Cytosolic proliferating cell nuclear antigen (PCNA) orchestrates neutrophil hyperactivation in COVID-19

R Rodrigo de Oliveira Formiga (Université Paris Cité, CNRS UMR 8104, Inserm U1016) L Lucie Pesenti (Université Paris Cité, CNRS UMR 8104, Inserm U1016) F François Chable de la Héronnière (Université Paris Cité, CNRS UMR 8104, Inserm U1016) M Maha Zohra Ladjemi (Université Paris Cité, CNRS UMR 8104, Inserm U1016) D Darko Stojkov S Shida Yousefi (Institute of Pharmacology, University of Bern) P Philippe Frachet (Université Grenoble Alpes, CNRS, Institute of Structural Biology (IBS) - UMR 5075) L Lisa Krafft (Institute of Immunology, University of Münster) L Laura Tiberio (Department of Molecular and Translational Medicine, University of Brescia) D Daniela Bosisio (Department of Molecular and Translational Medicine, University of Brescia) M Muriel Andrieu (Université Paris Cité, CNRS UMR 8104, Inserm U1016) S Souganya Many (Université Paris Cité, CNRS UMR 8104, Inserm U1016) V Vaarany Karunanithy (Université Paris Cité, CNRS UMR 8104, Inserm U1016) K Karine Bailly (Université Paris Cité, CNRS UMR 8104, Inserm U1016) T Théo Dhôte (Université Paris Cité, CNRS UMR 8104, Inserm U1016) G Giovanni Saraceni-Tasso (Université Paris Cité, CNRS UMR 8104, Inserm U1016) M Manon Castel (Université Paris Cité, CNRS UMR 8104, Inserm U1016) C Christophe Rousseau (Université Paris Cité, CNRS UMR 8104, Inserm U1016) M Marick Rodrigues Starick (Department of Pharmacology, Federal University of Santa Catarina) E Edroaldo Lummertz da Rocha (Laboratory of Systems Biology, Department of Microbiology, Immunology and Parasitology, Federal University of Santa Catarina) E Emilia Puig Lombardi (Bioinformatics Core Platform, INSERM Unit 1163, Imagine Institute, Université Paris Cité) C Cicero José Luíz dos Ramos Almeida (Center for Research in Inflammatory Diseases, Ribeirão Preto Medical School, University of São Paulo) A Anderson dos Santos Ramos (Center for Research in Inflammatory Diseases, Ribeirão Preto Medical School, University of São Paulo) F Fernando Queiroz Cunha (Center for Research in Inflammatory Diseases, Ribeirão Preto Medical School, University of São Paulo) J Jose Carlos Alves-Filho N Natália Ribeiro Cabacinha Nóbrega (Department of Morphology, Institute of Biological Sciences, Federal University of Minas Gerais) M Matheus Rodrigues Gonçalves (Department of Microbiology, Institute of Biological Sciences, Federal University of Minas Gerais) C Celso Martins Queiroz-Junior (Department of Morphology, Institute of Biological Sciences, Federal University of Minas Gerais) V Viviane Lima Batista (Department of Microbiology, Institute of Biological Sciences, Federal University of Minas Gerais) M Mauro Martins Teixeira (Department of Biochemistry and Immunology, Institute of Biological Sciences, Federal University of Minas Gerais) V Vanessa Granger (Assistance Publique–Hôpitaux de Paris (AP-HP), Department of Immunology, Bichat Hospital) S Sylvie Chollet-Martin (Assistance Publique–Hôpitaux de Paris (AP-HP), Department of Immunology, Bichat Hospital) L Luc De Chaisemartin (Assistance Publique–Hôpitaux de Paris (AP-HP), Department of Immunology, Bichat Hospital) L Luc Mouthon (Université Paris Cité, CNRS UMR 8104, Inserm U1016) A Anne Hosmalin (Université Paris Cité, CNRS UMR 8104, Inserm U1016) M Margarita Hurtado-Nedelec (Department of Immunology and Hematology, Functional Unit of Immune Dysfunctions, Hôpitaux Universitaires Paris Nord Val-de-Seine, Bichat Hospital) C Clémence Martin (Université Paris Cité, CNRS UMR 8104, Inserm U1016) F Fernando Spiller (Department of Pharmacology, Federal University of Santa Catarina) H Hans-Uwe Simon (Institute of Pharmacology, University of Bern) N Nicolas Tamassia (Department of Medicine, Section of General Pathology, University of Verona) M Marco Antonio Cassatella (Department of Medicine, Section of General Pathology, University of Verona) F Frédéric Pène (Université Paris Cité, CNRS UMR 8104, Inserm U1016) T Thomas Vogl (Institute of Immunology, University of Münster) P Pierre-Regis Burgel (Université Paris Cité, CNRS UMR 8104, Inserm U1016) V Vivian Vasconcelos Costa (Department of Morphology, Institute of Biological Sciences, Federal University of Minas Gerais) V Véronique Witko-Sarsat (Université Paris Cité, CNRS UMR 8104, Inserm U1016)

Abstract

Neutrophils are central mediators of the hyperinflammatory response in severe SARS-CoV-2 infection. We report elevated cytosolic levels of proliferating cell nuclear antigen (PCNA) in neutrophils from patients with severe and critical COVID-19, correlating with enhanced NADPH oxidase–dependent reactive oxygen species (ROS) generation and neutrophil extracellular trap (NET) formation. Using T2AA, a small-molecule inhibitor of the PCNA scaffold, we demonstrate potent suppression of NADPH oxidase activation and NET release, particularly in response to SARS-CoV-2 RNA. Mechanistically, we identify a previously unrecognized interaction between PCNA and the heterodimeric S100A8/S100A9 (calprotectin), predominantly enriched in CD16 high CD62L low neutrophils expanded during COVID-19. PCNA binds the dimeric S100A8/S100A9 complex mediated via S100A8 subunit with micromolar affinity, and this interaction is abrogated by tetramerization, suggesting regulation by intracellular calcium. Disruption of this complex by T2AA inhibited ROS production in an S100A8/S100A9-dependent manner, implicating calprotectin as a functional regulator of neutrophil activation. In a betacoronavirus mouse model, T2AA treatment attenuated lung inflammation, reduced NET and calprotectin levels, and shifted pulmonary neutrophils away from hyperactivated and immunosuppressive phenotypes, consistent with immune reprogramming toward resolution. These findings establish cytosolic PCNA as a central scaffold in neutrophil hyperactivation during COVID-19 and highlight its pharmacological disruption as a promising host-directed strategy to limit inflammation and prevent organ damage.

Article Details

Volume / Issue Vol. 122, Issue 43
Published October 28, 2025
ISSN 0027-8424
Publisher National Academy of Sciences

Authors (46)

R

Rodrigo de Oliveira Formiga

Université Paris Cité, CNRS UMR 8104, Inserm U1016

L

Lucie Pesenti

Université Paris Cité, CNRS UMR 8104, Inserm U1016

F

François Chable de la Héronnière

Université Paris Cité, CNRS UMR 8104, Inserm U1016

M

Maha Zohra Ladjemi

Université Paris Cité, CNRS UMR 8104, Inserm U1016

D

Darko Stojkov

S

Shida Yousefi

Institute of Pharmacology, University of Bern

P

Philippe Frachet

Université Grenoble Alpes, CNRS, Institute of Structural Biology (IBS) - UMR 5075

L

Lisa Krafft

Institute of Immunology, University of Münster

L

Laura Tiberio

Department of Molecular and Translational Medicine, University of Brescia

D

Daniela Bosisio

Department of Molecular and Translational Medicine, University of Brescia

M

Muriel Andrieu

Université Paris Cité, CNRS UMR 8104, Inserm U1016

S

Souganya Many

Université Paris Cité, CNRS UMR 8104, Inserm U1016

V

Vaarany Karunanithy

Université Paris Cité, CNRS UMR 8104, Inserm U1016

K

Karine Bailly

Université Paris Cité, CNRS UMR 8104, Inserm U1016

T

Théo Dhôte

Université Paris Cité, CNRS UMR 8104, Inserm U1016

G

Giovanni Saraceni-Tasso

Université Paris Cité, CNRS UMR 8104, Inserm U1016

M

Manon Castel

Université Paris Cité, CNRS UMR 8104, Inserm U1016

C

Christophe Rousseau

Université Paris Cité, CNRS UMR 8104, Inserm U1016

M

Marick Rodrigues Starick

Department of Pharmacology, Federal University of Santa Catarina

E

Edroaldo Lummertz da Rocha

Laboratory of Systems Biology, Department of Microbiology, Immunology and Parasitology, Federal University of Santa Catarina

E

Emilia Puig Lombardi

Bioinformatics Core Platform, INSERM Unit 1163, Imagine Institute, Université Paris Cité

C

Cicero José Luíz dos Ramos Almeida

Center for Research in Inflammatory Diseases, Ribeirão Preto Medical School, University of São Paulo

A

Anderson dos Santos Ramos

Center for Research in Inflammatory Diseases, Ribeirão Preto Medical School, University of São Paulo

F

Fernando Queiroz Cunha

Center for Research in Inflammatory Diseases, Ribeirão Preto Medical School, University of São Paulo

J

Jose Carlos Alves-Filho

N

Natália Ribeiro Cabacinha Nóbrega

Department of Morphology, Institute of Biological Sciences, Federal University of Minas Gerais

M

Matheus Rodrigues Gonçalves

Department of Microbiology, Institute of Biological Sciences, Federal University of Minas Gerais

C

Celso Martins Queiroz-Junior

Department of Morphology, Institute of Biological Sciences, Federal University of Minas Gerais

V

Viviane Lima Batista

Department of Microbiology, Institute of Biological Sciences, Federal University of Minas Gerais

M

Mauro Martins Teixeira

Department of Biochemistry and Immunology, Institute of Biological Sciences, Federal University of Minas Gerais

V

Vanessa Granger

Assistance Publique–Hôpitaux de Paris (AP-HP), Department of Immunology, Bichat Hospital

S

Sylvie Chollet-Martin

Assistance Publique–Hôpitaux de Paris (AP-HP), Department of Immunology, Bichat Hospital

L

Luc De Chaisemartin

Assistance Publique–Hôpitaux de Paris (AP-HP), Department of Immunology, Bichat Hospital

L

Luc Mouthon

Université Paris Cité, CNRS UMR 8104, Inserm U1016

A

Anne Hosmalin

Université Paris Cité, CNRS UMR 8104, Inserm U1016

M

Margarita Hurtado-Nedelec

Department of Immunology and Hematology, Functional Unit of Immune Dysfunctions, Hôpitaux Universitaires Paris Nord Val-de-Seine, Bichat Hospital

C

Clémence Martin

Université Paris Cité, CNRS UMR 8104, Inserm U1016

F

Fernando Spiller

Department of Pharmacology, Federal University of Santa Catarina

H

Hans-Uwe Simon

Institute of Pharmacology, University of Bern

N

Nicolas Tamassia

Department of Medicine, Section of General Pathology, University of Verona

M

Marco Antonio Cassatella

Department of Medicine, Section of General Pathology, University of Verona

F

Frédéric Pène

Université Paris Cité, CNRS UMR 8104, Inserm U1016

T

Thomas Vogl

Institute of Immunology, University of Münster

P

Pierre-Regis Burgel

Université Paris Cité, CNRS UMR 8104, Inserm U1016

V

Vivian Vasconcelos Costa

Department of Morphology, Institute of Biological Sciences, Federal University of Minas Gerais

V

Véronique Witko-Sarsat

Université Paris Cité, CNRS UMR 8104, Inserm U1016