X-linked recessive TLR7 deficiency in ~1% of men under 60 years old with life-threatening COVID-19

COVID Human Genetic Effort; Amsterdam UMC COVID-19 Biobank

doi:10.1126/sciimmunol.abl4348

X-linked recessive TLR7 deficiency in ~1% of men under 60 years old with life-threatening COVID-19

COVID Human Genetic Effort
, Amsterdam UMC COVID-19 Biobank

JEM The Rockefeller University Press
University Sordonne-Paris-Cite; Paris-Descartes Medical School, Paris, France
Necker Hospital for Sick Children
Helix, San Mateo, CA, USA
the Howard Hughes Medical Institute, Yale University School of Medicine, New Haven, Connecticut 06519, and Departments of Internal Medicine and Cellular and Molecular Physiology, Yale University School of Medicine, New Haven, Connecticut 06520.
Laboratory of Clinical Immunology, Department of Microbiology and Immunology, KU Leuven, Leuven, Belgium.
Vall d'Hebron University Hospital (HUVH)
Jeffrey Modell Diagnostic and Research Center for Primary Immunodeficiencies
Avicenne Hospital
Specialized Immunology Laboratory of Dr. Shahrooei, Ahvaz, Iran
Shahid Beheshti University of Medical Sciences
Department of Pharmacy, Faculty of Health Sciences, UiT The Arctic University of Norway, Norway; Department of Health and Caring Sciences, Faculty of Health and Life Sciences, Linnaeus University, Sweden
Bilkent University
University of Rome Tor Vergata
Bambino Gesù Children Hospital
Vita-Salute San Raffaele University
IRCCS Ospedale San Raffaele
Molecular Hematology Unit
University of Antioquia UDEA
Universidad de la Sabana
Laboratory of Immunology, The Rockefeller University, New York, New York 10021, USA.
NSERM Unit 1000 ″Neuroimaging and Psychiatry″, University Paris Sud, University Paris Descartes, Paris, France.
Helix
Howard Hughes Medical Institute, Yale University, New Haven, CT 06519, USA; Department of Medicine, Yale University School of Medicine, New Haven, CT 06520, USA; Department of Cellular and Molecular Physiology, Yale University School of Medicine, New Haven, CT 06520, USA.
Laboratory of Clinical Immunology and Microbiology, Bethesda, United States
Specialized Immunology Laboratory of Dr. Shahrooei
University of Health Sciences
Tor Vergata University of Rome, Rome, Italy
University of Antioquia
University Paris-Sud, Université Paris-Saclay, Le Kremlin-Bicêtre, Paris, France.
Infanta Leonor University Hospital
Bellvitge Biomedical Biomedical Research Institute (IDIBELL), 199 08908 Barcelona, Spain
The Barcelona Institute of Science and Technology (BIST)
Complutense University
University Hospital of Gran Canaria Dr. Negrín
Institute of Technology and Renewable Energies (ITER)
Universidad Politecnica de Madrid, 28040 Madrid, Spain
University Hospital La Paz
Necmettin Erbakan University
Konya City Hospital
Karolinska University Hospital and Karolinska Institute, Stockholm, Sweden
Central Hospital-Anesthesia and Intensive Care Unit
Unit of Clinical Therapy Research, Inflammatory Diseases (ClinTRID), Department of Medicine, Karolinska Institutet, Karolinska University Hospital, Stockholm, Sweden; Rheumatology Unit, Department of Medicine, Karolinska University Hospital and Karolinska Institutet, Stockholm, Sweden.
Department of Rheumatology & Clinical Immunology, Laboratory of Translational Immunology, Department of Immunology, University Medical Center Utrecht, Utrecht, The Netherlands.
Physiology & Genetics Uniformed Services University of the Health Sciences
San Gerardo Hospital-University of Milano-Bicocca
MetabERN-University of Milano-Bicocca-Fondazione MBBM- Ospedale San Gerardo
CHRU de Nancy, F-CRIN INI-CRCT (Cardiovascular and Renal Clinical Trialists), INSERM U1116, Centre d'Investigation Clinique et Plurithématique 1433, INSERM, Université de Lorraine, Nancy, France.
Invitae
Uniformed Services University of the Health Sciences

Research output: Contribution to journal › Article › Academic › peer-review

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Abstract

Autosomal inborn errors of type I IFN immunity and autoantibodies against these cytokines underlie at least 10% of critical COVID-19 pneumonia cases. We report very rare, biochemically deleterious X-linked TLR7 variants in 16 unrelated male individuals aged 7 to 71 years (mean, 36.7 years) from a cohort of 1202 male patients aged 0.5 to 99 years (mean, 52.9 years) with unexplained critical COVID-19 pneumonia. None of the 331 asymptomatically or mildly infected male individuals aged 1.3 to 102 years (mean, 38.7 years) tested carry such TLR7 variants (P = 3.5 × 10 ⁻⁵). The phenotypes of five hemizygous relatives of index cases infected with SARS-CoV-2 include asymptomatic or mild infection (n = 2) or moderate (n = 1), severe (n = 1), or critical (n = 1) pneumonia. Two patients from a cohort of 262 male patients with severe COVID-19 pneumonia (mean, 51.0 years) are hemizygous for a deleterious TLR7 variant. The cumulative allele frequency for deleterious TLR7 variants in the male general population is <6.5 × 10 ⁻⁴. We show that blood B cell lines and myeloid cell subsets from the patients do not respond to TLR7 stimulation, a phenotype rescued by wild-type TLR7. The patients’ blood plasmacytoid dendritic cells (pDCs) produce low levels of type I IFNs in response to SARS-CoV-2. Overall, X-linked recessive TLR7 deficiency is a highly penetrant genetic etiology of critical COVID-19 pneumonia, in about 1.8% of male patients below the age of 60 years. Human TLR7 and pDCs are essential for protective type I IFN immunity against SARS-CoV-2 in the respiratory tract.

Original language	English
Article number	eabl4348
Journal	Science immunology
Volume	6
Issue number	62
DOIs	https://doi.org/10.1126/sciimmunol.abl4348
Publication status	Published - 19 Aug 2021

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@article{049c1fcc139b442192a7a13cb0e52c0f,

title = "X-linked recessive TLR7 deficiency in \textasciitilde{}1\% of men under 60 years old with life-threatening COVID-19",

abstract = "Autosomal inborn errors of type I IFN immunity and autoantibodies against these cytokines underlie at least 10\% of critical COVID-19 pneumonia cases. We report very rare, biochemically deleterious X-linked TLR7 variants in 16 unrelated male individuals aged 7 to 71 years (mean, 36.7 years) from a cohort of 1202 male patients aged 0.5 to 99 years (mean, 52.9 years) with unexplained critical COVID-19 pneumonia. None of the 331 asymptomatically or mildly infected male individuals aged 1.3 to 102 years (mean, 38.7 years) tested carry such TLR7 variants (P = 3.5 {\texttimes} 10 −5). The phenotypes of five hemizygous relatives of index cases infected with SARS-CoV-2 include asymptomatic or mild infection (n = 2) or moderate (n = 1), severe (n = 1), or critical (n = 1) pneumonia. Two patients from a cohort of 262 male patients with severe COVID-19 pneumonia (mean, 51.0 years) are hemizygous for a deleterious TLR7 variant. The cumulative allele frequency for deleterious TLR7 variants in the male general population is <6.5 {\texttimes} 10 −4. We show that blood B cell lines and myeloid cell subsets from the patients do not respond to TLR7 stimulation, a phenotype rescued by wild-type TLR7. The patients{\textquoteright} blood plasmacytoid dendritic cells (pDCs) produce low levels of type I IFNs in response to SARS-CoV-2. Overall, X-linked recessive TLR7 deficiency is a highly penetrant genetic etiology of critical COVID-19 pneumonia, in about 1.8\% of male patients below the age of 60 years. Human TLR7 and pDCs are essential for protective type I IFN immunity against SARS-CoV-2 in the respiratory tract. ",

author = "\{COVID Human Genetic Effort\} and Takaki Asano and Bertrand Boisson and Fanny Onodi and Daniela Matuozzo and Marcela Moncada-Velez and \{Maglorius Renkilaraj\}, \{Majistor Raj Luxman\} and Peng Zhang and Laurent Meertens and Alexandre Bolze and Marie Materna and Sarantis Korniotis and Adrian Gervais and Estelle Talouarn and Benedetta Bigio and Yoann Seeleuthner and Kaya Bilguvar and Yu Zhang and Anna-Lena Neehus and Masato Ogishi and Pelham, \{Simon J\} and \{Le Voyer\}, Tom and J{\'e}r{\'e}mie Rosain and Quentin Philippot and Pere Soler-Palac{\'i}n and Roger Colobran and Andrea Martin-Nalda and Rivi{\`e}re, \{Jacques G\} and Yacine Tandjaoui-Lambiotte and Khalil Cha{\"i}bi and Mohammad Shahrooei and Darazam, \{Ilad Alavi\} and Olyaei, \{Nasrin Alipour\} and Davood Mansouri and Nevin Hatipoğlu and Figen Palabiyik and Tayfun Ozcelik and Giuseppe Novelli and Antonio Novelli and Giorgio Casari and Alessandro Aiuti and Paola Carrera and Simone Bondesan and Federica Barzaghi and Patrizia Rovere-Querini and Cristina Tresoldi and Franco, \{Jose Luis\} and Julian Rojas and Reyes, \{Luis Felipe\} and Bustos, \{Ingrid G\} and Arias, \{Andres Augusto\} and \{Amsterdam UMC COVID-19 Biobank\} and \{van de Beek\}, Diederik and Wiersinga, \{W. Joost\} and \{de Bree\}, \{Godelieve J.\} and Geerlings, \{Suzanne E.\} and Bram Goorhuis and Grobusch, \{Martin P.\} and Harris, \{Vanessa C.\} and Hermans, \{Sabine M.\} and Hovius, \{Joppe W. R.\} and Jeannine Nellen and \{van der Poll\}, Tom and Prins, \{Jan M.\} and Stijnis, \{Cornelis S.\} and \{van der Valk\}, Marc and \{van Vugt\}, Michele and Hollmann, \{Markus W.\} and Preckel, \{Prof. dr. , Benedikt\} and Veelo, \{Denise P.\} and Guillaume Morelle and Kyheng Christ{\`e}le and Jes{\'u}s Troya and Laura Planas-Serra and Agatha Schl{\"u}ter and Marta Gut and Aurora Pujol and Allende, \{Luis M\} and Carlos Rodriguez-Gallego and Carlos Flores and Oscar Cabrera-Marante and Pleguezuelo, \{Daniel E\} and \{de Diego\}, \{Rebeca P{\'e}rez\} and Sevgi Keles and Gokhan Aytekin and Akcan, \{Ozge Metin\} and Bryceson, \{Yenan T\} and Peter Bergman and Petter Brodin and Daniel Smole and Smith, \{C I Edvard\} and Anna-Carin Norlin and Campbell, \{Tessa M\} and Covill, \{Laura E\} and Lennart Hammarstr{\"o}m and Qiang Pan-Hammarstr{\"o}m and Hassan Abolhassani and Shrikant Mane and Nico Marr and Manar Ata and \{Al Ali\}, Fatima and Taushif Khan and Spaan, \{Andr{\'a}s N\} and Dalgard, \{Clifton L\} and Paolo Bonfanti and Andrea Biondi and Sarah Tubiana and Charles Burdet and Robert Nussbaum and Amanda Kahn-Kirby and Snow, \{Andrew L\} and Jacinta Bustamante and Anne Puel and St{\'e}phanie Boisson-Dupuis and Shen-Ying Zhang and Vivien B{\'e}ziat and Lifton, \{Richard P\} and Paul Bastard and Notarangelo, \{Luigi D\} and Laurent Abel and Su, \{Helen C\} and Emmanuelle Jouanguy and Ali Amara and Vassili Soumelis and Aur{\'e}lie Cobat and Qian Zhang and Jean-Laurent Casanova and M Bugiani",

note = "Members of Amsterdam UMC Covid-19 Biobank: Michiel van Agtmael2, Anne Geke Algera1, Brent Appelman2, Frank van Baarle1, Diane Bax3, Martijn Beudel4, Harm Jan Bogaard5, Marije Bomers2, Peter Bonta5, Lieuwe Bos1, Michela Botta1, Justin de Brabander2, Godelieve de Bree2, Sanne de Bruin1, David T. P. Buis1, Marianna Bugiani5, Esther Bulle1, Osoul Chouchane2 Alex Cloherty3, Mirjam Dijkstra12, Dave A. Dongelmans1, Romein W. G. Dujardin1, Paul Elbers1, Lucas Fleuren1, Suzanne Geerlings2 Theo Geijtenbeek3, Armand Girbes1, Bram Goorhuis2, Martin P. Grobusch2, Florianne Hafkamp3, Laura Hagens1, Jorg Hamann7, Vanessa Harris2, Robert Hemke8, Sabine M. Hermans2 Leo Heunks1, Markus Hollmann6, Janneke Horn1, Joppe W. Hovius2, Menno D. de Jong9, Rutger Koning4, Endry H. T. Lim1, Niels van Mourik1, Jeaninne Nellen2, Esther J. Nossent5, Frederique Paulus1, Edgar Peters2, Dan A. I. Pina-Fuentes4, Tom van der Poll2, Bennedikt Preckel6, Jan M. Prins2, Jorinde Raasveld1, Tom Reijnders2, Maurits C. F. J. de Rotte12, Michiel Schinkel2, Marcus J. Schultz1, Femke A. P. Schrauwen12, Alex Schuurmans10, Jaap Schuurmans1, Kim Sigaloff1, Marleen A. Slim1,2, Patrick Smeele5, Marry Smit1, Cornelis S. Stijnis2, Willemke Stilma1, Charlotte Teunissen11, Patrick Thoral1, Anissa M. Tsonas1, Pieter R. Tuinman2, Marc van der Valk2, Denise Veelo6, Carolien Volleman1, Heder de Vries1, Lonneke A. Vught1,2, Mich{\`e}le van Vugt2, Dorien Wouters12, A. H. (Koos) Zwinderman13, Matthijs C. Brouwer4, W. Joost Wiersinga2, Alexander P. J. Vlaar1, Diederik van de Beek ([email protected])4",

year = "2021",

month = aug,

day = "19",

doi = "10.1126/sciimmunol.abl4348",

language = "English",

volume = "6",

journal = "Science immunology",

issn = "2470-9468",

publisher = "American Association for the Advancement of Science",

number = "62",

}

TY - JOUR

T1 - X-linked recessive TLR7 deficiency in ~1% of men under 60 years old with life-threatening COVID-19

AU - COVID Human Genetic Effort

AU - Asano, Takaki

AU - Boisson, Bertrand

AU - Onodi, Fanny

AU - Matuozzo, Daniela

AU - Moncada-Velez, Marcela

AU - Maglorius Renkilaraj, Majistor Raj Luxman

AU - Zhang, Peng

AU - Meertens, Laurent

AU - Bolze, Alexandre

AU - Materna, Marie

AU - Korniotis, Sarantis

AU - Gervais, Adrian

AU - Talouarn, Estelle

AU - Bigio, Benedetta

AU - Seeleuthner, Yoann

AU - Bilguvar, Kaya

AU - Zhang, Yu

AU - Neehus, Anna-Lena

AU - Ogishi, Masato

AU - Pelham, Simon J

AU - Le Voyer, Tom

AU - Rosain, Jérémie

AU - Philippot, Quentin

AU - Soler-Palacín, Pere

AU - Colobran, Roger

AU - Martin-Nalda, Andrea

AU - Rivière, Jacques G

AU - Tandjaoui-Lambiotte, Yacine

AU - Chaïbi, Khalil

AU - Shahrooei, Mohammad

AU - Darazam, Ilad Alavi

AU - Olyaei, Nasrin Alipour

AU - Mansouri, Davood

AU - Hatipoğlu, Nevin

AU - Palabiyik, Figen

AU - Ozcelik, Tayfun

AU - Novelli, Giuseppe

AU - Novelli, Antonio

AU - Casari, Giorgio

AU - Aiuti, Alessandro

AU - Carrera, Paola

AU - Bondesan, Simone

AU - Barzaghi, Federica

AU - Rovere-Querini, Patrizia

AU - Tresoldi, Cristina

AU - Franco, Jose Luis

AU - Rojas, Julian

AU - Reyes, Luis Felipe

AU - Bustos, Ingrid G

AU - Arias, Andres Augusto

AU - Amsterdam UMC COVID-19 Biobank

AU - van de Beek, Diederik

AU - Wiersinga, W. Joost

AU - de Bree, Godelieve J.

AU - Geerlings, Suzanne E.

AU - Goorhuis, Bram

AU - Grobusch, Martin P.

AU - Harris, Vanessa C.

AU - Hermans, Sabine M.

AU - Hovius, Joppe W. R.

AU - Nellen, Jeannine

AU - van der Poll, Tom

AU - Prins, Jan M.

AU - Stijnis, Cornelis S.

AU - van der Valk, Marc

AU - van Vugt, Michele

AU - Hollmann, Markus W.

AU - Preckel, Prof. dr. , Benedikt

AU - Veelo, Denise P.

AU - Morelle, Guillaume

AU - Christèle, Kyheng

AU - Troya, Jesús

AU - Planas-Serra, Laura

AU - Schlüter, Agatha

AU - Gut, Marta

AU - Pujol, Aurora

AU - Allende, Luis M

AU - Rodriguez-Gallego, Carlos

AU - Flores, Carlos

AU - Cabrera-Marante, Oscar

AU - Pleguezuelo, Daniel E

AU - de Diego, Rebeca Pérez

AU - Keles, Sevgi

AU - Aytekin, Gokhan

AU - Akcan, Ozge Metin

AU - Bryceson, Yenan T

AU - Bergman, Peter

AU - Brodin, Petter

AU - Smole, Daniel

AU - Smith, C I Edvard

AU - Norlin, Anna-Carin

AU - Campbell, Tessa M

AU - Covill, Laura E

AU - Hammarström, Lennart

AU - Pan-Hammarström, Qiang

AU - Abolhassani, Hassan

AU - Mane, Shrikant

AU - Marr, Nico

AU - Ata, Manar

AU - Al Ali, Fatima

AU - Khan, Taushif

AU - Spaan, András N

AU - Dalgard, Clifton L

AU - Bonfanti, Paolo

AU - Biondi, Andrea

AU - Tubiana, Sarah

AU - Burdet, Charles

AU - Nussbaum, Robert

AU - Kahn-Kirby, Amanda

AU - Snow, Andrew L

AU - Bustamante, Jacinta

AU - Puel, Anne

AU - Boisson-Dupuis, Stéphanie

AU - Zhang, Shen-Ying

AU - Béziat, Vivien

AU - Lifton, Richard P

AU - Bastard, Paul

AU - Notarangelo, Luigi D

AU - Abel, Laurent

AU - Su, Helen C

AU - Jouanguy, Emmanuelle

AU - Amara, Ali

AU - Soumelis, Vassili

AU - Cobat, Aurélie

AU - Zhang, Qian

AU - Casanova, Jean-Laurent

AU - Bugiani, M

N1 - Members of Amsterdam UMC Covid-19 Biobank: Michiel van Agtmael2, Anne Geke Algera1, Brent Appelman2, Frank van Baarle1, Diane Bax3, Martijn Beudel4, Harm Jan Bogaard5, Marije Bomers2, Peter Bonta5, Lieuwe Bos1, Michela Botta1, Justin de Brabander2, Godelieve de Bree2, Sanne de Bruin1, David T. P. Buis1, Marianna Bugiani5, Esther Bulle1, Osoul Chouchane2 Alex Cloherty3, Mirjam Dijkstra12, Dave A. Dongelmans1, Romein W. G. Dujardin1, Paul Elbers1, Lucas Fleuren1, Suzanne Geerlings2 Theo Geijtenbeek3, Armand Girbes1, Bram Goorhuis2, Martin P. Grobusch2, Florianne Hafkamp3, Laura Hagens1, Jorg Hamann7, Vanessa Harris2, Robert Hemke8, Sabine M. Hermans2 Leo Heunks1, Markus Hollmann6, Janneke Horn1, Joppe W. Hovius2, Menno D. de Jong9, Rutger Koning4, Endry H. T. Lim1, Niels van Mourik1, Jeaninne Nellen2, Esther J. Nossent5, Frederique Paulus1, Edgar Peters2, Dan A. I. Pina-Fuentes4, Tom van der Poll2, Bennedikt Preckel6, Jan M. Prins2, Jorinde Raasveld1, Tom Reijnders2, Maurits C. F. J. de Rotte12, Michiel Schinkel2, Marcus J. Schultz1, Femke A. P. Schrauwen12, Alex Schuurmans10, Jaap Schuurmans1, Kim Sigaloff1, Marleen A. Slim1,2, Patrick Smeele5, Marry Smit1, Cornelis S. Stijnis2, Willemke Stilma1, Charlotte Teunissen11, Patrick Thoral1, Anissa M. Tsonas1, Pieter R. Tuinman2, Marc van der Valk2, Denise Veelo6, Carolien Volleman1, Heder de Vries1, Lonneke A. Vught1,2, Michèle van Vugt2, Dorien Wouters12, A. H. (Koos) Zwinderman13, Matthijs C. Brouwer4, W. Joost Wiersinga2, Alexander P. J. Vlaar1, Diederik van de Beek ([email protected])4

PY - 2021/8/19

Y1 - 2021/8/19

N2 - Autosomal inborn errors of type I IFN immunity and autoantibodies against these cytokines underlie at least 10% of critical COVID-19 pneumonia cases. We report very rare, biochemically deleterious X-linked TLR7 variants in 16 unrelated male individuals aged 7 to 71 years (mean, 36.7 years) from a cohort of 1202 male patients aged 0.5 to 99 years (mean, 52.9 years) with unexplained critical COVID-19 pneumonia. None of the 331 asymptomatically or mildly infected male individuals aged 1.3 to 102 years (mean, 38.7 years) tested carry such TLR7 variants (P = 3.5 × 10 −5). The phenotypes of five hemizygous relatives of index cases infected with SARS-CoV-2 include asymptomatic or mild infection (n = 2) or moderate (n = 1), severe (n = 1), or critical (n = 1) pneumonia. Two patients from a cohort of 262 male patients with severe COVID-19 pneumonia (mean, 51.0 years) are hemizygous for a deleterious TLR7 variant. The cumulative allele frequency for deleterious TLR7 variants in the male general population is <6.5 × 10 −4. We show that blood B cell lines and myeloid cell subsets from the patients do not respond to TLR7 stimulation, a phenotype rescued by wild-type TLR7. The patients’ blood plasmacytoid dendritic cells (pDCs) produce low levels of type I IFNs in response to SARS-CoV-2. Overall, X-linked recessive TLR7 deficiency is a highly penetrant genetic etiology of critical COVID-19 pneumonia, in about 1.8% of male patients below the age of 60 years. Human TLR7 and pDCs are essential for protective type I IFN immunity against SARS-CoV-2 in the respiratory tract.

AB - Autosomal inborn errors of type I IFN immunity and autoantibodies against these cytokines underlie at least 10% of critical COVID-19 pneumonia cases. We report very rare, biochemically deleterious X-linked TLR7 variants in 16 unrelated male individuals aged 7 to 71 years (mean, 36.7 years) from a cohort of 1202 male patients aged 0.5 to 99 years (mean, 52.9 years) with unexplained critical COVID-19 pneumonia. None of the 331 asymptomatically or mildly infected male individuals aged 1.3 to 102 years (mean, 38.7 years) tested carry such TLR7 variants (P = 3.5 × 10 −5). The phenotypes of five hemizygous relatives of index cases infected with SARS-CoV-2 include asymptomatic or mild infection (n = 2) or moderate (n = 1), severe (n = 1), or critical (n = 1) pneumonia. Two patients from a cohort of 262 male patients with severe COVID-19 pneumonia (mean, 51.0 years) are hemizygous for a deleterious TLR7 variant. The cumulative allele frequency for deleterious TLR7 variants in the male general population is <6.5 × 10 −4. We show that blood B cell lines and myeloid cell subsets from the patients do not respond to TLR7 stimulation, a phenotype rescued by wild-type TLR7. The patients’ blood plasmacytoid dendritic cells (pDCs) produce low levels of type I IFNs in response to SARS-CoV-2. Overall, X-linked recessive TLR7 deficiency is a highly penetrant genetic etiology of critical COVID-19 pneumonia, in about 1.8% of male patients below the age of 60 years. Human TLR7 and pDCs are essential for protective type I IFN immunity against SARS-CoV-2 in the respiratory tract.

UR - https://www.scopus.com/pages/publications/85113561257

U2 - 10.1126/sciimmunol.abl4348

DO - 10.1126/sciimmunol.abl4348

M3 - Article

SN - 2470-9468

VL - 6

JO - Science immunology

JF - Science immunology

IS - 62

M1 - eabl4348

ER -

X-linked recessive TLR7 deficiency in ~1% of men under 60 years old with life-threatening COVID-19

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