mimmunМедицинская иммунологияMedical Immunology (Russia)1563-06252313-741XSPb RAACI10.15789/10.15789/1563-0625-AAL-2569mimmun-2569Research ArticleОБЗОРЫREVIEWSАутофагия и LC3-ассоциированный фагоцитоз: сходства и различияAutophagy and LC3-associated phagocytosis: similarities and differenceshttps://orcid.org/0000-0003-4104-5443ИбрагимовБ. Р.IbragimovB. R.

Ибрагимов Б.Р. – аспирант кафедры биохимии, биотехнологии и фармакологии, младший научныйсотрудник НИЛ «Иммунопатология»

420008, Республика Татарстан, г. Казань, ул. Кремлевская, 18Тел.: 8 (999) 132-70-75

Ibragimov B.R., Postgraduate Student, Department ofBiochemistry, Biotechnology and Pharmacology, JuniorResearch Associate, Laboratory of Immunopathology

18 Kremlevskaya St., Kazan 420008 Republic of TatarstanPhone: +7 (999) 132-70-75

ibragimov94@inbox.ruhttps://orcid.org/0000-0001-8082-2511СкибоЮ. В.SkiboYu. V.

Скибо Ю.В. – к.б.н., старший научный сотрудник НИЛ«Иммунопатология»

420008, Республика Татарстан, г. Казань, ул. Кремлевская, 18

Skibo Yu.V., PhD (Biology), Senior Research Associate,Laboratory of Immunopathology

18 Kremlevskaya St., Kazan 420008 Republic of TatarstanPhone: +7 (999) 132-70-75

yuliya_ksu@mail.ruhttps://orcid.org/0000-0003-3749-3411АбрамоваЗ. И.AbramovaZ. I.

Абрамова З.И. – д.б.н., профессор кафедры биохимии, биотехнологии и фармакологии, главный научный сотрудник НИЛ «Иммунопатология»

420008, Республика Татарстан, г. Казань, ул. Кремлевская, 18

Abramova Z.I., PhD, MD (Biology), Professor, Departmentof Biochemistry, Biotechnology and Pharmacology, ChiefResearch Associate, Laboratory of Immunopathology

18 Kremlevskaya St., Kazan 420008 Republic of TatarstanPhone: +7 (999) 132-70-75

ziabramova@mail.ruИнститут фундаментальной медицины и биологии ФГАОУ ВО «Казанский (Приволжский) федеральный университет»РоссияInstitute of Fundamental Medicine and Biology, Kazan (Volga Region) Federal UniversityRussian Federation202305042023252233252Copyright © Ибрагимов Б.Р., Скибо Ю.В., Абрамова З.И., 20232023Ибрагимов Б.Р., Скибо Ю.В., Абрамова З.И.Ibragimov B.R., Skibo Y.V., Abramova Z.I.Данная работа распространяется под лицензией Creative Commons Attribution 4.0.This work is licensed under a Creative Commons Attribution 4.0 License.https://www.mimmun.ru/mimmun/article/view/2569

Ранее аутофагия представлялась в качестве механизма, используемого клеткой при дефиците питательных веществ, необходимого для сохранения гомеостаза. Результаты исследований последнего десятилетия показали, что аутофагия является более сложным, неоднозначным механизмом, активация которого зависит от природы стимула, типа иммунных клеток и конечного результата. И каноническая и схожая с ней молекулярно, но имеющая свои отличительные черты неканоническая аутофагия являются ключевыми процессами в защите организма от проникновения внутриклеточных патогенов, поддержание в клетке необходимого уровня питательных веществ и удаление поврежденных органелл и клеток. Каноническая аутофагия, вероятно, развилась как гомеостатический ответ на клеточный стресс и недостаток питательных веществ, а неканоническая – в ответ на подавление воспаления. Неканоническая аутофагия, именуемая в дальнейшем LC3-ассоциированный фагоцитоз (LAP), сочетает в себе молекулярный механизм фагоцитоза с механизмом аутофагии, характеризующейся поглощением экзогенных патогенов, формированием фагосомы (лапосом) и усиленным слиянием с лизосомами, с последующей деградацией содержимого.

Существуют различия в процессах не канонической и схожей с ней по своему механизму действия канонической аутофагии. Наличие PI3K комплексов в обоих процессах, утилизация и деградация не нужных для клетки и организма «груза» внутри самой клетки за счет лизосомальной органеллы (лизосомы), задействование практически одних и тех же белков делают механизмы схожими. Однако различия в запуске процессов, разновидность самих PI3K-комплексов (у аутофагии PI3K III класса 1-го и 2-го типа, а у LAP PI3K III класса 3-го типа), использование активных форм кислорода при LAP, разновидность использования регуляторных белков в процессах (при аутофагии это ULK1, FIP200, ATG13, Ambra1, WIPI2, ATG14; а при LC3-ассоциированном фагоцитозе это Rubicon и NOX2), разное количество слоев в мембранной структуре, в которой происходит лизис (двухмембранная аутофаголизосома и одномембранная лапосома) четко подчеркивают разновидность канонической и не канонической аутофагии. Разность выполняемых задач, а именно разновидность мишеней для утилизации (при аутофагии внутриклеточные патогены, дисфункциональные белки и органеллы, а при LAP внеклеточные патогены, апоптотические тельца, бактерии и др.) делают данные механизмы совершенно разными по своему значению.

В совокупности новые данные указывают на то, что аутофагия как каноническими, так и неканоническими путями превратилась в механизм защиты хозяина, способный противостоять иммунологическому и патогенному стрессу и опосредовать иммунологическую толерантность как к внутриклеточным, так и к внеклеточным угрозам. В представленном обзоре обсуждаются принципиальные молекулярные отличия каждого из механизмов, а также их роли в иммунитете с учетом последних литературных данных.

Previously, autophagy was termed as a mechanism used by the cells with a lack of essential nutrients supporting homeostasis. Over the decade of studies, autophagy proved to be a more complex, ambiguous mechanism. Its activation depends on the nature of stimulus, type of immune cells and the final result. Both canonical and non-canonical autophagy, being similar in molecular events, but showing their own distinctive features, are key processes in protecting the body from penetration of intracellular pathogens, maintaining the required level of nutrients in the cell, and removing damaged organelles and cells. Canonical autophagy probably evolved as a homeostatic response to cellular stress and nutritional deficiencies, whereas non-canonical autophagy emerged as a response to suppression of inflammation. Non-canonical autophagy, hereinafter referred to as LC3-associated phagocytosis (LAP), combines the molecular mechanism of phagocytosis with an autophagy mechanism characterized by ingestion of exogenous pathogens, formation of phagosomes (laposomes) and enhanced fusion with lysosomes, followed by degradation of their contents.

Significant differences were found between the processes of LAP- and canonical autophagy, which are similar in its mechanism of action. The presence of PI3K complexes in both processes, utilization and intracellular degradation of the “cargo” which is not required for the cells and organism proceeding in the lysosomes, and involvement of almost the same proteins provide similarity of their mechanisms. However, there are differences in the initiation of the processes, e.g., different types of PI3K complexes (in autophagy, PI3K III class 1 and 2 types; in LAP PI3K III, class 3 type), usage of reactive oxygen species in LAP, different types of regulatory proteins involved (ULK1, FIP200, ATG13 , Ambra1, WIPI2, ATG14 in autophagy; and Rubicon and NOX2 in LC3-associated phagocytosis), different number of layers in the membrane structure in which lysis occurs (double-membrane autophagolysosome and single-layer membrane in laposomes) clearly depict the variety of canonical and non-canonical autophagy. The two pathways are directed for different types of biological objects, i.e., intracellular pathogens, dysfunctional proteins and organelles in autophagy, and extracellular pathogens, apoptotic bodies, bacteria, utilized in LAP, thus making these mechanisms completely different in their significance.

Collectively, the new data indicate that autophagy performed via both canonical and non-canonical pathways, has evolved into a host defense mechanism capable of resisting immunological and pathogenic stress and mediating immunological tolerance to both intra- and extracellular threats. The present review discusses fundamental molecular differences between these mechanisms, as well as their role in immunity, based on the latest literature data.

аутофагияLC3-ассоциированный фагоцитозLC3Beclin-1Vps34аутоиммунитетвоспалениеautophagyLC3-associated phagocytosisLC3Beclin-1Vps34autoimmunityinflammationReferencesДеев Р.В., Билялов А.И., Жампеисов Т.М. Современные представления о клеточной гибели // Гены и Клетки, 2018. Т. 13, № 1. С. 6-19.Deev R.V., Bilyalov A.I., Zhampeisov T.M. Modern ideas about cell death. Geny i Kletki = Genes and Cells, 2018, Vol. 13, no. 1, pp. 6-19. (In Russ.)Alissafi T., Banos A., Boon L., Sparwasser T., Ghigo A., Wing K., Vassilopoulos D., Boumpas D., Chavakis T., Cadwell K., Verginis P. Tregs restrain dendritic cell autophagy to ameliorate autoimmunity. J. Clin. Invest., 2017, Vol. 127, no. 7, pp. 2789-2804.Alissafi T., Banos A., Boon L., Sparwasser T., Ghigo A., Wing K., Vassilopoulos D., Boumpas D., Chavakis T., Cadwell K., Verginis P. Tregs restrain dendritic cell autophagy to ameliorate autoimmunity. J. Clin. Invest., 2017, Vol. 127, no. 7, pp. 2789-2804.Bhattacharya A., Parillon X., Zeng S., Han S., Eissa N.T. Deficiency of autophagy in dendritic cells protects against experimental autoimmune encephalomyelitis. J. Biol. Chem., 2014, Vol. 289, no. 38, pp. 26525-26532.Bhattacharya A., Parillon X., Zeng S., Han S., Eissa N.T. Deficiency of autophagy in dendritic cells protects against experimental autoimmune encephalomyelitis. J. Biol. Chem., 2014, Vol. 289, no. 38, pp. 26525-26532.Birmingham C.L., Canadien V., Gouin E., Troy E.B., Yoshimori T., Cossart P., Higgins D.E., Brumell J.H. Listeria monocytogenes evades killing by autophagy during colonization of host cells. Autophagy, 2007, Vol. 3, no. 5, pp. 442-451.Birmingham C.L., Canadien V., Gouin E., Troy E.B., Yoshimori T., Cossart P., Higgins D.E., Brumell J.H. Listeria monocytogenes evades killing by autophagy during colonization of host cells. Autophagy, 2007, Vol. 3, no. 5, pp. 442-451.Burman C., Ktistakis N.T. Regulation of autophagy by phosphatidylinositol 3-phosphate. FEBS Lett., 2010, Vol. 584, no. 7, pp. 1302-1312.Burman C., Ktistakis N.T. Regulation of autophagy by phosphatidylinositol 3-phosphate. FEBS Lett., 2010, Vol. 584, no. 7, pp. 1302-1312.Castillo E.F., Dekonenko A., Arko-Mensah J., Mandell M.A., Dupont N., Jiang S., Delgado-Vargas M., Timmins G.S., Bhattacharya D., Yang H., Hutt J., Lyons C.R., Dobos K.M., Deretic V. Autophagy protects against active tuberculosis by suppressing bacterial burden and inflammation. Proc. Natl Acad. Sci. USA, 2012, Vol. 109, no. 46, pp. 168-176.Castillo E.F., Dekonenko A., Arko-Mensah J., Mandell M.A., Dupont N., Jiang S., Delgado-Vargas M., Timmins G.S., Bhattacharya D., Yang H., Hutt J., Lyons C.R., Dobos K.M., Deretic V. Autophagy protects against active tuberculosis by suppressing bacterial burden and inflammation. Proc. Natl Acad. Sci. USA, 2012, Vol. 109, no. 46, pp. 168-176.Chan E.Y., Kir S., Tooze S.A. siRNA screening of the kinome identifies ULK1 as a multidomain modulator of autophagy. J. Biol. Chem., 2007, Vol. 282, no. 35, pp. 25464-25474.Chan E.Y., Kir S., Tooze S.A. siRNA screening of the kinome identifies ULK1 as a multidomain modulator of autophagy. J. Biol. Chem., 2007, Vol. 282, no. 35, pp. 25464-25474.Cheung P.F., Yang J., Fang R., Borgers A., Krengel K., Stoffel A., Althoff K., Yip C.W., Siu E.H.L., Ng L.W.C., Lang K.S., Cham L.B., Engel D.R., Soun C., Cima I., Scheffler B., Striefler J.K., Sinn M., Bahra M., Pelzer U., Oettle H., Markus P., Smeets E.M.M., Aarntzen E.H.J.G., Savvatakis K., Liffers S.T., Lueong S.S., Neander C., Bazarna A., Zhang X., Paschen A., Crawford H.C., Chan A.W.H., Cheung S.T., Siveke J.T. Progranulin mediates immune evasion of pancreatic ductal adenocarcinoma through regulation of MHCI expression. Nat. Commun., 2022, Vol. 13, no. 1, 156. doi: 10.1038/s41467-021-27088-9.Cheung P.F., Yang J., Fang R., Borgers A., Krengel K., Stoffel A., Althoff K., Yip C.W., Siu E.H.L., Ng L.W.C., Lang K.S., Cham L.B., Engel D.R., Soun C., Cima I., Scheffler B., Striefler J.K., Sinn M., Bahra M., Pelzer U., Oettle H., Markus P., Smeets E.M.M., Aarntzen E.H.J.G., Savvatakis K., Liffers S.T., Lueong S.S., Neander C., Bazarna A., Zhang X., Paschen A., Crawford H.C., Chan A.W.H., Cheung S.T., Siveke J.T. Progranulin mediates immune evasion of pancreatic ductal adenocarcinoma through regulation of MHCI expression. Nat. Commun., 2022, Vol. 13, no. 1, 156. doi: 10.1038/s41467-021-27088-9.Chu H., Khosravi A., Kusumawardhani I.P., Kwon A.H., Vasconcelos A.C., Cunha L.D., Mayer A.E., Shen Y., Wu W.L., Kambal A., Targan S.R., Xavier R.J., Ernst P.B., Green D.R., McGovern D.P., Virgin H.W., Mazmanian S.K. Gene-microbiota interactions contribute to the pathogenesis of inflammatory bowel disease. Science, 2016, Vol. 352, no. 6289, pp. 1116-1120.Chu H., Khosravi A., Kusumawardhani I.P., Kwon A.H., Vasconcelos A.C., Cunha L.D., Mayer A.E., Shen Y., Wu W.L., Kambal A., Targan S.R., Xavier R.J., Ernst P.B., Green D.R., McGovern D.P., Virgin H.W., Mazmanian S.K. Gene-microbiota interactions contribute to the pathogenesis of inflammatory bowel disease. Science, 2016, Vol. 352, no. 6289, pp. 1116-1120.Cresswell P. A personal retrospective on the mechanisms of antigen processing. Immunogenetics, 2019, Vol. 71, no. 3, pp. 141-160.Cresswell P. A personal retrospective on the mechanisms of antigen processing. Immunogenetics, 2019, Vol. 71, no. 3, pp. 141-160.Crotzer V.L., Blum J.S. Autophagy and its role in MHC-mediated antigen presentation. J. Immunol., 2009, Vol. 182, no. 6, pp. 3335-3341.Crotzer V.L., Blum J.S. Autophagy and its role in MHC-mediated antigen presentation. J. Immunol., 2009, Vol. 182, no. 6, pp. 3335-3341.de Castro C.P., Jones S.A., Cheallaigh C.N., Hearnden C.A., Williams L., Winter J., Lavelle E.C., Mills K.H., Harris J. Autophagy regulates IL-23 secretion and innate T cell responses through effects on IL-1 secretion. J. Immunol., 2012, Vol. 189, no. 8, pp. 4144-4153.de Castro C.P., Jones S.A., Cheallaigh C.N., Hearnden C.A., Williams L., Winter J., Lavelle E.C., Mills K.H., Harris J. Autophagy regulates IL-23 secretion and innate T cell responses through effects on IL-1 secretion. J. Immunol., 2012, Vol. 189, no. 8, pp. 4144-4153.Dengjel J., Schoor O., Fischer R., Reich M., Kraus M., Müller M., Kreymborg K., Altenberend F., Brandenburg J., Kalbacher H., Brock R., Driessen C., Rammensee H.G., Stevanovic S. Autophagy promotes MHC class II presentation of peptides from intracellular source proteins. Proc. Natl Acad. Sci. USA, 2005, Vol. 102, no. 22, pp. 7922-7927.Dengjel J., Schoor O., Fischer R., Reich M., Kraus M., Müller M., Kreymborg K., Altenberend F., Brandenburg J., Kalbacher H., Brock R., Driessen C., Rammensee H.G., Stevanovic S. Autophagy promotes MHC class II presentation of peptides from intracellular source proteins. Proc. Natl Acad. Sci. USA, 2005, Vol. 102, no. 22, pp. 7922-7927.Dersh D., Hollý J., Yewdell J.W. A few good peptides: MHC class I-based cancer immunosurveillance and immunoevasion. Nat.Rev. Immunol., 2021, Vol. 21, no. 2, pp. 116-128.Dersh D., Hollý J., Yewdell J.W. A few good peptides: MHC class I-based cancer immunosurveillance and immunoevasion. Nat.Rev. Immunol., 2021, Vol. 21, no. 2, pp. 116-128.Dikic I., Elazar Z. Mechanism and medical implications of mammalian autophagy. Nat. Rev. Mol. Cell Biol., 2018, Vol. 19, no. 6, pp. 349-364.Dikic I., Elazar Z. Mechanism and medical implications of mammalian autophagy. Nat. Rev. Mol. Cell Biol., 2018, Vol. 19, no. 6, pp. 349-364.Durgan J., Lystad A.H., Sloan K., Carlsson S.R., Wilson M.I., Marcassa E., Ulferts R., Webster J., LopezClavijo A.F., Wakelam M.J., Beale R., Simonsen A., Oxley D., Florey O. Non-canonical autophagy drives alternative ATG8 conjugation to phosphatidylserine. Mol. Cell, 2021, Vol. 81, no. 9, pp. 2031-2040.Durgan J., Lystad A.H., Sloan K., Carlsson S.R., Wilson M.I., Marcassa E., Ulferts R., Webster J., LopezClavijo A.F., Wakelam M.J., Beale R., Simonsen A., Oxley D., Florey O. Non-canonical autophagy drives alternative ATG8 conjugation to phosphatidylserine. Mol. Cell, 2021, Vol. 81, no. 9, pp. 2031-2040.Filomeni G., de Zio D., Cecconi F. Oxidative stress and autophagy: the clash between damage and metabolic needs. Cell Death Differ., 2015, Vol. 22, no. 3, pp. 377-388.Filomeni G., de Zio D., Cecconi F. Oxidative stress and autophagy: the clash between damage and metabolic needs. Cell Death Differ., 2015, Vol. 22, no. 3, pp. 377-388.Florey O., Gammoh N., Kim S.E., Jiang X., Overholtzer M. V-ATPase and osmotic imbalances activate endolysosomal LC3 lipidation. Autophagy, 2015, Vol. 11, no. 1, pp. 88-99.Florey O., Gammoh N., Kim S.E., Jiang X., Overholtzer M. V-ATPase and osmotic imbalances activate endolysosomal LC3 lipidation. Autophagy, 2015, Vol. 11, no. 1, pp. 88-99.Ganley I.G., Lam du H., Wang J., Ding X., Chen S., Jiang X. ULK1.ATG13.FIP200 complex mediates mTOR signaling and is essential for autophagy. J. Biol. Chem., 2009, Vol. 284, no. 18, pp. 12297-12305.Ganley I.G., Lam du H., Wang J., Ding X., Chen S., Jiang X. ULK1.ATG13.FIP200 complex mediates mTOR signaling and is essential for autophagy. J. Biol. Chem., 2009, Vol. 284, no. 18, pp. 12297-12305.Gluschko A., Herb M., Wiegmann K., Krut O., Neiss W.F., Utermöhlen O., Krönke M., Schramm M. The β 2 Integrin Mac-1 Induces Protective LC3-Associated Phagocytosis of Listeria monocytogenes. Cell Host Microbe, 2018, Vol. 23, no. 3, pp. 324-337.Gluschko A., Herb M., Wiegmann K., Krut O., Neiss W.F., Utermöhlen O., Krönke M., Schramm M. The β 2 Integrin Mac-1 Induces Protective LC3-Associated Phagocytosis of Listeria monocytogenes. Cell Host Microbe, 2018, Vol. 23, no. 3, pp. 324-337.Hansen M., Rubinsztein D.C., Walker D.W. Autophagy as a promoter of longevity: insights from model organisms. Nat. Rev. Mol. Cell Biol., 2018, Vol. 19, pp. 579-593.Hansen M., Rubinsztein D.C., Walker D.W. Autophagy as a promoter of longevity: insights from model organisms. Nat. Rev. Mol. Cell Biol., 2018, Vol. 19, pp. 579-593.Harlé G., Kowalski C., Dubrot J., Brighouse D., Clavel G., Pick R., Bessis N., Niven J., Scheiermann C., Gannagé M., Hugues S. Macroautophagy in lymphatic endothelial cells inhibits T cell-mediated autoimmunity. J. Exp. Med., 2021, Vol. 218, no. 6, e20201776. doi: 10.1084/jem.20201776.Harlé G., Kowalski C., Dubrot J., Brighouse D., Clavel G., Pick R., Bessis N., Niven J., Scheiermann C., Gannagé M., Hugues S. Macroautophagy in lymphatic endothelial cells inhibits T cell-mediated autoimmunity. J. Exp. Med., 2021, Vol. 218, no. 6, e20201776. doi: 10.1084/jem.20201776.Harris J., Hartman M., Roche C., Zeng S.G., O’Shea A., Sharp F.A., Lambe E.M., Creagh E.M., Golenbock D.T., Tschopp J., Kornfeld H., Fitzgerald K.A., Lavelle E.C. Autophagy controls IL-1beta secretion by targeting pro-IL1beta for degradation. J. Biol. Chem., 2011, Vol. 286, no. 11, pp. 9587-9597.Harris J., Hartman M., Roche C., Zeng S.G., O’Shea A., Sharp F.A., Lambe E.M., Creagh E.M., Golenbock D.T., Tschopp J., Kornfeld H., Fitzgerald K.A., Lavelle E.C. Autophagy controls IL-1beta secretion by targeting pro-IL1beta for degradation. J. Biol. Chem., 2011, Vol. 286, no. 11, pp. 9587-9597.Hayashi K., Taura M., Iwasaki A. The interaction between IKKα and LC3 promotes type I interferon production through the TLR9-containing LAPosome. Sci. Signal., 2018, Vol. 11, no. 528, eaan4144. doi: 10.1126/ scisignal.aan4144.Hayashi K., Taura M., Iwasaki A. The interaction between IKKα and LC3 promotes type I interferon production through the TLR9-containing LAPosome. Sci. Signal., 2018, Vol. 11, no. 528, eaan4144. doi: 10.1126/ scisignal.aan4144.Herb M., Gluschko A., Schramm M. LC3-associated phagocytosis – The highway to hell for phagocytosed microbes. Semin. Cell Dev. Biol., 2020, Vol. 101, pp. 68-76.Herb M., Gluschko A., Schramm M. LC3-associated phagocytosis – The highway to hell for phagocytosed microbes. Semin. Cell Dev. Biol., 2020, Vol. 101, pp. 68-76.Heyworth P.G., Cross A.R., Curnutte J.T. Chronic granulomatous disease. Curr. Opin. Immunol., 2003, Vol. 15, no. 5, pp. 578-584.Heyworth P.G., Cross A.R., Curnutte J.T. Chronic granulomatous disease. Curr. Opin. Immunol., 2003, Vol. 15, no. 5, pp. 578-584.Holmström K.M., Finkel T. Cellular mechanisms and physiological consequences of redox-dependent signalling. Nat. Rev. Mol. Cell Biol., 2014, Vol. 15, no. 6, pp. 411-421.Holmström K.M., Finkel T. Cellular mechanisms and physiological consequences of redox-dependent signalling. Nat. Rev. Mol. Cell Biol., 2014, Vol. 15, no. 6, pp. 411-421.Huang L., Guo Z., Wang F., Fu L. KRAS mutation: from undruggable to druggable in cancer. Signal Transduct. Target. Ther., 2021, Vol. 6, no. 1, pp. 1-20.Huang L., Guo Z., Wang F., Fu L. KRAS mutation: from undruggable to druggable in cancer. Signal Transduct. Target. Ther., 2021, Vol. 6, no. 1, pp. 1-20.Itakura E., Kishi C., Inoue K., Mizushima N. Beclin 1 forms two distinct phosphatidylinositol 3-kinase complexes with mammalian Atg14 and UVRAG. Mol. Biol. Cell, 2008, Vol. 19, no. 12, pp. 5360-5372.Itakura E., Kishi C., Inoue K., Mizushima N. Beclin 1 forms two distinct phosphatidylinositol 3-kinase complexes with mammalian Atg14 and UVRAG. Mol. Biol. Cell, 2008, Vol. 19, no. 12, pp. 5360-5372.Johansen T., Lamark T. Selective Autophagy: ATG8 Family Proteins, LIR Motifs and Cargo Receptors. J. Mol. Biol., 2020, Vol. 432, no. 1, pp. 80-103.Johansen T., Lamark T. Selective Autophagy: ATG8 Family Proteins, LIR Motifs and Cargo Receptors. J. Mol. Biol., 2020, Vol. 432, no. 1, pp. 80-103.Kahn B.B., Alquier T., Carling D., Hardie D.G. AMP-activated protein kinase: ancient energy gauge provides clues to modern understanding of metabolism. Cell Metab., 2005, Vol. 1, no. 1, pp. 15-25.Kahn B.B., Alquier T., Carling D., Hardie D.G. AMP-activated protein kinase: ancient energy gauge provides clues to modern understanding of metabolism. Cell Metab., 2005, Vol. 1, no. 1, pp. 15-25.Keller C.W., Kotur M.B., Mundt S., Dokalis N., Ligeon L.A., Shah A.M., Prinz M., Becher B., Münz C., Lünemann J.D. CYBB/NOX2 in conventional DCs controls T cell encephalitogenicity during neuroinflammation. Autophagy, 2021, Vol. 17, no. 5, pp. 1244-1258.Keller C.W., Kotur M.B., Mundt S., Dokalis N., Ligeon L.A., Shah A.M., Prinz M., Becher B., Münz C., Lünemann J.D. CYBB/NOX2 in conventional DCs controls T cell encephalitogenicity during neuroinflammation. Autophagy, 2021, Vol. 17, no. 5, pp. 1244-1258.Keller C.W., Sina C., Kotur M.B., Ramelli G., Mundt S., Quast I., Ligeon L.A., Weber P., Becher B., Münz C., Lünemann J.D. ATG-dependent phagocytosis in dendritic cells drives myelin-specific CD4+ T cell pathogenicity during CNS inflammation. Proc. Natl Acad. Sci. USA, 2017, Vol. 114, no. 52, pp. 11228-11237.Keller C.W., Sina C., Kotur M.B., Ramelli G., Mundt S., Quast I., Ligeon L.A., Weber P., Becher B., Münz C., Lünemann J.D. ATG-dependent phagocytosis in dendritic cells drives myelin-specific CD4+ T cell pathogenicity during CNS inflammation. Proc. Natl Acad. Sci. USA, 2017, Vol. 114, no. 52, pp. 11228-11237.Kim Y.M., Jung C.H., Seo M., Kim E.K., Park J.M., Bae S.S., Kim D.H. mTORC1 phosphorylates UVRAG to negatively regulate autophagosome and endosome maturation. Mol. Cell, 2015, Vol. 57, no. 2, pp. 207-218.Kim Y.M., Jung C.H., Seo M., Kim E.K., Park J.M., Bae S.S., Kim D.H. mTORC1 phosphorylates UVRAG to negatively regulate autophagosome and endosome maturation. Mol. Cell, 2015, Vol. 57, no. 2, pp. 207-218.Kobayashi S.D., Voyich J.M., Braughton K.R., Whitney A.R., Nauseef W.M., Malech H.L., deLeo F.R. Gene expression profiling provides insight into the pathophysiology of chronic granulomatous disease. J. Immunol., 2004, Vol. 172, no. 1, pp. 636-643.Kobayashi S.D., Voyich J.M., Braughton K.R., Whitney A.R., Nauseef W.M., Malech H.L., deLeo F.R. Gene expression profiling provides insight into the pathophysiology of chronic granulomatous disease. J. Immunol., 2004, Vol. 172, no. 1, pp. 636-643.Lam G.Y., Cemma M., Muise A.M., Higgins D.E., Brumell J.H. Host and bacterial factors that regulate LC3 recruitment to Listeria monocytogenes during the early stages of macrophage infection. Autophagy, 2013, Vol. 9, no. 7, pp. 985-995.Lam G.Y., Cemma M., Muise A.M., Higgins D.E., Brumell J.H. Host and bacterial factors that regulate LC3 recruitment to Listeria monocytogenes during the early stages of macrophage infection. Autophagy, 2013, Vol. 9, no. 7, pp. 985-995.Lamprinaki D., Zhekova A., Wittmann A., James S., Dicks J., Iwakura Y., Saijo S., Wang X., Chow C.W., Roberts I., Korcsmaros T., Mayer U., Wileman T., Kawasaki N. LC3-associated phagocytosis is required for dendritic cell inflammatory cytokine response to gut commensal yeast saccharomyces cerevisiae. Immunology, 2022, Vol. 8, 1397. doi: 10.3389/fimmu.2017.01397.Lamprinaki D., Zhekova A., Wittmann A., James S., Dicks J., Iwakura Y., Saijo S., Wang X., Chow C.W., Roberts I., Korcsmaros T., Mayer U., Wileman T., Kawasaki N. LC3-associated phagocytosis is required for dendritic cell inflammatory cytokine response to gut commensal yeast saccharomyces cerevisiae. Immunology, 2022, Vol. 8, 1397. doi: 10.3389/fimmu.2017.01397.Lazarou M., Sliter D.A., Kane L.A., Sarraf S.A., Wang C., Burman J.L., Sideris D.P., Fogel A.I., Youle R.J. The ubiquitin kinase PINK1 recruits autophagy receptors to induce mitophagy. Nature, 2015, Vol. 524, no. 7565, pp. 309-314.Lazarou M., Sliter D.A., Kane L.A., Sarraf S.A., Wang C., Burman J.L., Sideris D.P., Fogel A.I., Youle R.J. The ubiquitin kinase PINK1 recruits autophagy receptors to induce mitophagy. Nature, 2015, Vol. 524, no. 7565, pp. 309-314.Lee J.P.W., Foote A., Fan H., Castro C.P., Lang T., Jones S.A., Gavrilescu N., Mills K.H.G., Leech M., Morand E.F., Harris J. Loss of autophagy enhances MIF/macrophage migration inhibitory factor release by macrophages. Autophagy, 2016, Vol. 12, no. 6, pp. 907-916.Lee J.P.W., Foote A., Fan H., Castro C.P., Lang T., Jones S.A., Gavrilescu N., Mills K.H.G., Leech M., Morand E.F., Harris J. Loss of autophagy enhances MIF/macrophage migration inhibitory factor release by macrophages. Autophagy, 2016, Vol. 12, no. 6, pp. 907-916.Leung C.S., Haigh T.A., Mackay L.K., Rickinson A.B., Taylor G.S. Nuclear location of an endogenously expressed antigen, EBNA1, restricts access to macroautophagy and the range of CD4 epitope display. Proc. Natl Acad. Sci. USA, 2010, Vol. 107, no. 5, pp. 2165-2170.Leung C.S., Haigh T.A., Mackay L.K., Rickinson A.B., Taylor G.S. Nuclear location of an endogenously expressed antigen, EBNA1, restricts access to macroautophagy and the range of CD4 epitope display. Proc. Natl Acad. Sci. USA, 2010, Vol. 107, no. 5, pp. 2165-2170.Liang C., Feng P., Ku B., Dotan I., Canaani D., Oh B.H., Jung J.U. Autophagic and tumour suppressor activity of a novel Beclin1-binding protein UVRAG. Nat. Cell Biol., 2006, Vol. 8, no. 7, pp. 688-699.Liang C., Feng P., Ku B., Dotan I., Canaani D., Oh B.H., Jung J.U. Autophagic and tumour suppressor activity of a novel Beclin1-binding protein UVRAG. Nat. Cell Biol., 2006, Vol. 8, no. 7, pp. 688-699.Ligeon L.A., Pena-Francesch M., Vanoaica L.D., Núñez N.G., Talwar D., Dick T.P., Münz C. Oxidation inhibits autophagy protein deconjugation from phagosomes to sustain MHC class II restricted antigen presentation. Nat. Commun., 2021, Vol. 12, no. 1, 1508. doi: 10.1038/s41467-021-21829-6.Ligeon L.A., Pena-Francesch M., Vanoaica L.D., Núñez N.G., Talwar D., Dick T.P., Münz C. Oxidation inhibits autophagy protein deconjugation from phagosomes to sustain MHC class II restricted antigen presentation. Nat. Commun., 2021, Vol. 12, no. 1, 1508. doi: 10.1038/s41467-021-21829-6.Ma J., Becker C., Lowell C.A., Underhill D.M. Dectin-1-triggered recruitment of light chain 3 protein to phagosomes facilitates major histocompatibility complex class II presentation of fungal-derived antigens. J. Biol. Chem., 2012, Vol. 287, no. 41, pp. 34149-34156.Ma J., Becker C., Lowell C.A., Underhill D.M. Dectin-1-triggered recruitment of light chain 3 protein to phagosomes facilitates major histocompatibility complex class II presentation of fungal-derived antigens. J. Biol. Chem., 2012, Vol. 287, no. 41, pp. 34149-34156.Martinez J., Almendinger J., Oberst A., Ness R., Dillon C.P., Fitzgerald P., Hengartner M.O., Green D.R. Microtubule-associated protein 1 light chain 3 alpha (LC3)-associated phagocytosis is required for the efficient clearance of dead cells. Proc. Natl Acad. Sci. USA, 2011, Vol. 108, no. 42, pp. 17396-17401.Martinez J., Almendinger J., Oberst A., Ness R., Dillon C.P., Fitzgerald P., Hengartner M.O., Green D.R. Microtubule-associated protein 1 light chain 3 alpha (LC3)-associated phagocytosis is required for the efficient clearance of dead cells. Proc. Natl Acad. Sci. USA, 2011, Vol. 108, no. 42, pp. 17396-17401.Martinez J., Cunha L.D., Park S., Yang M., Lu Q., Orchard R., Li Q.Z., Yan M., Janke L., Guy. C., Linkermann A., Virgin H.W., Green D.R. Noncanonical autophagy inhibits the autoinflammatory, lupus-like response to dying cells. Nature, 2016, Vol. 533, no. 7601, pp. 115-119.Martinez J., Cunha L.D., Park S., Yang M., Lu Q., Orchard R., Li Q.Z., Yan M., Janke L., Guy. C., Linkermann A., Virgin H.W., Green D.R. Noncanonical autophagy inhibits the autoinflammatory, lupus-like response to dying cells. Nature, 2016, Vol. 533, no. 7601, pp. 115-119.Martinez J., Malireddi R.K., Lu Q., Cunha L.D., Pelletier S., Gingras S., Orchard R., Guan J.L., Tan H., Peng J., Kanneganti T.D., Virgin H.W., Green D.R. Molecular characterization of LC3-associated phagocytosis reveals distinct roles for Rubicon, NOX2 and autophagy proteins. Nat. Cell Biol., 2015, Vol. 17, no. 7, pp. 893-906.Martinez J., Malireddi R.K., Lu Q., Cunha L.D., Pelletier S., Gingras S., Orchard R., Guan J.L., Tan H., Peng J., Kanneganti T.D., Virgin H.W., Green D.R. Molecular characterization of LC3-associated phagocytosis reveals distinct roles for Rubicon, NOX2 and autophagy proteins. Nat. Cell Biol., 2015, Vol. 17, no. 7, pp. 893-906.Martire B., Rondelli R., Soresina A., Pignata C., Broccoletti T., Finocchi A., Rossi P., Gattorno M., Rabusin M., Azzari C., Dellepiane R.M., Pietrogrande M.C., Trizzino A., di Bartolomeo P., Martino S., Carpino L., Cossu F., Locatelli F., Maccario R., Pierani P., Putti M.C., Stabile A., Notarangelo L.D., Ugazio A.G., Plebani A., de Mattia D. Clinical features, long-term follow-up and outcome of a large cohort of patients with Chronic Granulomatous Disease: an Italian multicenter study. Clin. Immunol., 2008, Vol. 126, no. 2, pp. 155-164.Martire B., Rondelli R., Soresina A., Pignata C., Broccoletti T., Finocchi A., Rossi P., Gattorno M., Rabusin M., Azzari C., Dellepiane R.M., Pietrogrande M.C., Trizzino A., di Bartolomeo P., Martino S., Carpino L., Cossu F., Locatelli F., Maccario R., Pierani P., Putti M.C., Stabile A., Notarangelo L.D., Ugazio A.G., Plebani A., de Mattia D. Clinical features, long-term follow-up and outcome of a large cohort of patients with Chronic Granulomatous Disease: an Italian multicenter study. Clin. Immunol., 2008, Vol. 126, no. 2, pp. 155-164.Masud S., Prajsnar T.K., Torraca V., Lamers G.E.M., Benning M., Vaart M.V.D., Meijer A.H. Macrophages target Salmonella by Lc3-associated phagocytosis in a systemic infection model. Autophagy, 2019, Vol. 15, no. 5, pp. 796-812.Masud S., Prajsnar T.K., Torraca V., Lamers G.E.M., Benning M., Vaart M.V.D., Meijer A.H. Macrophages target Salmonella by Lc3-associated phagocytosis in a systemic infection model. Autophagy, 2019, Vol. 15, no. 5, pp. 796-812.Mitchell G., Cheng M.I., Chen C., Nguyen B.N., Whiteley A.T., Kianian S., Cox J.S., Green D.R., McDonald K.L., Portnoy D.A. Listeria monocytogenes triggers noncanonical autophagy upon phagocytosis, but avoids subsequent growth-restricting xenophagy. Proc. Natl Acad. Sci. USA, 2018, Vol. 115, no. 2, pp. 210-217.Mitchell G., Cheng M.I., Chen C., Nguyen B.N., Whiteley A.T., Kianian S., Cox J.S., Green D.R., McDonald K.L., Portnoy D.A. Listeria monocytogenes triggers noncanonical autophagy upon phagocytosis, but avoids subsequent growth-restricting xenophagy. Proc. Natl Acad. Sci. USA, 2018, Vol. 115, no. 2, pp. 210-217.Mizushima N. Autophagy: process and function. Genes Dev., 2007, Vol. 21, no. 22, pp. 2861-2873.Mizushima N. Autophagy: process and function. Genes Dev., 2007, Vol. 21, no. 22, pp. 2861-2873.Mostowy S., Cossart P. Bacterial autophagy: restriction or promotion of bacterial replication? Trends Cell Biol., 2012, Vol. 22, no. 6, pp. 283-291.Mostowy S., Cossart P. Bacterial autophagy: restriction or promotion of bacterial replication? Trends Cell Biol., 2012, Vol. 22, no. 6, pp. 283-291.Münz C. Canonical and non-canonical functions of the autophagy machinery in MHC restricted antigen presentation. Front. Immunol., 2022, Vol. 13, 868888. doi: 10.3389/fimmu.2022.868888.Münz C. Canonical and non-canonical functions of the autophagy machinery in MHC restricted antigen presentation. Front. Immunol., 2022, Vol. 13, 868888. doi: 10.3389/fimmu.2022.868888.Niven J., Madelon N., Page N., Caruso A., Harlé G., Lemeille S., Seemayer C.A., Hugues S., Gannagé M. Macroautophagy in dendritic cells controls the homeostasis and stability of regulatory T cells. Cell Rep., 2019, Vol. 28, no. 1, pp. 21-29.Niven J., Madelon N., Page N., Caruso A., Harlé G., Lemeille S., Seemayer C.A., Hugues S., Gannagé M. Macroautophagy in dendritic cells controls the homeostasis and stability of regulatory T cells. Cell Rep., 2019, Vol. 28, no. 1, pp. 21-29.Noda T., Fujita N., Yoshimori T. The Ubi brothers reunited. Autophagy, 2008, Vol. 4, no. 4, pp. 540-541.Noda T., Fujita N., Yoshimori T. The Ubi brothers reunited. Autophagy, 2008, Vol. 4, no. 4, pp. 540-541.Paludan C., Schmid D., Landthaler M., Vockerodt M., Kube D., Tuschl T., Münz C. Endogenous MHC class II processing of a viral nuclear antigen after autophagy. Science, 2005, Vol. 307, no. 5709, pp. 593-596.Paludan C., Schmid D., Landthaler M., Vockerodt M., Kube D., Tuschl T., Münz C. Endogenous MHC class II processing of a viral nuclear antigen after autophagy. Science, 2005, Vol. 307, no. 5709, pp. 593-596.Paul S., Kashyap A.K., Jia W., He Y.W., Schaefer B.C. Selective autophagy of the adaptor protein Bcl10 modulates T cell receptor activation of NF-κB. Immunity, 2012, Vol. 36, no. 6, pp. 947-958.Paul S., Kashyap A.K., Jia W., He Y.W., Schaefer B.C. Selective autophagy of the adaptor protein Bcl10 modulates T cell receptor activation of NF-κB. Immunity, 2012, Vol. 36, no. 6, pp. 947-958.Pérez L., McLetchie S., Gardiner G.J., Deffit S.N., Zhou D., Blum J.S. LAMP-2C Inhibits MHC Class II Presentation of Cytoplasmic Antigens by Disrupting Chaperone-Mediated Autophagy. J. Immunol., 2016, Vol. 196, no. 6, pp. 2457-2465.Pérez L., McLetchie S., Gardiner G.J., Deffit S.N., Zhou D., Blum J.S. LAMP-2C Inhibits MHC Class II Presentation of Cytoplasmic Antigens by Disrupting Chaperone-Mediated Autophagy. J. Immunol., 2016, Vol. 196, no. 6, pp. 2457-2465.Quinn M.T., Gauss K.A. Structure and regulation of the neutrophil respiratory burst oxidase: comparison with nonphagocyte oxidases. J. Leukoc. Biol., 2004, Vol. 76, no. 4, pp. 760-781.Quinn M.T., Gauss K.A. Structure and regulation of the neutrophil respiratory burst oxidase: comparison with nonphagocyte oxidases. J. Leukoc. Biol., 2004, Vol. 76, no. 4, pp. 760-781.Riedel A., Nimmerjahn F., Burdach S., Behrends U., Bornkamm G.W., Mautner J. Endogenous presentation of a nuclear antigen on MHC class II by autophagy in the absence of CRM1-mediated nuclear export. Eur. J.Immunol., 2008, Vol. 38, no. 8, pp. 2090-2095.Riedel A., Nimmerjahn F., Burdach S., Behrends U., Bornkamm G.W., Mautner J. Endogenous presentation of a nuclear antigen on MHC class II by autophagy in the absence of CRM1-mediated nuclear export. Eur. J.Immunol., 2008, Vol. 38, no. 8, pp. 2090-2095.Rodriguez-Manzanet R., Sanjuan M.A., Wu H.Y., Quintana F.J., Xiao S., Anderson A.C., Weiner H.L., Green D.R., Kuchroo V.K. T and B cell hyperactivity and autoimmunity associated with niche-specific defects in apoptotic body clearance in TIM-4-deficient mice. Proc. Natl Acad. Sci. USA, 2010, Vol. 107, no. 19, pp. 8706-8711.Rodriguez-Manzanet R., Sanjuan M.A., Wu H.Y., Quintana F.J., Xiao S., Anderson A.C., Weiner H.L., Green D.R., Kuchroo V.K. T and B cell hyperactivity and autoimmunity associated with niche-specific defects in apoptotic body clearance in TIM-4-deficient mice. Proc. Natl Acad. Sci. USA, 2010, Vol. 107, no. 19, pp. 8706-8711.Romao S., Gasser N., Becker A.C., Guhl B., Bajagic M., Vanoaica D., Ziegler U., Roesler J., Dengjel J., Reichenbach J., Münz C. Autophagy proteins stabilize pathogen-containing phagosomes for prolonged MHC II antigen processing. J. Cell Biol., 2013, Vol. 203, no. 5, pp. 757-766.Romao S., Gasser N., Becker A.C., Guhl B., Bajagic M., Vanoaica D., Ziegler U., Roesler J., Dengjel J., Reichenbach J., Münz C. Autophagy proteins stabilize pathogen-containing phagosomes for prolonged MHC II antigen processing. J. Cell Biol., 2013, Vol. 203, no. 5, pp. 757-766.Sabatini D.M. mTOR and cancer: insights into a complex relationship. Nat. Rev. Cancer, 2006, Vol. 6, no. 9, pp. 729-734.Sabatini D.M. mTOR and cancer: insights into a complex relationship. Nat. Rev. Cancer, 2006, Vol. 6, no. 9, pp. 729-734.Sanjuan M.A., Dillon C.P., Tait S.W., Moshiach S., Dorsey F., Connell S., Komatsu M., Tanaka K., Cleveland J.L., Withoff S., Green D.R. Toll-like receptor signalling in macrophages links the autophagy pathway to phagocytosis. Nature, 2007, Vol. 450, no. 7173, pp. 1253-1257.Sanjuan M.A., Dillon C.P., Tait S.W., Moshiach S., Dorsey F., Connell S., Komatsu M., Tanaka K., Cleveland J.L., Withoff S., Green D.R. Toll-like receptor signalling in macrophages links the autophagy pathway to phagocytosis. Nature, 2007, Vol. 450, no. 7173, pp. 1253-1257.Sanjuan M.A., Green D.R. Eating for good health: linking autophagy and phagocytosis in host defense. Autophagy, 2008, Vol. 4, no. 5, pp. 607-611.Sanjuan M.A., Green D.R. Eating for good health: linking autophagy and phagocytosis in host defense. Autophagy, 2008, Vol. 4, no. 5, pp. 607-611.Sanjuan M.A., Milastra S., Green D.R. Toll-like receptor signaling in the lysosomal pathways. Immunol. Rev., 2009, Vol. 227, no. 1, pp. 203-220.Sanjuan M.A., Milastra S., Green D.R. Toll-like receptor signaling in the lysosomal pathways. Immunol. Rev., 2009, Vol. 227, no. 1, pp. 203-220.Satyavarapu E.M., Nath S., Mandal C. Desialylation of Atg5 by sialidase (Neu2) enhances autophagosome formation to induce anchorage-dependent cell death in ovarian cancer cells. Cell Death Discov., 2021, Vol. 7, no. 1, 26. doi: 10.1038/s41420-020-00391-y.Satyavarapu E.M., Nath S., Mandal C. Desialylation of Atg5 by sialidase (Neu2) enhances autophagosome formation to induce anchorage-dependent cell death in ovarian cancer cells. Cell Death Discov., 2021, Vol. 7, no. 1, 26. doi: 10.1038/s41420-020-00391-y.Shintani T., Klionsky D.J. Autophagy in health and disease: a double-edged sword. Science, 2004, Vol. 306, no. 5698, pp. 990-995.Shintani T., Klionsky D.J. Autophagy in health and disease: a double-edged sword. Science, 2004, Vol. 306, no. 5698, pp. 990-995.Song Z., An L., Ye Y., Wu J., Zou Y., He L., Zhu H. Essential role for UVRAG in autophagy and maintenance of cardiac function. Cardiovasc. Res., 2014, Vol. 101, no. 1, pp. 48-56.Song Z., An L., Ye Y., Wu J., Zou Y., He L., Zhu H. Essential role for UVRAG in autophagy and maintenance of cardiac function. Cardiovasc. Res., 2014, Vol. 101, no. 1, pp. 48-56.Sun Q., Zhang J., Fan W., Wong K.N., Ding X., Chen S., Zhong Q. The RUN domain of rubicon is important for hVps34 binding, lipid kinase inhibition, and autophagy suppression. J. Biol. Chem., 2011, Vol. 286, no. 1, pp. 185-191.Sun Q., Zhang J., Fan W., Wong K.N., Ding X., Chen S., Zhong Q. The RUN domain of rubicon is important for hVps34 binding, lipid kinase inhibition, and autophagy suppression. J. Biol. Chem., 2011, Vol. 286, no. 1, pp. 185-191.Suri A., Walters J.J., Rohrs H.W., Gross M.L., Unanue E.R. First signature of islet beta-cell-derived naturally processed peptides selected by diabetogenic class II MHC molecules. J. Immunol., 2008, Vol. 180, no. 6, pp. 3849- 3856.Suri A., Walters J.J., Rohrs H.W., Gross M.L., Unanue E.R. First signature of islet beta-cell-derived naturally processed peptides selected by diabetogenic class II MHC molecules. J. Immunol., 2008, Vol. 180, no. 6, pp. 3849- 3856.Tattoli I., Sorbara M.T., Yang C., Tooze S.A., Philpott D.J., Girardin S.E. Listeria phospholipases subvert host autophagic defenses by stalling pre-autophagosomal structures. EMBO J., 2013, Vol. 32, no. 23, pp. 3066-3078.Tattoli I., Sorbara M.T., Yang C., Tooze S.A., Philpott D.J., Girardin S.E. Listeria phospholipases subvert host autophagic defenses by stalling pre-autophagosomal structures. EMBO J., 2013, Vol. 32, no. 23, pp. 3066-3078.Wild P., McEwan D.G., Dikic I. The LC3 interactome at a glance. J. Cell Sci, 2014, Vol. 127, Pt 1, pp. 3-9.Wild P., McEwan D.G., Dikic I. The LC3 interactome at a glance. J. Cell Sci, 2014, Vol. 127, Pt 1, pp. 3-9.Wong S.W., Sil P., Martinez J. Rubicon: LC3-associated phagocytosis and beyond. FEBS J., 2018, Vol. 285, no. 8, pp. 1379-1388.Wong S.W., Sil P., Martinez J. Rubicon: LC3-associated phagocytosis and beyond. FEBS J., 2018, Vol. 285, no. 8, pp. 1379-1388.Wu D.J., Adamopoulos I.E. Autophagy and autoimmunity. Clin. Immunol., 2017, Vol. 176, pp. 55-62.Wu D.J., Adamopoulos I.E. Autophagy and autoimmunity. Clin. Immunol., 2017, Vol. 176, pp. 55-62.Yamamoto K., Venida A., Yano J., Biancur D.E., Kakiuchi M., Gupta S., Sohn A.S.W., Mukhopadhyay S., Lin E.Y., Parker S.J., Banh R.S., Paulo J.A., Wen K.W., Debnath J., Kim G.E., Mancias J.D., Fearon D.T., Perera R.M., Kimmelman A.C. Autophagy promotes immune evasion of pancreatic cancer by degrading MHC-I. Nature, 2020, Vol. 581, no. 7806, pp. 100-105.Yamamoto K., Venida A., Yano J., Biancur D.E., Kakiuchi M., Gupta S., Sohn A.S.W., Mukhopadhyay S., Lin E.Y., Parker S.J., Banh R.S., Paulo J.A., Wen K.W., Debnath J., Kim G.E., Mancias J.D., Fearon D.T., Perera R.M., Kimmelman A.C. Autophagy promotes immune evasion of pancreatic cancer by degrading MHC-I. Nature, 2020, Vol. 581, no. 7806, pp. 100-105.Yang C.S., Lee J.S., Rodgers M., Min C.K., Lee J.Y., Kim H.J., Lee K.H., Kim C.J., Oh B., Zandi E., Yue Z., Kramnik I., Liang C., Jung J.U. Autophagy protein Rubicon mediates phagocytic NADPH oxidase activation in response to microbial infection or TLR stimulation. Cell Host Microbe, 2012, Vol. 11, no. 3, pp. 264-276.Yang C.S., Lee J.S., Rodgers M., Min C.K., Lee J.Y., Kim H.J., Lee K.H., Kim C.J., Oh B., Zandi E., Yue Z., Kramnik I., Liang C., Jung J.U. Autophagy protein Rubicon mediates phagocytic NADPH oxidase activation in response to microbial infection or TLR stimulation. Cell Host Microbe, 2012, Vol. 11, no. 3, pp. 264-276.Yoshikawa Y., Ogawa M., Hain T., Yoshida M., Fukumatsu M., Kim M., Mimuro H., Nakagawa I., Yanagawa T., Ishii T., Kakizuka A., Sztul E., Chakraborty T., Sasakawa C. Listeria monocytogenes ActA-mediated escape from autophagic recognition. Nat. Cell Biol., 2009, Vol. 11, no. 10, pp. 1233-1240.Yoshikawa Y., Ogawa M., Hain T., Yoshida M., Fukumatsu M., Kim M., Mimuro H., Nakagawa I., Yanagawa T., Ishii T., Kakizuka A., Sztul E., Chakraborty T., Sasakawa C. Listeria monocytogenes ActA-mediated escape from autophagic recognition. Nat. Cell Biol., 2009, Vol. 11, no. 10, pp. 1233-1240.Yu L., Strandberg L., Lenardo M.J. The selectivity of autophagy and its role in cell death and survival. Autophagy, 2008, Vol. 4, no. 5, pp. 567-573.Yu L., Strandberg L., Lenardo M.J. The selectivity of autophagy and its role in cell death and survival. Autophagy, 2008, Vol. 4, no. 5, pp. 567-573.Zhao Z., Fux B., Goodwin M., Dunay I.R., Strong D., Miller B.C., Cadwell K., Delgado M.A., Ponpuak M., Green K.G., Schmidt R.E., Mizushima N., Deretic V., Sibley L.D., Virgin H.W. Autophagosome-independent essential function for the autophagy protein Atg5 in cellular immunity to intracellular pathogens. Cell Host Microbe, 2008, Vol. 4, no. 5, pp. 458-469.Zhao Z., Fux B., Goodwin M., Dunay I.R., Strong D., Miller B.C., Cadwell K., Delgado M.A., Ponpuak M., Green K.G., Schmidt R.E., Mizushima N., Deretic V., Sibley L.D., Virgin H.W. Autophagosome-independent essential function for the autophagy protein Atg5 in cellular immunity to intracellular pathogens. Cell Host Microbe, 2008, Vol. 4, no. 5, pp. 458-469.Zhong Y., Wang Q.J., Li X., Yan Y., Backer J.M., Chait B.T., Heintz N., Yue Z. Distinct regulation of autophagic activity by Atg14L and Rubicon associated with Beclin 1-phosphatidylinositol-3-kinase complex. Nat. Cell Biol., 2009, Vol. 11, no. 4, pp. 468-476.Zhong Y., Wang Q.J., Li X., Yan Y., Backer J.M., Chait B.T., Heintz N., Yue Z. Distinct regulation of autophagic activity by Atg14L and Rubicon associated with Beclin 1-phosphatidylinositol-3-kinase complex. Nat. Cell Biol., 2009, Vol. 11, no. 4, pp. 468-476.Zhong Z., Umemura A., Sanchez-Lopez E., Liang S., Shalapour S., Wong J., He F., Boassa D., Perkins G., Ali S.R., McGeough M.D., Ellisman M.H., Seki E., Gustafsson A.B., Hoffman H.M., Diaz-Meco M.T., Moscat J., Karin M. NF-κB Restricts Inflammasome Activation via Elimination of Damaged Mitochondria. Cell, 2016, Vol. 164, no. 5, pp. 896-910.Zhong Z., Umemura A., Sanchez-Lopez E., Liang S., Shalapour S., Wong J., He F., Boassa D., Perkins G., Ali S.R., McGeough M.D., Ellisman M.H., Seki E., Gustafsson A.B., Hoffman H.M., Diaz-Meco M.T., Moscat J., Karin M. NF-κB Restricts Inflammasome Activation via Elimination of Damaged Mitochondria. Cell, 2016, Vol. 164, no. 5, pp. 896-910.

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