medinfoАктуальные проблемы теоретической и клинической медициныActual Problems of Theoretical and Clinical Medicine2790-12892790-1297Казахстанско-Российский медицинский университет10.24412/2790-1289-2021-12125medinfo-75Research ArticleОБЗОР ЛИТЕРАТУРЫREVEWSНАРУШЕНИЕ АКТИВНОСТИ НЕКОТОРЫХ СУБПОПУЛЯЦИЙ Т-КЛЕТОК ПРИ ТЯЖЕЛОМ ТЕЧЕНИИ COVID-19VIOLATION OF THE ACTIVITY OF CERTAIN T-CELL SUBPOPULATIONS IN SEVERE COVID-19КанС. А.KanS. A.ОстапчукЕ. О.OstapchukE. O.«Институт молекулярной биологии и биохимии им. М.А. Айтхожина»Казахстан«M.A. Aitkhozhin Institute of Molecular Biology and Biochemistry»Kazakhstan202120102022012126Copyright © Кан С.А., Остапчук Е.О., 20222022Кан С.А., Остапчук Е.О.Kan S.A., Ostapchuk E.O.Данная работа распространяется под лицензией Creative Commons Attribution 4.0.This work is licensed under a Creative Commons Attribution 4.0 License.https://kazrosmedjournal.krmu.edu.kz/jour/article/view/75

COVID-19 является высокопатогенной вирусной инфекцией, вызываемой коронавирусом SARS-CoV-2. Изучение адаптивного иммунитета к SARS-CoV-2 важно для понимания патогенеза новой коронавирусной инфекции COVID-19, разработки вакцин и новых терапевтических подходов лечения. В данном обзоре рассматривается Т-клеточный иммунный ответ на SARS-CoV-2, а также активность Т-регуляторных клеток при COVID-19, опосредованные этими клетками иммунопатологические механизмы, участвующие в развитии заболевания и потенциальные новые терапевтические подходы

COVID-19 is a highly pathogenic viral infection caused by the SARS-CoV-2 coronavirus. The study of adaptive immunity to SARSCoV-2 is important for understanding the pathogenesis of the novel coronavirus infection COVID-19, the development of vaccines and new therapeutic treatment approaches. This review examines the T-cell immune response to SARS-CoV-2, as well as the activity of T-regulatory cells in COVID-19, the immunopathological mechanisms mediated by these cells involved in the development of the disease, and potential new therapeutic approaches

COVID-19SARS-CoV-2CD4адаптивный иммунитетTreg-клеткиCOVID-19SARS-CoV-2CD4adaptive immunityTreg cellsReferencesZhu N., Zhang D., Wang W., Li X., Yang B., Song J., Zhao X., Huang B., Shi W., Lu R., et al. A Novel Coronavirus from Patients with Pneumonia in China, 2019 // N. Engl. J. Med. - 2020. - №382. - С. 727 - 733.Zhu N., Zhang D., Wang W., Li X., Yang B., Song J., Zhao X., Huang B., Shi W., Lu R., et al. A Novel Coronavirus from Patients with Pneumonia in China, 2019 // N. Engl. J. Med. - 2020. - №382. - С. 727 - 733.Wu F., Zhao S., Yu B., Chen Y.M., Wang W., Song Z.G., Hu Y., Tao Z.W., Tian J.H., Pei Y.Y., et al. A new coronavirus associated with human respiratory disease in China//Nature. - 2020. - №579 (7798). С. 265 - 269.Wu F., Zhao S., Yu B., Chen Y.M., Wang W., Song Z.G., Hu Y., Tao Z.W., Tian J.H., Pei Y.Y., et al. A new coronavirus associated with human respiratory disease in China//Nature. - 2020. - №579 (7798). С. 265 - 269.Zhou P., Yang X.L., Wang X.G., Hu B., Zhang L., Zhang W., Si H.R., Zhu Y., Li B., Huang C.L., et al. A pneumonia outbreak associated with a new coronavirus of probable bat origin // Nature. - 2020. - № 579 (7798). - С. 270 - 273.Zhou P., Yang X.L., Wang X.G., Hu B., Zhang L., Zhang W., Si H.R., Zhu Y., Li B., Huang C.L., et al. A pneumonia outbreak associated with a new coronavirus of probable bat origin // Nature. - 2020. - № 579 (7798). - С. 270 - 273.Wan Y., Shang J., Graham R., Baric R.S., Li F. Receptor recognition by novel coronavirus from Wuhan: An analysis based on decade-long structural studies of SARS// J. Virol. - 2020. - №94. - С. 127-20.Wan Y., Shang J., Graham R., Baric R.S., Li F. Receptor recognition by novel coronavirus from Wuhan: An analysis based on decade-long structural studies of SARS// J. Virol. - 2020. - №94. - С. 127-20.Letko M., Marzi A., Munster V. Functional assessment of cell entry and receptor usage for SARS-CoV-2 and other lineage B betacoronaviruses// Nat. Microbiol. - 2020. - С. 1 - 8.Letko M., Marzi A., Munster V. Functional assessment of cell entry and receptor usage for SARS-CoV-2 and other lineage B betacoronaviruses// Nat. Microbiol. - 2020. - С. 1 - 8.Li W., Moore M.J., Vasilieva N., Sui J., Wong S.K., Berne M.A., Somasundaran M., Sullivan J.L., Luzuriaga K., Greenough T.C. Angiotensin-converting enzyme 2 is a functional receptor for the SARS coronavirus//Nature. - 2003. № 426. - С. 450 – 454.Li W., Moore M.J., Vasilieva N., Sui J., Wong S.K., Berne M.A., Somasundaran M., Sullivan J.L., Luzuriaga K., Greenough T.C. Angiotensin-converting enzyme 2 is a functional receptor for the SARS coronavirus//Nature. - 2003. № 426. - С. 450 – 454.Kumar B.V., Connors T.J., Farber D.L. Human T Cell Development, Localization, and Function throughout Life// Immunity. - 2018. - № 48. - С. 202 - 13.Kumar B.V., Connors T.J., Farber D.L. Human T Cell Development, Localization, and Function throughout Life// Immunity. - 2018. - № 48. - С. 202 - 13.Grant E.J., Nüssing S., Sant S., Clemens E.B., Kedzierska K. The role of CD27 in anti-viral T-cell immunity// Curr Opin Virol. - 2017. - №22. - С.77 - 88.Grant E.J., Nüssing S., Sant S., Clemens E.B., Kedzierska K. The role of CD27 in anti-viral T-cell immunity// Curr Opin Virol. - 2017. - №22. - С.77 - 88.Saeidi A., Zandi K., Cheok Y.Y., Saeidi H., Wong W.F., Lee CYQ, et al. Shankar E.M. T-Cell Exhaustion in Chronic Infections: Reversing the State of Exhaustion and Reinvigorating Optimal Protective Immune Responses//Front Immunol. - 2018. - № 9. - С. 2569.Saeidi A., Zandi K., Cheok Y.Y., Saeidi H., Wong W.F., Lee CYQ, et al. Shankar E.M. T-Cell Exhaustion in Chronic Infections: Reversing the State of Exhaustion and Reinvigorating Optimal Protective Immune Responses//Front Immunol. - 2018. - № 9. - С. 2569.Zheng M., Gao Y., Wang G., Song G., Liu S., Sun D., et al. Functional exhaustion of antiviral lymphocytes in COVID-19 patients // Cell Mol Immunol. - 2020. - №17. - С. 533 - 5.Zheng M., Gao Y., Wang G., Song G., Liu S., Sun D., et al. Functional exhaustion of antiviral lymphocytes in COVID-19 patients // Cell Mol Immunol. - 2020. - №17. - С. 533 - 5.Qin C., Zhou L., Hu Z., Zhang S., Yang S., Tao Y., et al. Dysregulation of immune response in patients with COVID-19 in Wuhan, China // Clin Infect Dis. - 2020. - №71(15). - С. 762 - 8.Qin C., Zhou L., Hu Z., Zhang S., Yang S., Tao Y., et al. Dysregulation of immune response in patients with COVID-19 in Wuhan, China // Clin Infect Dis. - 2020. - №71(15). - С. 762 - 8.Ruan Q., Yang K., Wang W., Jiang L., Song J. Clinical predictors of mortality due to COVID-19 based on an analysis of data of 150 patients from Wuhan, China // Intens Care Med. - 2020. - № 22. - С. 77-88.Ruan Q., Yang K., Wang W., Jiang L., Song J. Clinical predictors of mortality due to COVID-19 based on an analysis of data of 150 patients from Wuhan, China // Intens Care Med. - 2020. - № 22. - С. 77-88.Cao X. COVID-19: immunopathology and its implications for therapy // Nat Rev Immunol. - 2020. - № 20. - С. 269 - 70.Cao X. COVID-19: immunopathology and its implications for therapy // Nat Rev Immunol. - 2020. - № 20. - С. 269 - 70.Wing J.B., Tay C., Sakaguchi S. Control of regulatory T cells by co-signal molecules. In: Azuma M., Yagita H., editors // Advances in Experimental Medicine and Biology. - 2019. - vol. 1189. - p. 179 - 210.Wing J.B., Tay C., Sakaguchi S. Control of regulatory T cells by co-signal molecules. In: Azuma M., Yagita H., editors // Advances in Experimental Medicine and Biology. - 2019. - vol. 1189. - p. 179 - 210.Chen G., Wu D., Guo W., Cao Y., Huang D., Wang H. et al. Clinical and immunological features of severe and moderate coronavirus disease 2019 // J Clin Invest. - 2020. - № 130. - С. 2620 - 9.Chen G., Wu D., Guo W., Cao Y., Huang D., Wang H. et al. Clinical and immunological features of severe and moderate coronavirus disease 2019 // J Clin Invest. - 2020. - № 130. - С. 2620 - 9.Wang F., Hou H., Luo Y., Tang G., Wu S., Huang M., et al. The laboratory tests and host immunity of COVID-19 patients with different severity of illness // JCI Insight. - 2020. - № 5. - С. 10.Wang F., Hou H., Luo Y., Tang G., Wu S., Huang M., et al. The laboratory tests and host immunity of COVID-19 patients with different severity of illness // JCI Insight. - 2020. - № 5. - С. 10.Meckiff B.J. Single-cell transcriptomic analysis of SARS-CoV-2 reactive CD4+ T cells // SSRN. – 2020.Meckiff B.J. Single-cell transcriptomic analysis of SARS-CoV-2 reactive CD4+ T cells // SSRN. – 2020.Belkaid Y., Tarbell K. Regulatory T cells in the control of host-microorganism interactions // Annu. Rev. Immunol. - 2009. - №27. - С. 551 - 589.Belkaid Y., Tarbell K. Regulatory T cells in the control of host-microorganism interactions // Annu. Rev. Immunol. - 2009. - №27. - С. 551 - 589.Suvas S. CD4+CD25+ regulatory T cells control the severity of viral immunoinflammatory lesions // J. Immunol. - 2004. - №172. - С. 4123 - 4132.Suvas S. CD4+CD25+ regulatory T cells control the severity of viral immunoinflammatory lesions // J. Immunol. - 2004. - №172. - С. 4123 - 4132.Josefowicz S.Z., Lu L. - F., Rudensky A.Y. Regulatory T Cells: Mechanisms of Differentiation and Function// Annu. Rev. Immunol. - 2012. - №30. - С. 531 - 564.Josefowicz S.Z., Lu L. - F., Rudensky A.Y. Regulatory T Cells: Mechanisms of Differentiation and Function// Annu. Rev. Immunol. - 2012. - №30. - С. 531 - 564.Kalfaoglu B., Almeida-Santos J., Adele Tye C., Satou Y., Ono M. T-cell hyperactivation and paralysis in severe COVID - 19 infection revealed by single-cell analysis // Front Immunol. - 2020. - №11. - С. 589380.Kalfaoglu B., Almeida-Santos J., Adele Tye C., Satou Y., Ono M. T-cell hyperactivation and paralysis in severe COVID - 19 infection revealed by single-cell analysis // Front Immunol. - 2020. - №11. - С. 589380.Ono M., Tanaka R.J. Controversies concerning thymusderived regulatory T cells: Fundamental issues and a new perspective// Immunol. Cell Biol. -2016. - №94. - С. 3 - 10.Ono M., Tanaka R.J. Controversies concerning thymusderived regulatory T cells: Fundamental issues and a new perspective// Immunol. Cell Biol. -2016. - №94. - С. 3 - 10.Valencia X., Stephens G., Goldbach-Mansky R., Wilson M., Shevach E.M., Lipsky P.E. TNF downmodulates the function of human CD4+CD25hi T-regulatory cells // Blood. - 2006. - №108. - С. 253-261.Valencia X., Stephens G., Goldbach-Mansky R., Wilson M., Shevach E.M., Lipsky P.E. TNF downmodulates the function of human CD4+CD25hi T-regulatory cells // Blood. - 2006. - №108. - С. 253-261.Nie H., Zheng Y., Li R., Guo T.B., He D., Fang L., Liu X., Xiao L., Chen X., Wan B., et al. Phosphorylation of FOXP3 controls regulatory T cell function and is inhibited by TNF-α in rheumatoid arthritis//Nat. Med. - 2013. - №19. - С. 322 - 328.Nie H., Zheng Y., Li R., Guo T.B., He D., Fang L., Liu X., Xiao L., Chen X., Wan B., et al. Phosphorylation of FOXP3 controls regulatory T cell function and is inhibited by TNF-α in rheumatoid arthritis//Nat. Med. - 2013. - №19. - С. 322 - 328.D’Alessio F.R., Tsushima K., Aggarwal N.R., West E.E., Willett M.H., Britos M.F., Pipeling M.R., Brower R.G., Tuder R.M., McDyer J.F., et al. CD4+CD25+Foxp3+ Tregs resolve experimental lung injury in mice and are present in humans with acute lung injury // J. Clin. Investig. - 2009. - №119. - С. 2898 - 2913.D’Alessio F.R., Tsushima K., Aggarwal N.R., West E.E., Willett M.H., Britos M.F., Pipeling M.R., Brower R.G., Tuder R.M., McDyer J.F., et al. CD4+CD25+Foxp3+ Tregs resolve experimental lung injury in mice and are present in humans with acute lung injury // J. Clin. Investig. - 2009. - №119. - С. 2898 - 2913.Lyu M.A., Khoury J.D., Nishimoto M., Zeng K., Huang M., Iyer S.P., Parmar S. Single Injection of Cord Blood Regulatory T Cells Can Delay the Manifestations of Systemic Lupus Erythematosus // Blood. - 2019. - №134. - С. 1938.Lyu M.A., Khoury J.D., Nishimoto M., Zeng K., Huang M., Iyer S.P., Parmar S. Single Injection of Cord Blood Regulatory T Cells Can Delay the Manifestations of Systemic Lupus Erythematosus // Blood. - 2019. - №134. - С. 1938.Kadia T.M., Ma H., Zeng K., Nishimoto M., Lyu M.A., Huang M., Yilmaz M., DiNardo C.D., Issa G.C., Parmar S., et al. Phase I Clinical Trial of CK0801 (cord blood regulatory T cells) in Patients with Bone Marrow Failure Syndrome (BMF) Including Aplastic Anemia, Myelodysplasia and Myelofibrosis// Blood. - 2019. - №134. - С. 1221.Kadia T.M., Ma H., Zeng K., Nishimoto M., Lyu M.A., Huang M., Yilmaz M., DiNardo C.D., Issa G.C., Parmar S., et al. Phase I Clinical Trial of CK0801 (cord blood regulatory T cells) in Patients with Bone Marrow Failure Syndrome (BMF) Including Aplastic Anemia, Myelodysplasia and Myelofibrosis// Blood. - 2019. - №134. - С. 1221.Gladstone D.E., Kim B.S., Mooney K., Karaba A.H., D’Alessio F.R. Regulatory T Cells for Treating Patients with COVID - 19 and Acute Respiratory Distress Syndrome: Two Case Reports // Ann. Intern. Med. - 2020. - V. 173. - P. 852- 853.Gladstone D.E., Kim B.S., Mooney K., Karaba A.H., D’Alessio F.R. Regulatory T Cells for Treating Patients with COVID - 19 and Acute Respiratory Distress Syndrome: Two Case Reports // Ann. Intern. Med. - 2020. - V. 173. - P. 852- 853.He R., Lu Z., Zhang L., Fan T., Xiong R., Shen X., Feng H., Meng H., Lin W., Jiang W., et al. The clinical course and its correlated immune status in COVID-19 pneumonia // J. Clin. Virol. - 2020. - №127. - С. 104361.He R., Lu Z., Zhang L., Fan T., Xiong R., Shen X., Feng H., Meng H., Lin W., Jiang W., et al. The clinical course and its correlated immune status in COVID-19 pneumonia // J. Clin. Virol. - 2020. - №127. - С. 104361.Sadeghi A., Tahmasebi S., Mahmood A., Kuznetsova M., Valizadeh H., Taghizadieh A., et al. Th17 and Treg cells function in SARS-CoV2 patients compared with healthy controls // J Cell Physiol. – 2020.Sadeghi A., Tahmasebi S., Mahmood A., Kuznetsova M., Valizadeh H., Taghizadieh A., et al. Th17 and Treg cells function in SARS-CoV2 patients compared with healthy controls // J Cell Physiol. – 2020.Xu Z., Shi L., Wang Y., Zhang J., Huang L., Zhang C., et al. Pathological findings of COVID‐19 associated with acute respiratory distress syndrome // Lancet Respir Med. - 2020. - №8. - С. 420‐422.Xu Z., Shi L., Wang Y., Zhang J., Huang L., Zhang C., et al. Pathological findings of COVID‐19 associated with acute respiratory distress syndrome // Lancet Respir Med. - 2020. - №8. - С. 420‐422.di Mauro G., Scavone C., Rafaniello C., Rossi F., Capuano A. SARS-CoV-2 infection: Response of human immune system and possible implications for the rapid test and treatment // Int Immunopharmacol. - 2020. - №84. - С. 106519.di Mauro G., Scavone C., Rafaniello C., Rossi F., Capuano A. SARS-CoV-2 infection: Response of human immune system and possible implications for the rapid test and treatment // Int Immunopharmacol. - 2020. - №84. - С. 106519.Cafarotti S. Severe acute respiratory syndrome coronavirus 2 infection and patients with lung cancer: The potential role of interleukin - 17 target therapy // J Thorac Oncol. - 2020. - №15. - С. 101-103.Cafarotti S. Severe acute respiratory syndrome coronavirus 2 infection and patients with lung cancer: The potential role of interleukin - 17 target therapy // J Thorac Oncol. - 2020. - №15. - С. 101-103.Wang W., Su B., Pang L., Qiao L., Feng Y., Ouyang Y., et al. High-dimensional immune profiling by mass cytometry revealed immunosuppression and dysfunction of immunity in COVID-19 patients // Cell Mol Immunol. - 2020. - №17. - С. 650-652.Wang W., Su B., Pang L., Qiao L., Feng Y., Ouyang Y., et al. High-dimensional immune profiling by mass cytometry revealed immunosuppression and dysfunction of immunity in COVID-19 patients // Cell Mol Immunol. - 2020. - №17. - С. 650-652.Wang F., Nie J., Wang H., Zhao Q., Xiong Y., Deng L., et al. Characteristics of peripheral lymphocyte subset alteration in COVID-19 pneumonia // J Infect Dis. - 2020 - №221.Wang F., Nie J., Wang H., Zhao Q., Xiong Y., Deng L., et al. Characteristics of peripheral lymphocyte subset alteration in COVID-19 pneumonia // J Infect Dis. - 2020 - №221.Zheng H.Y., Zhang M., Yang C.X., Zhang N., Wang X.C., Yang X.P., et al. Elevated exhaustion levels and reduced functional diversity of T cells in peripheral blood may predict severe progression in COVID‐19 patients // Cell Mol Immunol. - 2020. - №17. - С. 541-543.Zheng H.Y., Zhang M., Yang C.X., Zhang N., Wang X.C., Yang X.P., et al. Elevated exhaustion levels and reduced functional diversity of T cells in peripheral blood may predict severe progression in COVID‐19 patients // Cell Mol Immunol. - 2020. - №17. - С. 541-543.De Biasi S., Meschiari M., Gibellini L., Bellinazzi C., Borella R., Fidanza L., et al. Marked T cell activation, senescence, exhaustion and skewing towards TH17 in patients with COVID-19 pneumonia// Nat Commun. - 2020. - №11. - С. 3434.De Biasi S., Meschiari M., Gibellini L., Bellinazzi C., Borella R., Fidanza L., et al. Marked T cell activation, senescence, exhaustion and skewing towards TH17 in patients with COVID-19 pneumonia// Nat Commun. - 2020. - №11. - С. 3434.Huang C., Wang Y., Li X., Ren L., Zhao J., Hu Y., Zhang L., Fan G., Xu J., Gu X., et al. Clinical features of patients infected with 2019 novel coronavirus in Wuhan, China // Lancet. - 2020. - №395. - С. 497-506.Huang C., Wang Y., Li X., Ren L., Zhao J., Hu Y., Zhang L., Fan G., Xu J., Gu X., et al. Clinical features of patients infected with 2019 novel coronavirus in Wuhan, China // Lancet. - 2020. - №395. - С. 497-506.Wong C.K., Lam C.W.K., Wu A.K.L., Ip W.K., Lee N.L.S., Chan I.H.S., Lit L.C.W., Hui D.S.C., Chan M.H.M., Chung S.S.C., et al. Plasma inflammatory cytokines and chemokines in severe acute respiratory syndrome // Clin. Exp. Immunol. - 2004. - №136. - С. 95-103.Wong C.K., Lam C.W.K., Wu A.K.L., Ip W.K., Lee N.L.S., Chan I.H.S., Lit L.C.W., Hui D.S.C., Chan M.H.M., Chung S.S.C., et al. Plasma inflammatory cytokines and chemokines in severe acute respiratory syndrome // Clin. Exp. Immunol. - 2004. - №136. - С. 95-103.Zhou Y., Fu B., Zheng X., Wang D., Zhao C., Qi Y., Sun R., Tian Z., Xu X., Wei H. Pathogenic T cells and inflammatory monocytes incite inflammatory storm in severe COVID - 19 patients // National Science Review. - 2020. - Vol. 7. - P. 998 - 1002Zhou Y., Fu B., Zheng X., Wang D., Zhao C., Qi Y., Sun R., Tian Z., Xu X., Wei H. Pathogenic T cells and inflammatory monocytes incite inflammatory storm in severe COVID - 19 patients // National Science Review. - 2020. - Vol. 7. - P. 998 - 1002Guo C., Li B., Ma H., Wang X., Cai P., Yu Q., Zhu L., Jin L., Jiang C., Fang J., et al. Single-cell analysis of two severe COVID-19 patients reveals a monocyte-associated and tocilizumab-responding cytokine storm // Nat. Commun. - 2020. - №1. - С. 3924.Guo C., Li B., Ma H., Wang X., Cai P., Yu Q., Zhu L., Jin L., Jiang C., Fang J., et al. Single-cell analysis of two severe COVID-19 patients reveals a monocyte-associated and tocilizumab-responding cytokine storm // Nat. Commun. - 2020. - №1. - С. 3924.Parr J.B. Time to Reassess Tocilizumab’s Role in COVID-19 Pneumonia // AMA Intern Med. - 2021. - №181 (1). - С. 12-15.Parr J.B. Time to Reassess Tocilizumab’s Role in COVID-19 Pneumonia // AMA Intern Med. - 2021. - №181 (1). - С. 12-15.Davis B.K., Wen H., Ting J.P.-Y. The Inflammasome NLRs in Immunity, Inflammation, and Associated Diseases // Annu. Rev. Immunol. - 2011. - №29. - С. 707–735.Davis B.K., Wen H., Ting J.P.-Y. The Inflammasome NLRs in Immunity, Inflammation, and Associated Diseases // Annu. Rev. Immunol. - 2011. - №29. - С. 707–735.Hou W., Jin Y.H., Kang H.S., Kim B.S. Interleukin-6 (IL-6) and IL-17 Synergistically Promote Viral Persistence by Inhibiting Cellular Apoptosis and Cytotoxic T Cell Function // J. Virol. - 2014. - №88. - С. 8479.Hou W., Jin Y.H., Kang H.S., Kim B.S. Interleukin-6 (IL-6) and IL-17 Synergistically Promote Viral Persistence by Inhibiting Cellular Apoptosis and Cytotoxic T Cell Function // J. Virol. - 2014. - №88. - С. 8479.Crowe C.R., Chen K., Pociask D.A., Alcorn J.F., Krivich C., Enelow R.I., Ross T.M., Witztum J.L., Kolls J.K. Critical role of IL-17RA in immunopathology of influenza infection // J. Immunol. -2009. - №183. - С. 5301-5310.Crowe C.R., Chen K., Pociask D.A., Alcorn J.F., Krivich C., Enelow R.I., Ross T.M., Witztum J.L., Kolls J.K. Critical role of IL-17RA in immunopathology of influenza infection // J. Immunol. -2009. - №183. - С. 5301-5310.Li C.K., Wu H., Yan H., Ma S., Wang L., Zhang M., Tang X., Temperton N.J., Weiss R.A., Brenchley J.M., et al. T cell responses to whole SARS coronavirus in humans// J. Immunol. - 2008. - №18. - С. 5490 - 5500.Li C.K., Wu H., Yan H., Ma S., Wang L., Zhang M., Tang X., Temperton N.J., Weiss R.A., Brenchley J.M., et al. T cell responses to whole SARS coronavirus in humans// J. Immunol. - 2008. - №18. - С. 5490 - 5500.Yang L.T., Peng H., Zhu Z.L., Li G., Huang Z.T., Zhao Z.X., Koup R.A., Bailer R.T., Wu C.Y. Long lived effector/ central memory T-cell responses to severe acute respiratory syndrome coronavirus (SARS-CoV) S antigen in recovered SARS patients // Clin. Immunol. - 2006. - №120. - С. 171 - 178.Yang L.T., Peng H., Zhu Z.L., Li G., Huang Z.T., Zhao Z.X., Koup R.A., Bailer R.T., Wu C.Y. Long lived effector/ central memory T-cell responses to severe acute respiratory syndrome coronavirus (SARS-CoV) S antigen in recovered SARS patients // Clin. Immunol. - 2006. - №120. - С. 171 - 178.Deming D., Sheahan T., Heise M., Yount B., Davis N., Sims A., Suthar M., Harkema J., Whitmore A., Pickles R., et al. Vaccine Efficacy in Senescent Mice Challenged with Recombinant SARS-CoV Bearing Epidemic and Zoonotic Spike Variants // PLOS Medicine. - 2020. - №4 (2).Deming D., Sheahan T., Heise M., Yount B., Davis N., Sims A., Suthar M., Harkema J., Whitmore A., Pickles R., et al. Vaccine Efficacy in Senescent Mice Challenged with Recombinant SARS-CoV Bearing Epidemic and Zoonotic Spike Variants // PLOS Medicine. - 2020. - №4 (2).Yasui F., Kai C., Kitabatake M., Inoue S., Yoneda M., Yokochi S., Kase R., Sekiguchi S., Morita K., Hishima T., et al. Prior Immunization with Severe Acute Respiratory Syndrome (SARS)-Associated Coronavirus (SARS-CoV) Nucleocapsid Protein Causes Severe Pneumonia in Mice Infected with SARS-CoV // J. Immunol. - 2008. - №181. - С. 6337.Yasui F., Kai C., Kitabatake M., Inoue S., Yoneda M., Yokochi S., Kase R., Sekiguchi S., Morita K., Hishima T., et al. Prior Immunization with Severe Acute Respiratory Syndrome (SARS)-Associated Coronavirus (SARS-CoV) Nucleocapsid Protein Causes Severe Pneumonia in Mice Infected with SARS-CoV // J. Immunol. - 2008. - №181. - С. 6337.

The authors declare that there are no conflicts of interest present.