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1.
Front Immunol ; 13: 984476, 2022.
Article in English | MEDLINE | ID: covidwho-2154723

ABSTRACT

Regulatory T cells that express the transcription factor Foxp3 (Treg cells) are a highly heterogenous population of immunoregulatory cells critical for maintaining immune homeostasis and preventing immunopathology during infections. Tissue resident Treg (TR-Treg) cells are maintained within nonlymphoid tissues and have been shown to suppress proinflammatory tissue resident T cell responses and promote tissue repair. Human populations are repetitively exposed to influenza infections and lung tissue resident effector T cell responses are associated with flu-induced long-term pulmonary sequelae. The kinetics of TR-Treg cell development and molecular features of TR-Treg cells during repeated and/or long-term flu infections are unclear. Utilizing a Foxp3RFP/IL-10GFP dual reporter mouse model along with intravascular fluorescent in vivo labeling, we characterized the TR-Treg cell responses to repetitive heterosubtypic influenza infections. We found lung tissue resident Treg cells accumulated and expressed high levels of co-inhibitory and co-stimulatory receptors post primary and secondary infections. Blockade of PD-1 or ICOS signaling reveals that PD-1 and ICOS signaling pathways counter-regulate TR-Treg cell expansion and IL-10 production, during secondary influenza infection. Furthermore, the virus-specific TR-Treg cell response displayed distinct kinetics, when compared to conventional CD4+ tissue resident memory T cells, during secondary flu infection. Our results provide insight into the tissue resident Foxp3+ regulatory T cell response during repetitive flu infections, which may be applicable to other respiratory infectious diseases such as tuberculosis and COVID.


Subject(s)
COVID-19 , Animals , Forkhead Transcription Factors/metabolism , Humans , Inducible T-Cell Co-Stimulator Protein/metabolism , Interleukin-10 , Mice , Orthomyxoviridae Infections , Programmed Cell Death 1 Receptor/metabolism , T-Lymphocytes, Regulatory
2.
Frontiers in immunology ; 13, 2022.
Article in English | EuropePMC | ID: covidwho-2046163

ABSTRACT

Regulatory T cells that express the transcription factor Foxp3 (Treg cells) are a highly heterogenous population of immunoregulatory cells critical for maintaining immune homeostasis and preventing immunopathology during infections. Tissue resident Treg (TR-Treg) cells are maintained within nonlymphoid tissues and have been shown to suppress proinflammatory tissue resident T cell responses and promote tissue repair. Human populations are repetitively exposed to influenza infections and lung tissue resident effector T cell responses are associated with flu-induced long-term pulmonary sequelae. The kinetics of TR-Treg cell development and molecular features of TR-Treg cells during repeated and/or long-term flu infections are unclear. Utilizing a Foxp3RFP/IL-10GFP dual reporter mouse model along with intravascular fluorescent in vivo labeling, we characterized the TR-Treg cell responses to repetitive heterosubtypic influenza infections. We found lung tissue resident Treg cells accumulated and expressed high levels of co-inhibitory and co-stimulatory receptors post primary and secondary infections. Blockade of PD-1 or ICOS signaling reveals that PD-1 and ICOS signaling pathways counter-regulate TR-Treg cell expansion and IL-10 production, during secondary influenza infection. Furthermore, the virus-specific TR-Treg cell response displayed distinct kinetics, when compared to conventional CD4+ tissue resident memory T cells, during secondary flu infection. Our results provide insight into the tissue resident Foxp3+ regulatory T cell response during repetitive flu infections, which may be applicable to other respiratory infectious diseases such as tuberculosis and COVID.

3.
Pathogens ; 11(8)2022 Aug 05.
Article in English | MEDLINE | ID: covidwho-2023970

ABSTRACT

Feline infectious peritonitis (FIP) virus is the most common infectious cause of uveitis in cats. Confirmatory diagnosis is usually only reached at postmortem examination. The relationship between the histologic inflammatory pattern, which depends on the stage of the disease, and the likelihood of detection of the viral antigen and/or RNA has not been investigated. We hypothesized that viral detection rate by either immunohistochemistry, in situ hybridization or RT-qPCR is dependent upon the predominant type of uveal inflammatory response (i.e., pyogranulomatous vs. plasmacytic). Thus, the aims of this study were to evaluate cases of FIP-induced uveitis, localize the viral antigen and RNA, and assess the relationship between the inflammatory pattern (macrophage- vs. plasma cell-rich) and the likelihood of detecting the FIP antigen and/or RNA. We evaluated 30 cats with FIP-induced uveitis. The viral antigen and/or RNA were detected within uveal macrophages in 11/30 cases, of which 8 tested positive by RT-qPCR. Correlation analysis determined a weak to moderate but significant negative correlation between the degree of plasmacytic uveal inflammation and the likelihood of detecting the FIP antigen and RNA. This study suggests that predominance of plasmacytic inflammation in cases of FIP uveitis reduces the odds of a confirmatory diagnosis through the viral detection methods available.

4.
J Vet Sci ; 23(4): e52, 2022 Jul.
Article in English | MEDLINE | ID: covidwho-1975112

ABSTRACT

This paper reports a presumptive severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2) infection in a cat. A cat with respiratory disease living with three individuals with coronavirus disease 2019 showed bilateral ground-glass opacities in the lung on X-ray and computed tomography. The clinical swabs were negative for SARS-CoV-2 RNA, but the serum was positive for SARS-CoV-2 antibodies. Interstitial pneumonia and prominent type 2 pneumocyte hyperplasia were noted on histopathology. Respiratory tissues were negative for SARS-CoV-2 RNA or antigen, but the cat was positive for feline parvovirus DNA. In conclusion, the respiratory disease and associated pathology in this cat could have been due to exposure to SARS-CoV-2.


Subject(s)
COVID-19 , Cat Diseases , Animals , Antibodies, Viral , COVID-19/veterinary , Cat Diseases/diagnostic imaging , Cats , RNA, Viral , SARS-CoV-2 , Tomography, X-Ray Computed/veterinary
5.
Veterinary Microbiology ; n/a(n/a):622-658, 2022.
Article in English | Wiley | ID: covidwho-1905768

ABSTRACT

Summary The International Committee on Taxonomy of Viruses first established the order Nidovirales in 1996. Initially, the order contained only two viral families, Coronaviridae and Arteriviridae . Since the emergence of the human severe acute respiratory syndrome-related coronavirus (SARS-CoV) responsible for the severe acute respiratory syndrome (SARS) in the spring of 2003, coronaviruses became more recognized and generated significant interest among researchers. Coronaviruses are spherical, enveloped virions with large club-shaped surface projections (peplomers) extending from the viral envelope. SARS-CoV-2 disease in hamsters is associated with high levels of virus replication and some of the histopathological changes similar to COVID-19 in humans (e.g., ground-glass opacities). ToVs have recently been classified into the new subfamily Torovirinae within the family Tobaniviridae in the suborder Tornidovirineae . ToVs are associated with enteric diseases in animals and humans, and they are recognized as pathogens of veterinary and medical importance. Since the onset of the coronavirus disease 2019 (COVID-19) pandemic, preventive social paradigms and vaccine development have undergone serious renovations, which drastically reduced the viral spread and increased collective immunity. Although the technological advancements in diagnostic systems for severe acute respiratory syndrome coronavirus-2 (SARS-CoV-2) detection are groundbreaking, the lack of sensitive, robust, and consumer-end point-of-care (POC) devices with smartphone connectivity are conspicuously felt. Despite its revolutionary impact on biotechnology and molecular diagnostics, the reverse transcription polymerase chain reaction technique as the gold standard in COVID-19 diagnosis is not suitable for rapid testing. Today's POC tests are dominated by the lateral flow assay technique, with inadequate sensitivity and lack of internet connectivity. Herein, the biosensing advancements in Internet of Things (IoT)-integrated electroanalytical tools as superior POC devices for SARS-CoV-2 detection will be demonstrated. Meanwhile, the impeding factors pivotal for the successful deployment of such novel bioanalytical devices, including the incongruous standards, redundant guidelines, and the limitations of IoT modules will be discussed.

6.
Viruses ; 14(3)2022 03 05.
Article in English | MEDLINE | ID: covidwho-1732242

ABSTRACT

Animal models recapitulating COVID-19 are critical to enhance our understanding of SARS-CoV-2 pathogenesis. Intranasally inoculated transgenic mice expressing human angiotensin-converting enzyme 2 under the cytokeratin 18 promoter (K18-hACE2) represent a lethal model of SARS-CoV-2 infection. We evaluated the clinical and virological dynamics of SARS-CoV-2 using two intranasal doses (104 and 106 PFUs), with a detailed spatiotemporal pathologic analysis of the 106 dose cohort. Despite generally mild-to-moderate pneumonia, clinical decline resulting in euthanasia or death was commonly associated with hypothermia and viral neurodissemination independent of inoculation dose. Neuroinvasion was first observed at 4 days post-infection, initially restricted to the olfactory bulb suggesting axonal transport via the olfactory neuroepithelium as the earliest portal of entry. Absence of viremia suggests neuroinvasion occurs independently of transport across the blood-brain barrier. SARS-CoV-2 tropism was neither restricted to ACE2-expressing cells (e.g., AT1 pneumocytes), nor inclusive of some ACE2-positive cell lineages (e.g., bronchiolar epithelium and brain vasculature). Absence of detectable ACE2 protein expression in neurons but overexpression in neuroepithelium suggest this as the most likely portal of neuroinvasion, with subsequent ACE2 independent lethal neurodissemination. A paucity of epidemiological data and contradicting evidence for neuroinvasion and neurodissemination in humans call into question the translational relevance of this model.


Subject(s)
COVID-19 , SARS-CoV-2 , Angiotensin-Converting Enzyme 2 , Animals , Humans , Keratin-18 , Melphalan , Mice , Mice, Transgenic , SARS-CoV-2/genetics , Viral Tropism , gamma-Globulins
7.
Emerg Microbes Infect ; 11(1): 662-675, 2022 Dec.
Article in English | MEDLINE | ID: covidwho-1665836

ABSTRACT

Severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2) is responsible for a global pandemic that has had significant impacts on human health and economies worldwide. SARS-CoV-2 is highly transmissible and the cause of coronavirus disease 2019 in humans. A wide range of animal species have also been shown to be susceptible to SARS-CoV-2 by experimental and/or natural infections. Sheep are a commonly farmed domestic ruminant that have not been thoroughly investigated for their susceptibility to SARS-CoV-2. Therefore, we performed in vitro and in vivo studies which consisted of infection of ruminant-derived cells and experimental challenge of sheep to investigate their susceptibility to SARS-CoV-2. Our results showed that sheep-derived kidney cells support SARS-CoV-2 replication. Furthermore, the experimental challenge of sheep demonstrated limited infection with viral RNA shed in nasal and oral swabs at 1 and 3-days post challenge (DPC); viral RNA was also detected in the respiratory tract and lymphoid tissues at 4 and 8 DPC. Sero-reactivity was observed in some of the principal infected sheep but not the contact sentinels, indicating that transmission to co-mingled naïve sheep was not highly efficient; however, viral RNA was detected in respiratory tract tissues of sentinel animals at 21 DPC. Furthermore, we used a challenge inoculum consisting of a mixture of two SARS-CoV-2 isolates, representatives of the ancestral lineage A and the B.1.1.7-like alpha variant of concern, to study competition of the two virus strains. Our results indicate that sheep show low susceptibility to SARS-CoV-2 infection and that the alpha variant outcompeted the lineage A strain.


Subject(s)
COVID-19 , Coinfection , Sheep/virology , Animals , COVID-19/veterinary , Coinfection/veterinary , SARS-CoV-2
8.
Emerg Microbes Infect ; 11(1): 95-112, 2022 Dec.
Article in English | MEDLINE | ID: covidwho-1541489

ABSTRACT

ABSTRACTSARS-CoV-2 was first reported circulating in human populations in December 2019 and has since become a global pandemic. Recent history involving SARS-like coronavirus outbreaks have demonstrated the significant role of intermediate hosts in viral maintenance and transmission. Evidence of SARS-CoV-2 natural infection and experimental infections of a wide variety of animal species has been demonstrated, and in silico and in vitro studies have indicated that deer are susceptible to SARS-CoV-2 infection. White-tailed deer (WTD) are amongst the most abundant and geographically widespread wild ruminant species in the US. Recently, WTD fawns were shown to be susceptible to SARS-CoV-2. In the present study, we investigated the susceptibility and transmission of SARS-CoV-2 in adult WTD. In addition, we examined the competition of two SARS-CoV-2 isolates, representatives of the ancestral lineage A and the alpha variant of concern (VOC) B.1.1.7 through co-infection of WTD. Next-generation sequencing was used to determine the presence and transmission of each strain in the co-infected and contact sentinel animals. Our results demonstrate that adult WTD are highly susceptible to SARS-CoV-2 infection and can transmit the virus through direct contact as well as vertically from doe to fetus. Additionally, we determined that the alpha VOC B.1.1.7 isolate of SARS-CoV-2 outcompetes the ancestral lineage A isolate in WTD, as demonstrated by the genome of the virus shed from nasal and oral cavities from principal infected and contact animals, and from the genome of virus present in tissues of principal infected deer, fetuses and contact animals.


Subject(s)
Animal Diseases/epidemiology , Animal Diseases/transmission , Animal Diseases/virology , COVID-19/veterinary , Deer , Pregnancy Complications, Infectious , SARS-CoV-2 , Animals , Antibodies, Neutralizing/immunology , Antibodies, Viral/immunology , Cell Line , Disease Susceptibility , Enzyme-Linked Immunosorbent Assay , Female , High-Throughput Nucleotide Sequencing , Organ Specificity , Pregnancy , RNA, Viral , SARS-CoV-2/classification , SARS-CoV-2/genetics , SARS-CoV-2/immunology , Virus Shedding
9.
Transbound Emerg Dis ; 68(6): 3443-3452, 2021 Nov.
Article in English | MEDLINE | ID: covidwho-1526424

ABSTRACT

The recently emerged novel coronavirus, SARS-CoV-2, is phylogenetically related to bat coronaviruses (CoVs), specifically SARS-related CoVs from the Eurasian bat family Rhinolophidae. As this human pandemic virus has spread across the world, the potential impacts of SARS-CoV-2 on native North American bat populations are unknown, as is the ability of North American bats to serve as reservoirs or intermediate hosts able to transmit the virus to humans or to other animal species. To help determine the impacts of the pandemic virus on North American bat populations, we experimentally challenged big brown bats (Eptesicus fuscus) with SARS-CoV-2 under BSL-3 conditions. We inoculated the bats both oropharyngeally and nasally, and over the ensuing three weeks, we measured infectivity, pathology, virus concentrations in tissues, oral and rectal virus excretion, virus transmission, and clinical signs of disease. We found no evidence of SARS-CoV-2 infection in any examined bat, including no viral excretion, no transmission, no detectable virus in tissues, and no signs of disease or pathology. Based on our findings, it appears that big brown bats are resistant to infection with the SARS-CoV-2. The potential susceptibility of other North American bat species to SARS-CoV-2 remains to be investigated.


Subject(s)
COVID-19 , Chiroptera , Coronaviridae , Animals , COVID-19/veterinary , Humans , North America/epidemiology , Phylogeny , SARS-CoV-2
10.
Cell ; 184(10): 2618-2632.e17, 2021 05 13.
Article in English | MEDLINE | ID: covidwho-1157174

ABSTRACT

The ongoing pandemic caused by severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2) is currently affecting millions of lives worldwide. Large retrospective studies indicate that an elevated level of inflammatory cytokines and pro-inflammatory factors are associated with both increased disease severity and mortality. Here, using multidimensional epigenetic, transcriptional, in vitro, and in vivo analyses, we report that topoisomerase 1 (TOP1) inhibition suppresses lethal inflammation induced by SARS-CoV-2. Therapeutic treatment with two doses of topotecan (TPT), an FDA-approved TOP1 inhibitor, suppresses infection-induced inflammation in hamsters. TPT treatment as late as 4 days post-infection reduces morbidity and rescues mortality in a transgenic mouse model. These results support the potential of TOP1 inhibition as an effective host-directed therapy against severe SARS-CoV-2 infection. TPT and its derivatives are inexpensive clinical-grade inhibitors available in most countries. Clinical trials are needed to evaluate the efficacy of repurposing TOP1 inhibitors for severe coronavirus disease 2019 (COVID-19) in humans.


Subject(s)
COVID-19/drug therapy , DNA Topoisomerases, Type I/metabolism , SARS-CoV-2/metabolism , Topoisomerase I Inhibitors/pharmacology , Topotecan/pharmacology , Animals , COVID-19/enzymology , COVID-19/pathology , Chlorocebus aethiops , Humans , Inflammation/drug therapy , Inflammation/enzymology , Inflammation/pathology , Inflammation/virology , Mesocricetus , Mice , Mice, Transgenic , THP-1 Cells , Vero Cells
11.
Emerg Microbes Infect ; 10(1): 638-650, 2021 Dec.
Article in English | MEDLINE | ID: covidwho-1127285

ABSTRACT

SARS-CoV-2 is the causative agent of COVID-19 and responsible for the current global pandemic. We and others have previously demonstrated that cats are susceptible to SARS-CoV-2 infection and can efficiently transmit the virus to naïve cats. Here, we address whether cats previously exposed to SARS-CoV-2 can be re-infected with SARS-CoV-2. In two independent studies, SARS-CoV-2-infected cats were re-challenged with SARS-CoV-2 at 21 days post primary challenge (DPC) and necropsies performed at 4, 7 and 14 days post-secondary challenge (DP2C). Sentinels were co-mingled with the re-challenged cats at 1 DP2C. Clinical signs were recorded, and nasal, oropharyngeal, and rectal swabs, blood, and serum were collected and tissues examined for histologic lesions. Viral RNA was transiently shed via the nasal, oropharyngeal and rectal cavities of the re-challenged cats. Viral RNA was detected in various tissues of re-challenged cats euthanized at 4 DP2C, mainly in the upper respiratory tract and lymphoid tissues, but less frequently and at lower levels in the lower respiratory tract when compared to primary SARS-CoV-2 challenged cats at 4 DPC. Viral RNA and antigen detected in the respiratory tract of the primary SARS-CoV-2 infected cats at early DPCs were absent in the re-challenged cats. Naïve sentinels co-housed with the re-challenged cats did not shed virus or seroconvert. Together, our results indicate that cats previously infected with SARS-CoV-2 can be experimentally re-infected with SARS-CoV-2; however, the levels of virus shed was insufficient for transmission to co-housed naïve sentinels. We conclude that SARS-CoV-2 infection in cats induces immune responses that provide partial, non-sterilizing immune protection against re-infection.


Subject(s)
Antibodies, Viral/blood , COVID-19/transmission , Disease Susceptibility/immunology , Reinfection/veterinary , Virus Shedding , Animals , Antibodies, Neutralizing/blood , Antibodies, Neutralizing/immunology , Antibodies, Viral/immunology , COVID-19/veterinary , Cats , Cell Line , Chlorocebus aethiops , RNA, Viral/isolation & purification , Reinfection/immunology , Reinfection/virology , SARS-CoV-2/immunology , Vero Cells , Viral Load
12.
Emerg Microbes Infect ; 9(1): 2322-2332, 2020 Dec.
Article in English | MEDLINE | ID: covidwho-838603

ABSTRACT

Severe Acute Respiratory Syndrome Coronavirus 2 (SARS-CoV-2) is the cause of Coronavirus Disease 2019 (COVID-19) and responsible for the current pandemic. Recent SARS-CoV-2 susceptibility studies in cats show that the virus can replicate in these companion animals and transmit to other cats. Here, we present an in-depth study of SARS-CoV-2 infection, disease and transmission in domestic cats. Cats were challenged with SARS-CoV-2 via intranasal and oral routes. One day post challenge (DPC), two sentinel cats were introduced. Animals were monitored for clinical signs, clinicopathological abnormalities and viral shedding. Postmortem examinations were performed at 4, 7 and 21 DPC. Viral RNA was not detected in blood but transiently in nasal, oropharyngeal and rectal swabs and bronchoalveolar lavage fluid as well as various tissues. Tracheobronchoadenitis of submucosal glands with the presence of viral RNA and antigen was observed in airways of the infected cats. Serology showed that both, principals and sentinels, developed antibodies to SARS-CoV-2. All animals were clinically asymptomatic during the course of the study and capable of transmitting SARS-CoV-2 to sentinels. The results of this study are critical for understanding the clinical course of SARS-CoV-2 in a naturally susceptible host species, and for risk assessment.


Subject(s)
Betacoronavirus/isolation & purification , Cat Diseases/transmission , Coronavirus Infections/transmission , Coronavirus Infections/veterinary , Disease Susceptibility , Pandemics/veterinary , Pneumonia, Viral/transmission , Pneumonia, Viral/veterinary , Animals , Antibodies, Neutralizing/blood , Antibodies, Viral/blood , Bronchoalveolar Lavage Fluid/chemistry , COVID-19 , Cat Diseases/pathology , Cat Diseases/virology , Cats , Cell Line , Chlorocebus aethiops , Coronavirus Infections/pathology , Male , Pneumonia, Viral/pathology , RNA, Viral/analysis , RNA, Viral/isolation & purification , SARS-CoV-2 , Vero Cells , Virus Replication
13.
Emerg Microbes Infect ; 9(1): 2278-2288, 2020 Dec.
Article in English | MEDLINE | ID: covidwho-811383

ABSTRACT

The emergence of SARS-CoV-2 has resulted in an ongoing global pandemic with significant morbidity, mortality, and economic consequences. The susceptibility of different animal species to SARS-CoV-2 is of concern due to the potential for interspecies transmission, and the requirement for pre-clinical animal models to develop effective countermeasures. In the current study, we determined the ability of SARS-CoV-2 to (i) replicate in porcine cell lines, (ii) establish infection in domestic pigs via experimental oral/intranasal/intratracheal inoculation, and (iii) transmit to co-housed naïve sentinel pigs. SARS-CoV-2 was able to replicate in two different porcine cell lines with cytopathic effects. Interestingly, none of the SARS-CoV-2-inoculated pigs showed evidence of clinical signs, viral replication or SARS-CoV-2-specific antibody responses. Moreover, none of the sentinel pigs displayed markers of SARS-CoV-2 infection. These data indicate that although different porcine cell lines are permissive to SARS-CoV-2, five-week old pigs are not susceptible to infection via oral/intranasal/intratracheal challenge. Pigs are therefore unlikely to be significant carriers of SARS-CoV-2 and are not a suitable pre-clinical animal model to study SARS-CoV-2 pathogenesis or efficacy of respective vaccines or therapeutics.


Subject(s)
Betacoronavirus/pathogenicity , Coronavirus Infections/veterinary , Pandemics/veterinary , Pneumonia, Viral/veterinary , Swine Diseases/virology , Animals , Betacoronavirus/genetics , Betacoronavirus/immunology , COVID-19 , Cell Line , Coronavirus Infections/immunology , Coronavirus Infections/pathology , Coronavirus Infections/transmission , Disease Models, Animal , Disease Reservoirs , Disease Susceptibility , Female , Male , Pneumonia, Viral/immunology , Pneumonia, Viral/pathology , Pneumonia, Viral/transmission , RNA, Viral/blood , Reverse Transcriptase Polymerase Chain Reaction/veterinary , SARS-CoV-2 , Swine , Swine Diseases/immunology , Swine Diseases/pathology , Swine Diseases/transmission , Virus Cultivation , Virus Replication
14.
Arch Virol ; 165(10): 2373-2377, 2020 Oct.
Article in English | MEDLINE | ID: covidwho-695405

ABSTRACT

In situ hybridization (ISH) and immunohistochemistry (IHC) are essential tools to characterize SARS-CoV-2 infection and tropism in naturally and experimentally infected animals and also for diagnostic purposes. Here, we describe three RNAscope®-based ISH assays targeting the ORF1ab, spike, and nucleocapsid genes and IHC assays targeting the spike and nucleocapsid proteins of SARS-CoV-2.


Subject(s)
Betacoronavirus/genetics , Betacoronavirus/isolation & purification , Clinical Laboratory Techniques/methods , Coronavirus Infections/diagnosis , Pneumonia, Viral/diagnosis , RNA, Viral/genetics , Animals , Antibodies, Monoclonal , Antibodies, Viral , Antisense Elements (Genetics)/genetics , COVID-19 , COVID-19 Testing , Chlorocebus aethiops , Coronavirus Infections/virology , Coronavirus Nucleocapsid Proteins , Genes, Viral , Humans , Immunohistochemistry/methods , In Situ Hybridization/methods , Nucleocapsid Proteins/genetics , Nucleocapsid Proteins/metabolism , Pandemics , Phosphoproteins , Pneumonia, Viral/virology , Polyproteins , RNA, Viral/metabolism , SARS-CoV-2 , Spike Glycoprotein, Coronavirus/genetics , Spike Glycoprotein, Coronavirus/metabolism , Vero Cells , Viral Proteins/genetics , Viral Proteins/metabolism
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