics and Bioimaging, School of Medicine and Surgery, University of Milano-Bicocca , Milan , Italy
  • María Hernandez-Tejero
    Hospital Clinic, University of Barcelona, and IDIBAPS , Barcelona , Spain
  • Maria J G T Vehreschild
    Department of Internal Medicine , Infectious Diseases, , Frankfurt am Main , Germany
  • Maria Manunta
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Marialbert Acosta-Herrera
    Institute of Parasitology and Biomedicine Lopez-Neyra , CSIC, Granada , Spain
  • Mariella D'Angiò
    Pediatric Departement, Centro Tettamanti-European Reference Network (ERN) PaedCan, EuroBloodNet, MetabERN-University of Milano-Bicocca-Fondazione MBBM/Ospedale , San Gerardo , Italy
  • Marina Baldini
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Marina Cazzaniga
    Phase 1 Research Centre, ASST Monza, School of Medicine and Surgery, University of Milano-Bicocca , Monza , Italy
  • Marit M Grimsrud
    Institute of Clinical Medicine, University of Oslo , Oslo , Norway
  • Markus Cornberg
    Department of Gastroenterology , Hepatology and Endocrinology, , Hannover , Germany
  • Markus M Nöthen
    Institute of Human Genetics, University of Bonn School of Medicine & University Hospital Bonn , Bonn , Germany
  • Marta Marquié
    Research Center and Memory Clinic, Ace Alzheimer Center Barcelona, Universitat Internacional de Catalunya , Barcelona , Spain
  • Massimo Castoldi
    Humanitas Gavazzeni-Castelli , Bergamo , Italy
  • Mattia Cordioli
    Institute for Molecular Medicine Finland, University of Helsinki , Helsinki , Finland
  • Maurizio Cecconi
    IRCCS Humanitas Research Hospital , Milan , Italy
  • Mauro D'Amato
    Ikerbasque, Basque Foundation for Science , Bilbao , Spain
  • Max Augustin
    Department I of Internal Medicine, Faculty of Medicine and University Hospital Cologne, University of Cologne , Cologne , Germany
  • Melissa Tomasi
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Mercè Boada
    Research Center and Memory Clinic, Ace Alzheimer Center Barcelona, Universitat Internacional de Catalunya , Barcelona , Spain
  • Michael Dreher
    Department of Pneumology and Intensive Care Medicine, University Hospital Aachen , Aachen , Germany
  • Michael J Seilmaier
    Munich Clinic Schwabing, Academic Teaching Hospital, Ludwig-Maximilians-University (LMU) , Munich , Germany
  • Michael Joannidis
    Division of Intensive Care and Emergency Medicine , Department of Internal Medicine, , Innsbruck , Austria
  • Michael Wittig
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Michela Mazzocco
    HLA Laboratory, E.O. Ospedali Galliera , Genova , Italy
  • Michele Ciccarelli
    Humanitas Clinical and Research Center, IRCCS , Milan , Italy
  • Miguel Rodríguez-Gandía
    Centro de Investigación Biomédica en Red en Enfermedades Hepáticas y Digestivas (CIBEREHD), Instituto de Salud Carlos III (ISCIII) , Madrid , Spain
  • Monica Bocciolone
    Humanitas Clinical and Research Center, IRCCS , Milan , Italy
  • Monica Miozzo
    University of Milan , Milan , Italy
  • Natale Imaz Ayo
    Biocruces Bizkaia Health Research Institute , Barakaldo , Spain
  • Natalia Blay
    Genomes for Life-GCAT Lab, Germans Trias i Pujol Research Institute (IGTP) , Badalona , Spain
  • Natalia Chueca
    Microbiology Unit. Hospital Universitario Clinico San Cecilio , Granada , Spain
  • Nicola Montano
    University of Milan , Milan , Italy
  • Nicole Braun
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Nicole Ludwig
    Department of Human Genetics, Center of Human and Molecular Biology, University Hospital Saarland , Homburg/Saar , Germany
  • Nikolaus Marx
    Department of Internal Medicine I, RWTH Aachen University Hospital , Aachen , Germany
  • Nilda Martínez
    Department of Anesthesiology, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS) , Madrid , Spain
  • Oliver A Cornely
    Cologne Excellence Cluster on Cellular Stress Responses in Aging-Associated Diseases (CECAD), Faculty of Medicine and University Hospital Cologne, University of Cologne , Cologne , Germany
  • Oliver Witzke
    Department of Infectious Diseases, University Hospital Essen, University Duisburg-Essen , Essen , Germany
  • Orazio Palmieri
    Gastroenterology Unit, Fondazione IRCCS Casa Sollievo della Sofferenza , San Giovanni Rotondo , Italy
  • Paola Faverio
    School of Medicine and Surgery, University of Milano-Bicocca , Milan , Italy
  • Paoletta Preatoni
    Humanitas Clinical and Research Center, IRCCS , Milan , Italy
  • Paolo Bonfanti
    School of Medicine and Surgery, University of Milano-Bicocca , Milan , Italy
  • Paolo Omodei
    Humanitas Clinical and Research Center, IRCCS , Milan , Italy
  • Paolo Tentorio
    IRCCS Humanitas Research Hospital , Milan , Italy
  • Pedro Castro
    Hospital Clinic, University of Barcelona, and IDIBAPS , Barcelona , Spain
  • Pedro M Rodrigues
    Centro de Investigación Biomédica en Red en Enfermedades Hepáticas y Digestivas (CIBEREHD), Instituto de Salud Carlos III (ISCIII) , Madrid , Spain
  • Pedro Pablo España
    Osakidetza Basque Health Service, Galdakao Hospital, Respiratory Service , Galdakao , Spain
  • Per Hoffmann
    Institute of Human Genetics, University of Bonn School of Medicine & University Hospital Bonn , Bonn , Germany
  • Philip Rosenstiel
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Philipp Schommers
    Department I of Internal Medicine, Faculty of Medicine and University Hospital Cologne, University of Cologne , Cologne , Germany
  • Phillip Suwalski
    Charite Universitätsmedizin Berlin , Berlin , Germany
  • Raúl de Pablo
    Department of Intensive Care, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of Alcalá , Madrid , Spain
  • Ricard Ferrer
    Intensive Care Department, Vall d'Hebron University Hospital, SODIR-VHIR research group , Barcelona , Spain
  • Robert Bals
    Department of Internal Medicine V—Pneumology , Allergology and Intensive Care Medicine, , Homburg , Germany
  • Roberta Gualtierotti
    University of Milan , Milan , Italy
  • Rocío Gallego-Durán
    Hospital Universitario Virgen del Rocío de Sevilla , Sevilla , Spain
  • Rosa Nieto
    Department of Respiratory Diseases, Hospital Universitario Ramón y Cajal, Instituto Ramón y Cajal de Investigación Sanitaria (IRYCIS), University of Alcalá , Madrid , Spain
  • Rossana Carpani
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Rubén Morilla
    Hospital Universitario Virgen del Rocío de Sevilla , Sevilla , Spain
  • Salvatore Badalamenti
    IRCCS Humanitas Research Hospital , Milan , Italy
  • Sammra Haider
    Department of Medicine, Møre & Romsdal Hospital Trust , Molde , Norway
  • Sandra Ciesek
    Institute of Medical Virology, University Hospital Frankfurt, Goethe University , Frankfurt am Main , Germany
  • Sandra May
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Sara Bombace
    IRCCS Humanitas Research Hospital , Milan , Italy
  • Sara Marsal
    Vall d’Hebron Institut de Recerca (VHIR), Vall d’Hebron Hospital Universitari , Barcelona , Spain
  • Sara Pigazzini
    Institute for Molecular Medicine Finland, University of Helsinki , Helsinki , Finland
  • Sebastian Klein
    Division of Intensive Care and Emergency Medicine , Department of Internal Medicine, , Innsbruck , Austria
  • Serena Pelusi
    University of Milan , Milan , Italy
  • Sibylle Wilfling
    Department for Infectious Diseases and Infection Control, University Hospital Regensburg , Regensburg , Germany
  • Silvano Bosari
    University of Milan , Milan , Italy
  • Sonja Volland
    Hannover Unified Biobank, Hannover Medical School , Hannover , Germany
  • Søren Brunak
    Novo Nordisk Foundation Center for Protein Research, Disease Systems Biology, Faculty of Health and Medical Sciences, University of Copenhagen , Copenhagen , Denmark
  • Soumya Raychaudhuri
    Program in Medical and Population Genetics, Broad Institute of MIT and Harvard , Cambridge, MA , USA
  • Stefan Schreiber
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Stefanie Heilmann-Heimbach
    Institute of Human Genetics, University of Bonn School of Medicine & University Hospital Bonn , Bonn , Germany
  • Stefano Aliberti
    IRCCS Humanitas Research Hospital , Milan , Italy
  • Stephan Ripke
    Charite Universitätsmedizin Berlin , Berlin , Germany
  • Susanne Dudman
    Institute of Clinical Medicine, University of Oslo , Oslo , Norway
  • Tanja Wesse
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Tenghao Zheng
    School of Biological Sciences, Monash University , Clayton, VIC , Australia
  • Thomas Bahmer
    Klinik für Innere Medizin I, Universitätsklinikum Schleswig-Holstein , Kiel , Germany
  • Thomas Eggermann
    Institute of Human Genetics, Medical Faculty, RWTH Aachen University , Aachen , Germany
  • Thomas Illig
    Hannover Unified Biobank, Hannover Medical School , Hannover , Germany
  • Thorsten Brenner
    Department of Anesthesiology and Intensive Care Medicine, University Hospital Essen, University Duisburg-Essen , Essen , Germany
  • Tomas Pumarola
    Department of Microbiology, University Hospital Vall d’Hebron , Barcelona , Spain
  • Torsten Feldt
    Department of Gastroenterology , Hepatology and Infectious Diseases, , Duesseldorf , Germany
  • Trine Folseraas
    Institute of Clinical Medicine, University of Oslo , Oslo , Norway
  • Trinidad Gonzalez Cejudo
    Biochemistry Unit, Hospital Universitario Clinico San Cecilio , Granada , Spain
  • Ulf Landmesser
    Berlin Institute of Health, Charite Universitätsmedizin Berlin , Berlin Germany
  • Ulrike Protzer
    Institute of Virology, Technical University Munich/Helmholtz Zentrum München , Munich , Germany
  • Ute Hehr
    Zentrum für Humangenetik Regensburg , Regensburg , Germany
  • Valeria Rimoldi
    Department of Biomedical Sciences, Humanitas University , Milan , Italy
  • Valter Monzani
    Fondazione IRCCS Ca' Granda Ospedale Maggiore Policlinico , Milan , Italy
  • Vegard Skogen
    Department of Infectious Diseases, University Hospital of North Norway , Tromsø , Norway
  • Verena Keitel
    Department of Gastroenterology , Hepatology and Infectious Diseases, , Duesseldorf , Germany
  • Verena Kopfnagel
    Hannover Unified Biobank, Hannover Medical School , Hannover , Germany
  • Vicente Friaza
    Hospital Universitario Virgen del Rocío de Sevilla , Sevilla , Spain
  • Victor Andrade
    Division of Neurogenetics and Molecular Psychiatry , Department of Psychiatry and Psychotherapy, , Cologne , Germany
  • Victor Moreno
    Centro de Investigación Biomédica en Red de Epidemiología y Salud Pública (CIBERESP) , Madrid , Spain
  • Wolfgang Albrecht
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Wolfgang Peter
    Stefan-Morsch-Stiftung , Birkenfeld , Germany
  • Wolfgang Poller
    Charite Universitätsmedizin Berlin , Berlin , Germany
  • Xavier Farre
    Genomes for Life-GCAT Lab, Germans Trias i Pujol Research Institute (IGTP) , Badalona , Spain
  • Xiaoli Yi
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Xiaomin Wang
    Charite Universitätsmedizin Berlin , Berlin , Germany
  • Yascha Khodamoradi
    Department of Internal Medicine , Infectious Diseases, , Frankfurt am Main , Germany
  • Zehra Karadeniz
    Charite Universitätsmedizin Berlin , Berlin , Germany
  • Anna Latiano
    Gastroenterology Unit, Fondazione IRCCS Casa Sollievo della Sofferenza , San Giovanni Rotondo , Italy
  • Siegfried Goerg
    Institute of Transfusionsmedicine, University Hospital Schleswig-Holstein (UKSH) , Kiel , Germany
  • Petra Bacher
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Philipp Koehler
    Cologne Excellence Cluster on Cellular Stress Responses in Aging-Associated Diseases (CECAD), Faculty of Medicine and University Hospital Cologne, University of Cologne , Cologne , Germany
  • Florian Tran
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany
  • Heinz Zoller
    Department of Medicine I, Gastroenterology, Hepatology and Endocrinology, Medical University of Innsbruck , Innsbruck , Austria
  • Eva C Schulte
    Institute of Virology, Technical University Munich/Helmholtz Zentrum München , Munich , Germany
  • Bettina Heidecker
    Charite Universitätsmedizin Berlin , Berlin , Germany
  • Kerstin U Ludwig
    Institute of Human Genetics, University of Bonn School of Medicine & University Hospital Bonn , Bonn , Germany
  • Javier Fernández
    Hospital Clinic, University of Barcelona, and IDIBAPS , Barcelona , Spain
  • Manuel Romero-Gómez
    Hospital Universitario Virgen del Rocío de Sevilla , Sevilla , Spain
  • Agustín Albillos
    Centro de Investigación Biomédica en Red en Enfermedades Hepáticas y Digestivas (CIBEREHD), Instituto de Salud Carlos III (ISCIII) , Madrid , Spain
  • Pietro Invernizzi
    European Reference Network on Hepatological Diseases (ERN RARE-LIVER), San Gerardo Hospital , Monza , Italy
  • Maria Buti
    Centro de Investigación Biomédica en Red en Enfermedades Hepáticas y Digestivas (CIBEREHD), Instituto de Salud Carlos III (ISCIII) , Madrid , Spain
  • Stefano Duga
    IRCCS Humanitas Research Hospital , Milan , Italy
  • Luis Bujanda
    Centro de Investigación Biomédica en Red en Enfermedades Hepáticas y Digestivas (CIBEREHD), Instituto de Salud Carlos III (ISCIII) , Madrid , Spain
  • Johannes R Hov
    Institute of Clinical Medicine, University of Oslo , Oslo , Norway
  • Tobias L Lenz
    Research Group for Evolutionary Immunogenomics, Max Planck Institute for Evolutionary Biology , Plön , Germany
  • Rosanna Asselta
    IRCCS Humanitas Research Hospital , Milan , Italy
  • Rafael de Cid
    Genomes for Life-GCAT Lab, Germans Trias i Pujol Research Institute (IGTP) , Badalona , Spain
  • Luca Valenti
    University of Milan , Milan , Italy
  • Tom H Karlsen
    Institute of Clinical Medicine, University of Oslo , Oslo , Norway
  • Mario Cáceres
    Institut de Biotecnologia i de Biomedicina, Universitat Autònoma de Barcelona , Barcelona , Spain
  • Andre Franke
    Institute of Clinical Molecular Biology, Christian-Albrechts-University , Kiel , Germany

Abstract

<jats:title>Abstract</jats:title> <jats:p>Given the highly variable clinical phenotype of Coronavirus disease 2019 (COVID-19), a deeper analysis of the host genetic contribution to severe COVID-19 is important to improve our understanding of underlying disease mechanisms. Here, we describe an extended genome-wide association meta-analysis of a well-characterized cohort of 3255 COVID-19 patients with respiratory failure and 12 488 population controls from Italy, Spain, Norway and Germany/Austria, including stratified analyses based on age, sex and disease severity, as well as targeted analyses of chromosome Y haplotypes, the human leukocyte antigen region and the SARS-CoV-2 peptidome. By inversion imputation, we traced a reported association at 17q21.31 to a ~0.9-Mb inversion polymorphism that creates two highly differentiated haplotypes and characterized the potential effects of the inversion in detail. Our data, together with the 5th release of summary statistics from the COVID-19 Host Genetics Initiative including non-Caucasian individuals, also identified a new locus at 19q13.33, including NAPSA, a gene which is expressed primarily in alveolar cells responsible for gas exchange in the lung.</jats:p>

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