PMC:7736111 / 55001-60580 JSONTXT

{"target":"https://pubannotation.org/docs/sourcedb/PMC/sourceid/7736111","sourcedb":"PMC","sourceid":"7736111","source_url":"https://www.ncbi.nlm.nih.gov/pmc/7736111","text":"T Cell Response\nGeneration of early adaptive immune response is critical for the selective elimination of virus-infected cells and neutralization of viral antigens, thereby preventing the damage to the underlying lung parenchyma. Cytokines, chemokines, PAMPs, and DAMPs released by infected ATII and activated AMs in the lung are adequate to mount a well-coordinated and regulated adaptive immune response by priming lung resident DCs. After encountering the antigen-presenting DCs, naive CD4+ T cells differentiate into effector and memory CD4+ T cells. At least five different CD4+ T cell lineages are known (TH1, TH2, TH17, TFH, and TREG cells) with prominent roles of TH1 and TFH cells in mounting antiviral response during SARS-CoV infection (Channappanavar et al., 2014). Additionally, some studies have also shown a functional TH2 response in PBMCs derived from COVID-19 patients. Release of TH2 specific cytokines like IL-4 and IL-5 was observed in vitro after these cells were stimulated (Weiskopf et al., 2020). Similarly, these patients show enhanced production of IL-17 along with other TH17 cell-specific cytokines (Liu J. et al., 2020; Wu and Yang, 2020). These findings suggest that the TH cell response in COVID-19 patients is complex concerning other infections, and this complexity may partly depend upon the prevailing pathophysiological state of a patient.\nDuring viral infections like SARS-CoV, TH1 differentiation is influenced by IL-12 and IFN-γ secreted by DCs along with co-stimulatory signaling via B7-1/2 and CD28. Whereas IL-6 secreted by DCs influence TFH differentiation to aid in antibody secretion by B cells (Tang et al., 2008; Lau et al., 2012). Under the influence of chemokines (CCL3, CCL4, CCL5, CCL8), TH1 cells are recruited to the site of infection and are distinguished by the secretion of IL-2, IFN-γ, IL-12, and TNF-α as the main effector cytokines during SARS-CoV infections (Li et al., 2008). Similarly, naive CD8+ T cells are activated by DCs by engaging MHC-I and TCR receptors, along with CD28-B7 co-stimulatory signaling and cytokines released by CD4+ T cells. IL-2 secreted chiefly by CD4+ T cells is also implicated in their long-term maintenance and proliferation (Eickhoff et al., 2015; Hor et al., 2015). Notably, CD8+ T cells could also be activated independently of help from CD4+ T cells under conditions where a robust IFN Type I response is present (Wiesel and Oxenius, 2012). These activated CD8+ T cells [also referred to as cytotoxic T lymphocytes (CTLs)] get subsequently recruited to the effector organ under the influence of chemokines (CCL3, CCL4, CCL5, CXCL9, and CXCL10) (Nolz, 2015). At the infected site, CTLs mount an antiviral response by directly killing the infected cells via secretion of cytotoxic molecules like granzymes, perforins, granulysin, and other cytotoxic granules. Very recently, a study shows that CTLs secrete the granzymes and perforins as supramolecular attack particles (SMAPs) in a glycoprotein complex along with over 283 other proteins (including cytokines such as IFN-γ and TNF-α) (Bálint et al., 2020). It will be interesting to know whether infection by CoVs also influences the release of SMAP by CTLs.\nAnimal studies have revealed the critical molecular insights of CD4+ cells in SARS-CoV clearance and attenuation of a pathological condition. The depletion of CD4+ cells was associated with reduced virus clearance and interstitial pneumonitis (Jin et al., 2005; Wang et al., 2006). In comparison, the adoptive transfer of virus-specific CD4+ and CD8+ T cells resulted in viral clearance (Zhao et al., 2010). Similarly, clinical data has consistently shown the presence of antigen-specific CD4+ and CD8+ T cells in the recovered patients, akin to what was found in immunized animals, reviewed in Channappanavar et al. (2014). On the other hand, severe cases of SARS-CoV infection were associated with a decline in T cells, as will be discussed in later sections. Thus, based on these animal and clinical data, CD4+ T and CD8+ T cells were central to the antiviral response during SARS-CoV infection (Peng et al., 2006; Oh et al., 2012).\nA subset of primed CD4+ and CD8+ T cells differentiates into long-acting memory cells after the infection subsides. TCR-p: MHCII signaling helps in CD4+ T memory cell formation along with presence of cytokines like IL-2, IL-21 and interaction via CD40R-CD40L (Jaigirdar and MacLeod, 2015). Similarly, This CD8+ T cell transition to memory cells take place under the influence of CD8+ TREG cells via secreted IL-10 (Laidlaw et al., 2015). Long lasting CD4+ and CD8+ T memory cells were detected in the recovered SARS-CoV infected patients (Peng et al., 2006; Li et al., 2008).\nBesides, other T cells subsets which are involved in antiviral response include unconventional NKT cells (CD56+) and MAIT (mucosa-associated invariant T) cells. NKT cells act at the interface between innate and adaptive immune response and traffic to the site of infection under the influence of cytokines (Tsay and Zouali, 2018). MAIT cells reside in the mucosal lining, such as in the lungs where they serve an immunoregulatory function. Both these cell types play an essential role in the early clearance of the SARS-CoV-2, along with other T cell subsets (Grifoni et al., 2020). Strategies to enhance their function are proposed to enhance the virial clearance during COVID-19 (Cao, 2020). Role of these cells will be further discussed under the dysfunctional immune response in section “T and B Cell Response in Mild/Moderate and Recovered COVID-19 Patients.”","divisions":[{"label":"title","span":{"begin":0,"end":15}},{"label":"p","span":{"begin":16,"end":1376}},{"label":"p","span":{"begin":1377,"end":3202}},{"label":"p","span":{"begin":3203,"end":4138}},{"label":"p","span":{"begin":4139,"end":4714}}],"tracks":[{"project":"LitCovid-PD-HP","denotations":[{"id":"T14","span":{"begin":3421,"end":3445},"obj":"Phenotype"}],"attributes":[{"id":"A14","pred":"hp_id","subj":"T14","obj":"http://purl.obolibrary.org/obo/HP_0006515"},{"subj":"T14","pred":"source","obj":"LitCovid-PD-HP"}]},{"project":"LitCovid-PubTator","denotations":[{"id":"1437","span":{"begin":489,"end":492},"obj":"Gene"},{"id":"1438","span":{"begin":541,"end":544},"obj":"Gene"},{"id":"1439","span":{"begin":579,"end":582},"obj":"Gene"},{"id":"1440","span":{"begin":611,"end":614},"obj":"Gene"},{"id":"1441","span":{"begin":927,"end":931},"obj":"Gene"},{"id":"1442","span":{"begin":936,"end":940},"obj":"Gene"},{"id":"1443","span":{"begin":1076,"end":1081},"obj":"Gene"},{"id":"1444","span":{"begin":672,"end":675},"obj":"Gene"},{"id":"1445","span":{"begin":878,"end":886},"obj":"Species"},{"id":"1446","span":{"begin":1039,"end":1047},"obj":"Species"},{"id":"1447","span":{"begin":1231,"end":1239},"obj":"Species"},{"id":"1448","span":{"begin":1368,"end":1375},"obj":"Species"},{"id":"1449","span":{"begin":834,"end":837},"obj":"Chemical"},{"id":"1450","span":{"begin":106,"end":120},"obj":"Disease"},{"id":"1451","span":{"begin":282,"end":290},"obj":"Disease"},{"id":"1452","span":{"begin":728,"end":746},"obj":"Disease"},{"id":"1453","span":{"begin":869,"end":877},"obj":"Disease"},{"id":"1454","span":{"begin":1222,"end":1230},"obj":"Disease"},{"id":"1455","span":{"begin":1268,"end":1278},"obj":"Disease"},{"id":"1491","span":{"begin":1463,"end":1468},"obj":"Gene"},{"id":"1492","span":{"begin":1525,"end":1531},"obj":"Gene"},{"id":"1493","span":{"begin":1550,"end":1554},"obj":"Gene"},{"id":"1494","span":{"begin":1715,"end":1719},"obj":"Gene"},{"id":"1495","span":{"begin":1721,"end":1725},"obj":"Gene"},{"id":"1496","span":{"begin":1727,"end":1731},"obj":"Gene"},{"id":"1497","span":{"begin":1733,"end":1737},"obj":"Gene"},{"id":"1498","span":{"begin":1831,"end":1835},"obj":"Gene"},{"id":"1499","span":{"begin":1837,"end":1842},"obj":"Gene"},{"id":"1500","span":{"begin":1855,"end":1860},"obj":"Gene"},{"id":"1501","span":{"begin":1955,"end":1958},"obj":"Gene"},{"id":"1502","span":{"begin":2096,"end":2099},"obj":"Gene"},{"id":"1503","span":{"begin":2110,"end":2114},"obj":"Gene"},{"id":"1504","span":{"begin":2135,"end":2138},"obj":"Gene"},{"id":"1505","span":{"begin":2268,"end":2271},"obj":"Gene"},{"id":"1506","span":{"begin":2332,"end":2335},"obj":"Gene"},{"id":"1507","span":{"begin":2452,"end":2455},"obj":"Gene"},{"id":"1508","span":{"begin":2602,"end":2606},"obj":"Gene"},{"id":"1509","span":{"begin":2608,"end":2612},"obj":"Gene"},{"id":"1510","span":{"begin":2614,"end":2618},"obj":"Gene"},{"id":"1511","span":{"begin":2620,"end":2625},"obj":"Gene"},{"id":"1512","span":{"begin":2631,"end":2637},"obj":"Gene"},{"id":"1513","span":{"begin":3061,"end":3066},"obj":"Gene"},{"id":"1514","span":{"begin":3071,"end":3076},"obj":"Gene"},{"id":"1515","span":{"begin":1740,"end":1743},"obj":"Gene"},{"id":"1516","span":{"begin":1416,"end":1419},"obj":"Gene"},{"id":"1517","span":{"begin":1406,"end":1414},"obj":"Species"},{"id":"1518","span":{"begin":3153,"end":3157},"obj":"Species"},{"id":"1519","span":{"begin":2377,"end":2380},"obj":"Gene"},{"id":"1520","span":{"begin":1384,"end":1400},"obj":"Disease"},{"id":"1521","span":{"begin":1779,"end":1788},"obj":"Disease"},{"id":"1522","span":{"begin":1899,"end":1918},"obj":"Disease"},{"id":"1523","span":{"begin":2660,"end":2668},"obj":"Disease"},{"id":"1524","span":{"begin":2732,"end":2740},"obj":"Disease"},{"id":"1525","span":{"begin":3140,"end":3149},"obj":"Disease"},{"id":"1539","span":{"begin":3267,"end":3270},"obj":"Gene"},{"id":"1540","span":{"begin":3362,"end":3365},"obj":"Gene"},{"id":"1541","span":{"begin":4012,"end":4015},"obj":"Gene"},{"id":"1542","span":{"begin":4023,"end":4026},"obj":"Gene"},{"id":"1543","span":{"begin":3692,"end":3695},"obj":"Gene"},{"id":"1544","span":{"begin":3701,"end":3704},"obj":"Gene"},{"id":"1545","span":{"begin":3549,"end":3552},"obj":"Gene"},{"id":"1546","span":{"begin":3540,"end":3543},"obj":"Gene"},{"id":"1547","span":{"begin":3281,"end":3289},"obj":"Species"},{"id":"1548","span":{"begin":3731,"end":3739},"obj":"Species"},{"id":"1549","span":{"begin":3434,"end":3445},"obj":"Disease"},{"id":"1550","span":{"begin":3863,"end":3881},"obj":"Disease"},{"id":"1551","span":{"begin":4082,"end":4100},"obj":"Disease"},{"id":"1567","span":{"begin":4158,"end":4161},"obj":"Gene"},{"id":"1568","span":{"begin":4167,"end":4170},"obj":"Gene"},{"id":"1569","span":{"begin":4287,"end":4290},"obj":"Gene"},{"id":"1570","span":{"begin":4354,"end":4358},"obj":"Gene"},{"id":"1571","span":{"begin":4360,"end":4365},"obj":"Gene"},{"id":"1572","span":{"begin":4392,"end":4397},"obj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