{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T13:47:39Z","timestamp":1785764859882,"version":"3.56.0"},"reference-count":155,"publisher":"MDPI AG","issue":"15","license":[{"start":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T00:00:00Z","timestamp":1785542400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Stability Support Program of Shenzhen Universities and Colleges, China","award":["20231122174953002"],"award-info":[{"award-number":["20231122174953002"]}]},{"name":"Research Fund for High-end Talents of Shenzhen, China","award":["000700"],"award-info":[{"award-number":["000700"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Int. J. Mol. Sci.","IJMS"],"abstract":"<jats:p>Lung cancer ranks first in both incidence and mortality among all malignancies, and tumor microenvironment (TME)-induced CD8+ T cell exhaustion is a critical factor driving immune evasion and compromising the efficacy of immunotherapy. MicroRNAs (miRNAs), as key post-transcriptional regulators, shuttle between lung cancer cells and CD8+ T cells via extracellular vesicles (EVs), serving as critical communication hubs that reshape the TME. This review systematically synthesizes recent literature to summarize the regulatory patterns of miRNAs on functions of lung cancer cells and CD8+ T cells, and dissect the molecular mechanisms underlying miRNA-mediated bidirectional crosstalk between these two cell types. This review focuses on the dual-pronged immune evasion strategy employed by lung cancer cells to counteract CD8+ T cells. On the one hand, lung cancer cells aberrantly express endogenous miRNAs, such as miR-20a, miR-149-5p, and miR-326, to remodel their surface ligands and establish immune camouflage. On the other hand, they actively secrete EVs enriched in specific miRNAs, including miR-7108-3p, miR-651-5p, and miR-24-3p, which directly suppress CD8+ T cell function. Furthermore, lung cancer cells secrete additional miRNAs, notably miR-6794-5p, miR-708-5p, and miR-1234-3p, to reprogram other TME components, namely tumor-associated macrophages (TAMs), natural killer (NK) cells, and myeloid-derived suppressor cells (MDSCs). These reprogrammed cells, in turn, indirectly attenuate CD8+ T cells through a relay-like mechanism via immunosuppressive cytokines or surface checkpoint molecules produced by these cells. In addition, competing endogenous RNA (ceRNA) networks formed by long non-coding RNAs (lncRNAs) and circular RNAs (circRNAs) in lung cancer cells regulate miRNA activity at multiple levels, further impairing the immune effector functions of CD8+ T cells. Conversely, activated CD8+ T cells also secrete miRNA-containing EVs, which deliver these miRNAs to tumor cells, thereby inhibiting tumor progression. Elucidation of this miRNA-based bidirectional communication network will not only advance our understanding of immune evasion mechanisms in lung cancer but also provide novel insights into cell-free immunotherapeutic approaches based on CD8+ T cell-derived vesicles.<\/jats:p>","DOI":"10.3390\/ijms27156918","type":"journal-article","created":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T12:57:56Z","timestamp":1785761876000},"page":"6918","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Advances in miRNA-Mediated Bidirectional Crosstalk and Immune Evasion Mechanisms Between Lung Cancer Cells and CD8+ T Cells"],"prefix":"10.3390","volume":"27","author":[{"given":"Xinyi","family":"Zhou","sequence":"first","affiliation":[{"name":"School of Sports, Shenzhen University, Shenzhen 518060, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tao","family":"Pang","sequence":"additional","affiliation":[{"name":"School of Sports, Shenzhen University, Shenzhen 518060, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhe","family":"Ge","sequence":"additional","affiliation":[{"name":"School of Sports, Shenzhen University, Shenzhen 518060, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,1]]},"reference":[{"key":"ref_1","first-page":"229","article-title":"Global cancer statistics 2022: GLOBOCAN estimates of incidence and mortality worldwide for 36 cancers in 185 countries","volume":"74","author":"Bray","year":"2024","journal-title":"CA Cancer J. Clin."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Inamura, K. (2017). Lung Cancer: Understanding Its Molecular Pathology and the 2015 WHO Classification. Front. Oncol., 7.","DOI":"10.3389\/fonc.2017.00193"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"2339","DOI":"10.1200\/JCO.21.00174","article-title":"Five-Year Outcomes with Pembrolizumab Versus Chemotherapy for Metastatic Non-Small-Cell Lung Cancer with PD-L1 Tumor Proportion Score \u2265 50","volume":"39","author":"Reck","year":"2021","journal-title":"J. Clin. Oncol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"541","DOI":"10.1038\/s41591-018-0014-x","article-title":"Understanding the tumor immune microenvironment (TIME) for effective therapy","volume":"24","author":"Binnewies","year":"2018","journal-title":"Nat. Med."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1186\/s12935-026-04280-9","article-title":"MicroRNA-regulated signaling and tumor microenvironment modulation in lung cancer: Mechanistic insights and translational opportunities","volume":"26","author":"Shrestha","year":"2026","journal-title":"Cancer Cell Int."},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Zhao, Y.-L., Li, Y.-J., Wu, C.-X., Xie, N., Li, X.-Y., Cao, D.-C., Guo, Y., Liang, Y., Li, Y.-M., and Xue, J.-N. (2025). m6A-modified circZNF548 regulates exosomal miR-7108-3p to activate CD3+CD8+ T cells and suppress NSCLC growth by JMY. BMC Biol., 23.","DOI":"10.1186\/s12915-025-02355-z"},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Zhao, C., Cheng, L., Li, A., Wang, H., Li, X., and Xu, J. (2025). EGFR-Mutant Lung Adenocarcinoma Cell-Derived Exosomal miR-651-5p Induces CD8+ T Cell Apoptosis via Downregulating BCL2 Expression. Biomedicines, 13.","DOI":"10.3390\/biomedicines13020482"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"5330","DOI":"10.7150\/thno.58337","article-title":"CD45RO\u2212CD8+ T cell-derived exosomes restrict estrogen-driven endometrial cancer development via the ER\u03b2\/miR-765\/PLP2\/Notch axis","volume":"11","author":"Zhou","year":"2021","journal-title":"Theranostics"},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Su, D., Wu, G., Xiong, R., Sun, X., Xu, M., Mei, Y., and Wu, X. (2021). Tumor Immune Microenvironment Characteristics and Their Prognostic Value in Non-Small-Cell Lung Cancer. Front. Oncol., 11.","DOI":"10.3389\/fonc.2021.634059"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1186\/s12964-025-02069-3","article-title":"Intrinsic STING of CD8+ T cells regulates self-metabolic reprogramming and memory to exert anti-tumor effects","volume":"23","author":"Xu","year":"2025","journal-title":"Cell Commun. Signal."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"348","DOI":"10.1007\/s12672-025-02060-x","article-title":"Ki-67 expression in anti-programmed cell death protein-1 antibody-bound CD8+ T cells as a predictor of clinical benefit","volume":"16","author":"Tsurui","year":"2025","journal-title":"Discov. Oncol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1038\/s41577-021-00574-3","article-title":"CD8+ T cell differentiation and dysfunction in cancer","volume":"22","author":"Philip","year":"2022","journal-title":"Nat. Rev. Immunol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"389","DOI":"10.1146\/annurev.immunol.26.021607.090404","article-title":"Death by a thousand cuts: Granzyme pathways of programmed cell death","volume":"26","author":"Chowdhury","year":"2008","journal-title":"Annu. Rev. Immunol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1016\/j.coi.2021.03.016","article-title":"Modulation of the tumor micro-environment by CD8+ T cell-derived cytokines","volume":"69","author":"Hoekstra","year":"2021","journal-title":"Curr. Opin. Immunol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"e2100629","DOI":"10.1002\/advs.202100629","article-title":"Pulmonary Aerosol Delivery of Let-7b microRNA Confers a Striking Inhibitory Effect on Lung Carcinogenesis through Targeting the Tumor Immune Microenvironment","volume":"8","author":"Zhang","year":"2021","journal-title":"Adv. Sci."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"5352","DOI":"10.1172\/JCI76561","article-title":"Targeting miR-23a in CD8+ cytotoxic T lymphocytes prevents tumor-dependent immunosuppression","volume":"124","author":"Lin","year":"2014","journal-title":"J. Clin. Investig."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"134","DOI":"10.1007\/s12026-023-09423-y","article-title":"MiR-7 deficiency promotes Th1 polarization of CD4+T cells and enhances the antitumor effect in adoptive cell therapy for lung cancer","volume":"72","author":"Chang","year":"2024","journal-title":"Immunol. Res."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1483","DOI":"10.4049\/jimmunol.2100602","article-title":"Long Noncoding RNA lncNDEPD1 Regulates PD-1 Expression via miR-3619-5p in CD8+ T Cells","volume":"208","author":"Cheng","year":"2022","journal-title":"J. Immunol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1038\/cdd.2009.69","article-title":"miR-15a and miR-16-1 in cancer: Discovery, function and future perspectives","volume":"17","author":"Aqeilan","year":"2010","journal-title":"Cell Death Differ."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"RP87900","DOI":"10.7554\/eLife.87900","article-title":"The lncRNA Malat1 inhibits miR-15\/16 to enhance cytotoxic T cell activation and memory cell formation","volume":"12","author":"Wheeler","year":"2023","journal-title":"eLife"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"5585","DOI":"10.1038\/s41467-023-40959-7","article-title":"Let-7 enhances murine anti-tumor CD8 T cell responses by promoting memory and antagonizing terminal differentiation","volume":"14","author":"Wells","year":"2023","journal-title":"Nat. Commun."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"144","DOI":"10.1186\/s12943-021-01448-x","article-title":"Cancer cell-derived exosomal circUSP7 induces CD8+ T cell dysfunction and anti-PD1 resistance by regulating the miR-934\/SHP2 axis in NSCLC","volume":"20","author":"Chen","year":"2021","journal-title":"Mol. Cancer"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1111\/cas.16036","article-title":"Cancer cell-derived exosomal miR-20a-5p inhibits CD8+ T-cell function and confers anti-programmed cell death 1 therapy resistance in triple-negative breast cancer","volume":"115","author":"Li","year":"2024","journal-title":"Cancer Sci."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1186\/s12943-019-1024-0","article-title":"miR-301a promotes lung tumorigenesis by suppressing Runx3","volume":"18","author":"Li","year":"2019","journal-title":"Mol. Cancer"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1186\/1479-5876-11-71","article-title":"Dynamically expressed microRNA-15b modulates the activities of CD8+ T lymphocytes in mice with Lewis lung carcinoma","volume":"11","author":"Zhong","year":"2013","journal-title":"J. Transl. Med."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1017","DOI":"10.1534\/genetics.115.179176","article-title":"MicroRNAs and Their Targets Are Differentially Regulated in Adult and Neonatal Mouse CD8+ T Cells","volume":"201","author":"Wissink","year":"2015","journal-title":"Genetics"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1190","DOI":"10.1038\/s41565-024-01649-7","article-title":"Functionalized nanowires for miRNA-mediated therapeutic programming of na\u00efve T cells","volume":"19","author":"Kim","year":"2024","journal-title":"Nat. Nanotechnol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1007\/s00109-019-01865-y","article-title":"MicroRNA-181a regulates IFN-\u03b3 expression in effector CD8+ T cell differentiation","volume":"98","author":"Amado","year":"2020","journal-title":"J. Mol. Med."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1351","DOI":"10.1007\/s00262-020-02770-x","article-title":"Nicotine exhausts CD8+ T cells against tumor cells through increasing miR-629-5p to repress IL2RB-mediated granzyme B expression","volume":"70","author":"Cheng","year":"2021","journal-title":"Cancer Immunol. Immunother."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"3139","DOI":"10.1097\/CM9.0000000000003894","article-title":"Exosomal Ppar\u03b1 derived from cancer cells induces CD8+ T cell exhaustion in hepatocellular carcinoma through the miR-27b-3p \/TOX axis","volume":"138","author":"Zhong","year":"2025","journal-title":"Chin. Med. J."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"e010363","DOI":"10.1136\/jitc-2024-010363","article-title":"Memory-promoting function of miR-379-5p attenuates CD8+ T cell exhaustion by targeting immune checkpoints","volume":"13","author":"Lin","year":"2025","journal-title":"J. Immunother. Cancer"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"190061","DOI":"10.1098\/rsob.190061","article-title":"miR-149-3p reverses CD8+ T-cell exhaustion by reducing inhibitory receptors and promoting cytokine secretion in breast cancer cells","volume":"9","author":"Zhang","year":"2019","journal-title":"Open Biol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"84","DOI":"10.1186\/s12967-016-0841-9","article-title":"miR-29b and miR-198 overexpression in CD8+ T cells of renal cell carcinoma patients down-modulates JAK3 and MCL-1 leading to immune dysfunction","volume":"14","author":"Gigante","year":"2016","journal-title":"J. Transl. Med."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"143070","DOI":"10.1016\/j.ijbiomac.2025.143070","article-title":"Exosomes enriched with miR-31-3p from keratinocytes under oxidative stress promote vitiligo progression by destructing melanocytes and activating CD8+ T cells","volume":"310","author":"Ma","year":"2025","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"30923","DOI":"10.1038\/srep30923","article-title":"MicroRNA-491 regulates the proliferation and apoptosis of CD8+ T cells","volume":"6","author":"Yu","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"2584","DOI":"10.1016\/j.celrep.2017.08.060","article-title":"miR-150 Regulates Memory CD8 T Cell Differentiation via c-Myb","volume":"20","author":"Chen","year":"2017","journal-title":"Cell Rep."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"4212","DOI":"10.1038\/onc.2017.62","article-title":"MiR-21 is required for anti-tumor immune response in mice: An implication for its bi-directional roles","volume":"36","author":"He","year":"2017","journal-title":"Oncogene"},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Yin, L.-B., Song, C.-B., Zheng, J.-F., Fu, Y.-J., Qian, S., Jiang, Y.-J., Xu, J.-J., Ding, H.-B., Shang, H., and Zhang, Z.-N. (2018). Elevated Expression of miR-19b Enhances CD8+ T Cell Function by Targeting PTEN in HIV Infected Long Term Non-progressors with Sustained Viral Suppression. Front. Immunol., 9.","DOI":"10.3389\/fimmu.2018.03140"},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Xie, C., Wang, S., Zhang, H., Zhu, Y., Jiang, P., Shi, S., Si, Y., and Chen, J. (2023). Lnc-AIFM2-1 promotes HBV immune escape by acting as a ceRNA for miR-330-3p to regulate CD244 expression. Front. Immunol., 14.","DOI":"10.3389\/fimmu.2023.1121795"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"593","DOI":"10.1038\/ni.2576","article-title":"The microRNA miR-155 controls CD8+ T cell responses by regulating interferon signaling","volume":"14","author":"Gracias","year":"2013","journal-title":"Nat. Immunol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"2157","DOI":"10.1038\/s41467-019-09882-8","article-title":"miR-155 harnesses Phf19 to potentiate cancer immunotherapy through epigenetic reprogramming of CD8+ T cell fate","volume":"10","author":"Ji","year":"2019","journal-title":"Nat. Commun."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"861","DOI":"10.1038\/ni.2073","article-title":"The microRNA miR-29 controls innate and adaptive immune responses to intracellular bacterial infection by targeting interferon-\u03b3","volume":"12","author":"Ma","year":"2011","journal-title":"Nat. Immunol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"633","DOI":"10.1189\/jlb.3A0114-025R","article-title":"The TGF-\u03b2-inducible miR-23a cluster attenuates IFN-\u03b3 levels and antigen-specific cytotoxicity in human CD8+ T cells","volume":"96","author":"Chandran","year":"2014","journal-title":"J. Leukoc. Biol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"18608","DOI":"10.1073\/pnas.1317191110","article-title":"MicroRNA-directed program of cytotoxic CD8+ T-cell differentiation","volume":"110","author":"Trifari","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1601","DOI":"10.1210\/clinem\/dgad725","article-title":"Exosome-mediated Delivery of miR-519e-5p Promotes Malignant Tumor Phenotype and CD8+ T-Cell Exhaustion in Metastatic PTC","volume":"109","author":"Li","year":"2024","journal-title":"J. Clin. Endocrinol. Metab."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1684\/mrh.2020.0469","article-title":"miR-199a-5p targeted regulation of MAGT1 expression in the functional depletion of CD8+T cells in HBV infection","volume":"33","author":"Mo","year":"2020","journal-title":"Magnes. Res."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"405","DOI":"10.1038\/ni1575","article-title":"Lymphoproliferative disease and autoimmunity in mice with increased miR-17-92 expression in lymphocytes","volume":"9","author":"Xiao","year":"2008","journal-title":"Nat. Immunol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"549","DOI":"10.1016\/j.bbrc.2015.02.003","article-title":"Transgene-derived overexpression of miR-17-92 in CD8+ T-cells confers enhanced cytotoxic activity","volume":"458","author":"Kosaka","year":"2015","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"9965","DOI":"10.1073\/pnas.1207327109","article-title":"Temporal expression of microRNA cluster miR-17-92 regulates effector and memory CD8+ T-cell differentiation","volume":"109","author":"Wu","year":"2012","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Mitchell, M.I., and Loudig, O. (2026). Extracellular MicroRNAs: A Stable and Diverse Source of Transcriptional Control. Biomolecules, 16.","DOI":"10.3390\/biom16060787"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1038\/s41556-018-0250-9","article-title":"Specificities of secretion and uptake of exosomes and other extracellular vesicles for cell-to-cell communication","volume":"21","author":"Mathieu","year":"2019","journal-title":"Nat. Cell Biol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"2162","DOI":"10.1093\/hmg\/ddad052","article-title":"Lung adenocarcinoma cell-derived exosomes promote M2 macrophage polarization through transmission of miR-3153 to activate the JNK signaling pathway","volume":"32","author":"Xu","year":"2023","journal-title":"Hum. Mol. Genet."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"6354","DOI":"10.1038\/s41388-020-01406-9","article-title":"miR-21a in exosomes from Lewis lung carcinoma cells accelerates tumor growth through targeting PDCD4 to enhance expansion of myeloid-derived suppressor cells","volume":"39","author":"Zhang","year":"2020","journal-title":"Oncogene"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1164","DOI":"10.1038\/cr.2014.121","article-title":"Tumor-secreted miR-214 induces regulatory T cells: A major link between immune evasion and tumor growth","volume":"24","author":"Yin","year":"2014","journal-title":"Cell Res."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"3235","DOI":"10.4049\/jimmunol.168.7.3235","article-title":"TCR activation of human T cells induces the production of exosomes bearing the TCR\/CD3\/zeta complex","volume":"168","author":"Blanchard","year":"2002","journal-title":"J. Immunol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"118762","DOI":"10.1016\/j.ijpharm.2019.118762","article-title":"Effects of gefitinib treatment on cellular uptake of extracellular vesicles in EGFR-mutant non-small cell lung cancer cells","volume":"572","author":"Takenaka","year":"2019","journal-title":"Int. J. Pharm."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Hodakoski, C., Hopkins, B.D., Zhang, G., Su, T., Cheng, Z., Morris, R., Rhee, K.Y., Goncalves, M.D., and Cantley, L.C. (2019). Rac-Mediated Macropinocytosis of Extracellular Protein Promotes Glucose Independence in Non-Small Cell Lung Cancer. Cancers, 11.","DOI":"10.3390\/cancers11010037"},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Jin, H.Y., Oda, H., Chen, P., Yang, C., Zhou, X., Kang, S.G., Valentine, E., Kefauver, J.M., Liao, L., and Zhang, Y. (2017). Differential Sensitivity of Target Genes to Translational Repression by miR-17~92. PLoS Genet., 13.","DOI":"10.1371\/journal.pgen.1006623"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"1137","DOI":"10.1038\/s41590-018-0208-x","article-title":"The effect of cellular context on miR-155-mediated gene regulation in four major immune cell types","volume":"19","author":"Hsin","year":"2018","journal-title":"Nat. Immunol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1016\/j.cell.2007.03.008","article-title":"miR-181a is an intrinsic modulator of T cell sensitivity and selection","volume":"129","author":"Li","year":"2007","journal-title":"Cell"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"1067","DOI":"10.1016\/j.immuni.2017.11.008","article-title":"Roquin Suppresses the PI3K-mTOR Signaling Pathway to Inhibit T Helper Cell Differentiation and Conversion of Treg to Tfr Cells","volume":"47","author":"Essig","year":"2017","journal-title":"Immunity"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"458","DOI":"10.1080\/26895293.2022.2059578","article-title":"Silencing of long non-coding RNA linc01106 suppresses non-small cell lung cancer proliferation, migration and invasion by regulating microRNA-765","volume":"15","author":"Zhang","year":"2022","journal-title":"All Life"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"1033","DOI":"10.1093\/jleuko\/qiae117","article-title":"The antitumor effect of extracellular vesicles derived from cytokine-activated CD8+ T cells","volume":"116","author":"Zhang","year":"2024","journal-title":"J. Leukoc. Biol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"190","DOI":"10.3892\/or.2021.8141","article-title":"Overview of serpin B9 and its roles in cancer (Review)","volume":"46","author":"Wang","year":"2021","journal-title":"Oncol. Rep."},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Zhou, Q., Wei, S., Wang, H., Li, Y., Fan, S., Cao, Y., and Wang, C. (2023). T cell-derived exosomes in tumor immune modulation and immunotherapy. Front. Immunol., 14.","DOI":"10.3389\/fimmu.2023.1130033"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1038\/s41467-018-02865-1","article-title":"Activated CD8+ T cell extracellular vesicles prevent tumour progression by targeting of lesional mesenchymal cells","volume":"9","author":"Seo","year":"2018","journal-title":"Nat. Commun."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"374","DOI":"10.1016\/j.ccell.2023.02.016","article-title":"The evolving tumor microenvironment: From cancer initiation to metastatic outgrowth","volume":"41","author":"Joyce","year":"2023","journal-title":"Cancer Cell"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"236","DOI":"10.1002\/resp.70220","article-title":"T-Cell Immunity and Lung Cancer","volume":"31","author":"Yamada","year":"2026","journal-title":"Respirology"},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Saeed, A.F. (2025). Tumor-Associated Macrophages: Polarization, Immunoregulation, and Immunotherapy. Cells, 14.","DOI":"10.3390\/cells14100741"},{"key":"ref_70","doi-asserted-by":"crossref","unstructured":"Chen, K., Luo, L., Li, Y., and Yang, G. (2025). Reprogramming the immune microenvironment in lung cancer. Front. Immunol., 16.","DOI":"10.3389\/fimmu.2025.1684889"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"5151","DOI":"10.1172\/JCI128644","article-title":"Hedgehog signaling promotes tumor-associated macrophage polarization to suppress intratumoral CD8+ T cell recruitment","volume":"129","author":"Petty","year":"2019","journal-title":"J. Clin. Investig."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1007\/s00432-009-0634-0","article-title":"Population alterations of L-arginase- and inducible nitric oxide synthase-expressed CD11b+\/CD14\u2212\/CD15+\/CD33+ myeloid-derived suppressor cells and CD8+ T lymphocytes in patients with advanced-stage non-small cell lung cancer","volume":"136","author":"Liu","year":"2010","journal-title":"J. Cancer Res. Clin. Oncol."},{"key":"ref_73","doi-asserted-by":"crossref","unstructured":"Qin, D., Zhang, Y., Shu, P., Lei, Y., Li, X., and Wang, Y. (2024). Targeting tumor-infiltrating tregs for improved antitumor responses. Front. Immunol., 15.","DOI":"10.3389\/fimmu.2024.1325946"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"RP99010","DOI":"10.7554\/eLife.99010.3","article-title":"Syngeneic natural killer cell therapy activates dendritic and T cells in metastatic lungs and effectively treats low-burden metastases","volume":"13","author":"Huang","year":"2025","journal-title":"eLife"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"189568","DOI":"10.1016\/j.bbcan.2026.189568","article-title":"Cancer-associated fibroblasts in non-small cell lung cancer therapy: Challenges and opportunities","volume":"1881","author":"Deng","year":"2026","journal-title":"Biochim. Biophys. Acta Rev. Cancer"},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Yan, Q., Li, S., He, L., and Chen, N. (2024). Prognostic implications of tumor-infiltrating lymphocytes in non-small cell lung cancer: A systematic review and meta-analysis. Front. Immunol., 15.","DOI":"10.3389\/fimmu.2024.1476365"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"101479","DOI":"10.1016\/j.xcrm.2024.101479","article-title":"CXCL9\/10-engineered dendritic cells promote T cell activation and enhance immune checkpoint blockade for lung cancer","volume":"5","author":"Lim","year":"2024","journal-title":"Cell Rep. Med."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"e002045","DOI":"10.1136\/jitc-2020-002045","article-title":"CXCL9-expressing tumor-associated macrophages: New players in the fight against cancer","volume":"9","author":"Marcovecchio","year":"2021","journal-title":"J. Immunother. Cancer"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"1062","DOI":"10.1158\/2326-6066.CIR-17-0190","article-title":"Differential Expression of Homing Receptor Ligands on Tumor-Associated Vasculature that Control CD8 Effector T-cell Entry","volume":"5","author":"Woods","year":"2017","journal-title":"Cancer Immunol. Res."},{"key":"ref_80","first-page":"1973","article-title":"MicroRNA-181b regulates NF-\u03baB-mediated vascular inflammation","volume":"122","author":"Sun","year":"2012","journal-title":"J. Clin. Investig."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"1243","DOI":"10.1007\/s13258-022-01227-y","article-title":"The role of exosomal miR-181b in the crosstalk between NSCLC cells and tumor-associated macrophages","volume":"44","author":"Ma","year":"2022","journal-title":"Genes Genom."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"e001617","DOI":"10.1136\/jitc-2020-001617","article-title":"Wrinkle in the plan: miR-34a-5p impacts chemokine signaling by modulating CXCL10\/CXCL11\/CXCR3-axis in CD4+, CD8+ T cells, and M1 macrophages","volume":"8","author":"Hart","year":"2020","journal-title":"J. Immunother. Cancer"},{"key":"ref_83","doi-asserted-by":"crossref","unstructured":"Yi, C., Zhang, H., Guo, Q., Lu, L., Gao, C., Zhao, Y., Su, Y., and Lu, J. (2025). MiR-4664-3p as a potential diagnostic, prognostic, and immunotherapeutic biomarker in NSCLC: Modulation of tumor progression through CD8+ T cell regulation. Front. Oncol., 15.","DOI":"10.3389\/fonc.2025.1642999"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"1023","DOI":"10.1158\/2326-6066.CIR-14-0161","article-title":"The immunological synapse","volume":"2","author":"Dustin","year":"2014","journal-title":"Cancer Immunol. Res."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1186\/s13045-020-00848-8","article-title":"The role of cancer-derived microRNAs in cancer immune escape","volume":"13","author":"Yi","year":"2020","journal-title":"J. Hematol. Oncol."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"2","DOI":"10.1186\/s13578-018-0202-x","article-title":"Characterization and selective incorporation of small non-coding RNAs in non-small cell lung cancer extracellular vesicles","volume":"8","author":"Li","year":"2018","journal-title":"Cell Biosci."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"190","DOI":"10.1186\/s12964-024-01570-5","article-title":"Tumor-derived miR-6794-5p enhances cancer growth by promoting M2 macrophage polarization","volume":"22","author":"Choi","year":"2024","journal-title":"Cell Commun. Signal."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"123599","DOI":"10.1016\/j.lfs.2025.123599","article-title":"The crosstalk between lung adenocarcinoma cells and M2 macrophages promotes cancer cell development via the SFRS1\/miR-708-5p\/PD-L1 axis","volume":"371","author":"Xu","year":"2025","journal-title":"Life Sci."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"241","DOI":"10.1002\/hep.30607","article-title":"Endoplasmic Reticulum Stress Causes Liver Cancer Cells to Release Exosomal miR-23a-3p and Up-regulate Programmed Death Ligand 1 Expression in Macrophages","volume":"70","author":"Liu","year":"2019","journal-title":"Hepatology"},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"e2304222","DOI":"10.1002\/advs.202304222","article-title":"The Uridylyl Transferase TUT7-Mediated Accumulation of Exosomal miR-1246 Reprograms TAMs to Support CRC Progression","volume":"11","author":"Feng","year":"2024","journal-title":"Adv. Sci."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"e1062968","DOI":"10.1080\/2162402X.2015.1062968","article-title":"Hypoxic tumor-derived microvesicles negatively regulate NK cell function by a mechanism involving TGF-\u03b2 and miR23a transfer","volume":"5","author":"Berchem","year":"2016","journal-title":"Oncoimmunology"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"2651","DOI":"10.1002\/tox.23625","article-title":"MiR-1246 is responsible for lung cancer cells-derived exosomes-mediated promoting effects on lung cancer stemness via targeting TRIM17","volume":"37","author":"Liu","year":"2022","journal-title":"Environ. Toxicol."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"1307","DOI":"10.21037\/atm-22-5565","article-title":"Tumor cell-derived exosomal microRNA-146a promotes non-small cell lung cancer cell invasion and proliferation by inhibiting M1 macrophage polarization","volume":"10","author":"Yan","year":"2022","journal-title":"Ann. Transl. Med."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"568","DOI":"10.1016\/j.ymthe.2017.11.016","article-title":"Hypoxic Lung-Cancer-Derived Extracellular Vesicle MicroRNA-103a Increases the Oncogenic Effects of Macrophages by Targeting PTEN","volume":"26","author":"Hsu","year":"2018","journal-title":"Mol. Ther."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"e478","DOI":"10.1002\/ctm2.478","article-title":"Tumor-derived exosomal miR-19b-3p facilitates M2 macrophage polarization and exosomal LINC00273 secretion to promote lung adenocarcinoma metastasis via Hippo pathway","volume":"11","author":"Chen","year":"2021","journal-title":"Clin. Transl. Med."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"241","DOI":"10.1186\/s12957-022-02706-y","article-title":"Hypoxic lung cancer cell-derived exosomal miR-21 mediates macrophage M2 polarization and promotes cancer cell proliferation through targeting IRF1","volume":"20","author":"Jin","year":"2022","journal-title":"World J. Surg. Oncol."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"12639","DOI":"10.1002\/cam4.5954","article-title":"Hypoxic lung adenocarcinoma-derived exosomal miR-1290 induces M2 macrophage polarization by targeting SOCS3","volume":"12","author":"Gu","year":"2023","journal-title":"Cancer Med."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"4424221","DOI":"10.1155\/2022\/4424221","article-title":"The Exosomal miR-1246 of Laryngeal Squamous Cell Carcinoma Induces Polarization of M2 Type Macrophages and Promotes the Invasiveness of Laryngeal Squamous Cell Carcinoma","volume":"2022","author":"Wu","year":"2022","journal-title":"J. Oncol."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"110753","DOI":"10.1016\/j.abb.2026.110753","article-title":"Exosomal miR-93-5p modulates macrophage polarization to enhance prostate cancer progression","volume":"779","author":"Wang","year":"2026","journal-title":"Arch. Biochem. Biophys."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1186\/s13048-025-01616-3","article-title":"Tumour-derived exosomal miR-205 promotes ovarian cancer cell progression through M2 macrophage polarization via the PI3K\/Akt\/mTOR pathway","volume":"18","author":"He","year":"2025","journal-title":"J. Ovarian Res."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"e11940","DOI":"10.1002\/advs.202511940","article-title":"Red blood cell-derived exosomal miR-93-5p promotes lung cancer progression through PTEN suppression","volume":"13","author":"Li","year":"2026","journal-title":"Adv. Sci."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"42","DOI":"10.3816\/CLC.2009.n.006","article-title":"Exosomal microRNA: A diagnostic marker for lung cancer","volume":"10","author":"Rabinowits","year":"2009","journal-title":"Clin. Lung Cancer"},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"4681","DOI":"10.1038\/s41388-020-1322-4","article-title":"Cancer exosome-derived miR-9 and miR-181a promote the development of early-stage MDSCs via interfering with SOCS3 and PIAS3 respectively in breast cancer","volume":"39","author":"Jiang","year":"2020","journal-title":"Oncogene"},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"3449","DOI":"10.1016\/j.ymthe.2021.06.023","article-title":"Exosomal miR-1246 from glioma patient body fluids drives the differentiation and activation of myeloid-derived suppressor cells","volume":"29","author":"Qiu","year":"2021","journal-title":"Mol. Ther."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"e70221","DOI":"10.1096\/fj.202400951R","article-title":"FRK exerts oncogenic effects by targeting NQO2 via exosomal miR-9-3p to regulate mitochondrial function","volume":"38","author":"Huang","year":"2024","journal-title":"FASEB J."},{"key":"ref_106","doi-asserted-by":"crossref","unstructured":"Chang, W.-A., Tsai, M.-J., Hung, J.-Y., Wu, K.-L., Tsai, Y.-M., Huang, Y.-C., Chang, C.-Y., Tsai, P.-H., and Hsu, Y.-L. (2021). MiR-150-5p-containing extracellular vesicles are a new immunoregulator that favor the progression of lung cancer in hypoxic microenvironments by altering the phenotype of NK cells. Cancers, 13.","DOI":"10.3390\/cancers13246252"},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"3250","DOI":"10.1111\/ajt.15490","article-title":"Circulating exosomal miR-92b: Its role for cancer immunoediting and clinical value for prediction of posttransplant hepatocellular carcinoma recurrence","volume":"19","author":"Nakano","year":"2019","journal-title":"Am. J. Transplant."},{"key":"ref_108","doi-asserted-by":"crossref","unstructured":"Li, M., Shan, W., Hua, Y., Chao, F., Cui, Y., Lv, L., Dou, X., Bian, X., Zou, J., and Li, H. (2021). Exosomal miR-92b-3p promotes chemoresistance of small cell lung cancer through the PTEN\/AKT pathway. Front. Cell Dev. Biol., 9.","DOI":"10.3389\/fcell.2021.661602"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"34595","DOI":"10.1074\/jbc.M114.614503","article-title":"Activation of glioma cells generates immune tolerant NKT cells","volume":"289","author":"Tang","year":"2014","journal-title":"J. Biol. Chem."},{"key":"ref_110","doi-asserted-by":"crossref","unstructured":"Liu, F., Peng, W., Chen, J., Xu, Z., Jiang, R., Shao, Q., Zhao, N., and Qian, K. (2021). Exosomes Derived from Alveolar Epithelial Cells Promote Alveolar Macrophage Activation Mediated by miR-92a-3p in Sepsis-Induced Acute Lung Injury. Front. Cell. Infect. Microbiol., 11.","DOI":"10.3389\/fcimb.2021.646546"},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"2868","DOI":"10.1002\/ijc.28945","article-title":"Lung tumours reprogram pulmonary dendritic cell immunogenicity at the microRNA level","volume":"135","author":"Pyfferoen","year":"2014","journal-title":"Int. J. Cancer"},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"158","DOI":"10.1186\/s13046-024-03077-w","article-title":"Extracellular vesicles activated cancer-associated fibroblasts promote lung cancer metastasis through mitophagy and mtDNA transfer","volume":"43","author":"Zhou","year":"2024","journal-title":"J. Exp. Clin. Cancer Res."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"807","DOI":"10.1042\/CS20200039","article-title":"miR-210 transferred by lung cancer cell-derived exosomes may act as proangiogenic factor in cancer-associated fibroblasts by modulating JAK2\/STAT3 pathway","volume":"134","author":"Fan","year":"2020","journal-title":"Clin. Sci."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"242","DOI":"10.1186\/s13046-018-0911-3","article-title":"Melanoma cell-secreted exosomal miR-155-5p induce proangiogenic switch of cancer-associated fibroblasts via SOCS1\/JAK2\/STAT3 signaling pathway","volume":"37","author":"Zhou","year":"2018","journal-title":"J. Exp. Clin. Cancer Res."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"e12509","DOI":"10.1002\/jev2.12509","article-title":"Extracellular vesicles derived from melanoma cells induce carcinoma-associated fibroblasts via miR-92b-3p mediated downregulation of PTEN","volume":"13","author":"Windisch","year":"2024","journal-title":"J. Extracell. Vesicles"},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1166\/jbt.2022.2886","article-title":"Exosome miR-155-5p derived from lung cancer Hcc827 promotes macrophage activation and lung cancer progression","volume":"12","author":"Ma","year":"2022","journal-title":"J. Biomater. Tissue Eng."},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"D1670","DOI":"10.1093\/nar\/gkae973","article-title":"Generic Diagramming Platform (GDP): A comprehensive database of high-quality biomedical graphics","volume":"53","author":"Jiang","year":"2025","journal-title":"Nucleic Acids Res."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"582","DOI":"10.1101\/gad.349321.121","article-title":"The microRNA-183\/96\/182 cluster inhibits lung cancer progression and metastasis by inducing an interleukin-2-mediated antitumor CD8+ cytotoxic T-cell response","volume":"36","author":"Kundu","year":"2022","journal-title":"Genes Dev."},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"2960773","DOI":"10.1155\/2022\/2960773","article-title":"Mechanism of miR-760 Reversing Lung Cancer Immune Escape by Downregulating IDO1 and Eliminating Regulatory T Cells Based on Mathematical Biology","volume":"2022","author":"Ge","year":"2022","journal-title":"Comput. Math. Methods Med."},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"9785","DOI":"10.2147\/OTT.S265859","article-title":"MiRNA505\/NET1 Axis Acts as a CD8+ T-TIL Regulator in Non-Small Cell Lung Cancer","volume":"13","author":"Zhu","year":"2020","journal-title":"OncoTargets Ther."},{"key":"ref_121","doi-asserted-by":"crossref","unstructured":"Zhang, B., Liu, J., Mo, Y., Zhang, K., Huang, B., and Shang, D. (2024). CD8+ T cell exhaustion and its regulatory mechanisms in the tumor microenvironment: Key to the success of immunotherapy. Front. Immunol., 15.","DOI":"10.3389\/fimmu.2024.1476904"},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"689","DOI":"10.1158\/2326-6066.CIR-25-0431","article-title":"Upregulation of an Epithelial miRNA Is Associated with Immune Evasion in Progressive Bronchial Premalignant Lesions","volume":"14","author":"Ning","year":"2026","journal-title":"Cancer Immunol. Res."},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"4101","DOI":"10.1111\/cas.15921","article-title":"MicroRNA-326 negatively regulates CD155 expression in lung adenocarcinoma","volume":"114","author":"Nakanishi","year":"2023","journal-title":"Cancer Sci."},{"key":"ref_124","first-page":"173","article-title":"MiR-326 regulates cell proliferation and migration in lung cancer by targeting phox2a and is regulated by HOTAIR","volume":"6","author":"Wang","year":"2016","journal-title":"Am. J. Cancer Res."},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"3425","DOI":"10.1158\/1078-0432.CCR-15-2428","article-title":"Molecular Pathways: Targeting B7-H3 (CD276) for Human Cancer Immunotherapy","volume":"22","author":"Picarda","year":"2016","journal-title":"Clin. Cancer Res."},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"0590","DOI":"10.34133\/research.0590","article-title":"Altered Atlas of Exercise-Responsive MicroRNAs Revealing miR-29a-3p Attacks Armored and Cold Tumors and Boosts Anti-B7-H3 Therapy","volume":"8","author":"Mei","year":"2025","journal-title":"Research"},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"379","DOI":"10.3892\/ol.2022.13499","article-title":"miR-29a-3p inhibits the malignant characteristics of non-small cell lung cancer cells by reducing the activity of the Wnt\/\u03b2-catenin signaling pathway","volume":"24","author":"Zhang","year":"2022","journal-title":"Oncol. Lett."},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1038\/s41577-019-0224-6","article-title":"TIM3 comes of age as an inhibitory receptor","volume":"20","author":"Wolf","year":"2020","journal-title":"Nat. Rev. Immunol."},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1111\/cas.70232","article-title":"The LGALS9\/miR-491-5p Axis Regulates CD8+ T Cell Function and Inhibits the Progression of Gastric Cancer","volume":"117","author":"Huang","year":"2026","journal-title":"Cancer Sci."},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"622","DOI":"10.3892\/etm.2021.10054","article-title":"miR-491-5p inhibits the proliferation and migration of A549 cells by FOXP4","volume":"21","author":"Wu","year":"2021","journal-title":"Exp. Ther. Med."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"216186","DOI":"10.1016\/j.canlet.2023.216186","article-title":"Oncogenic miR-93-5p\/Gal-9 axis drives CD8 (+) T-cell inactivation and is a therapeutic target for hepatocellular carcinoma immunotherapy","volume":"564","author":"Dong","year":"2023","journal-title":"Cancer Lett."},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1016\/j.gene.2018.01.024","article-title":"MiR-93-5p up-regulation is involved in non-small cell lung cancer cells proliferation and migration and poor prognosis","volume":"647","author":"Yang","year":"2018","journal-title":"Gene"},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"148","DOI":"10.3892\/ol.2022.13269","article-title":"MicroRNA-20a promotes non-small cell lung cancer proliferation by upregulating PD-L1 by targeting PTEN","volume":"23","author":"Gong","year":"2022","journal-title":"Oncol. Lett."},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"3847","DOI":"10.1111\/jcmm.13657","article-title":"The miR-3127-5p\/p-STAT3 axis up-regulates PD-L1 inducing chemoresistance in non-small-cell lung cancer","volume":"22","author":"Tang","year":"2018","journal-title":"J. Cell. Mol. Med."},{"key":"ref_135","first-page":"888","article-title":"LINC00961 functions as an anti-oncogene in non-small cell lung carcinoma by regulation of miR-3127","volume":"14","author":"Liu","year":"2022","journal-title":"Am. J. Transl. Res."},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"5241","DOI":"10.1038\/ncomms6241","article-title":"Metastasis is regulated via microRNA-200\/ZEB1 axis control of tumour cell PD-L1 expression and intratumoral immunosuppression","volume":"5","author":"Chen","year":"2014","journal-title":"Nat. Commun."},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"djv303","DOI":"10.1093\/jnci\/djv303","article-title":"PDL1 Regulation by p53 via miR-34","volume":"108","author":"Cortez","year":"2016","journal-title":"J. Natl. Cancer Inst."},{"key":"ref_138","doi-asserted-by":"crossref","unstructured":"Zhang, Q., Pan, J., Xiong, D., Zheng, J., McPherson, K.N., Lee, S., Huang, M., Xu, Y., Chen, S.-H., and Wang, Y. (2023). Aerosolized miR-138-5p and miR-200c targets PD-L1 for lung cancer prevention. Front. Immunol., 14.","DOI":"10.3389\/fimmu.2023.1166951"},{"key":"ref_139","first-page":"5272","article-title":"Downregulation of miR-138-5p promotes non-small cell lung cancer progression by regulating CDK8","volume":"20","author":"Xing","year":"2019","journal-title":"Mol. Med. Rep."},{"key":"ref_140","doi-asserted-by":"crossref","first-page":"469","DOI":"10.1080\/10799893.2021.2019274","article-title":"XIST\/miR-34a-5p\/PDL1 axis regulated the development of lung cancer cells and the immune function of CD8+ T cells","volume":"42","author":"Li","year":"2022","journal-title":"J. Recept. Signal Transduct. Res."},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"1718","DOI":"10.1007\/s12672-025-03295-4","article-title":"LncRNA B4GALT 1-AS1\/miR-144-3p axis suppresses cytotoxicity of CD8+ T cells in lung adenocarcinoma by modulating PD-L1 expression","volume":"16","author":"Huang","year":"2025","journal-title":"Discov. Oncol."},{"key":"ref_142","first-page":"58","article-title":"circ_0001006 Promotes Immune Escape in Non-small Cell Lung Cancer by Regulating the miR-320a\/PD-L1 Axis","volume":"22","author":"Geng","year":"2025","journal-title":"Iran. J. Immunol."},{"key":"ref_143","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1038\/s41419-020-2230-9","article-title":"circRNA-002178 act as a ceRNA to promote PDL1\/PD1 expression in lung adenocarcinoma","volume":"11","author":"Wang","year":"2020","journal-title":"Cell Death Dis."},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"4094","DOI":"10.1093\/hmg\/ddac155","article-title":"Immunomodulatory functions of the circ_001678\/miRNA-326\/ZEB1 axis in non-small cell lung cancer via the regulation of PD-1\/PD-L1 pathway","volume":"31","author":"Tian","year":"2022","journal-title":"Hum. Mol. Genet."},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"109652","DOI":"10.1016\/j.cbi.2021.109652","article-title":"Thalidomide suppresses angiogenesis and immune evasion via lncRNA FGD5-AS1\/miR-454-3p\/ZEB1 axis-mediated VEGFA expression and PD-1\/PD-L1 checkpoint in NSCLC","volume":"349","author":"Xia","year":"2021","journal-title":"Chem. Biol. Interact."},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"2691","DOI":"10.1007\/s10528-023-10401-0","article-title":"CircDENND2D Inhibits PD-L1-Mediated Non-Small Cell Lung Cancer Metastasis and Immune Escape by Regulating miR-130b-3p\/STK11 Axis","volume":"61","author":"Chen","year":"2023","journal-title":"Biochem. Genet."},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"115041","DOI":"10.1016\/j.isci.2026.115041","article-title":"Immunometabolism in lung cancer\u2014The link between metabolism and immune response","volume":"29","author":"Eivazzadeh","year":"2026","journal-title":"iScience"},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"155613","DOI":"10.1016\/j.prp.2024.155613","article-title":"Exploring hypoxia-induced ncRNAs as biomarkers and therapeutic targets in lung cancer","volume":"263","author":"Thangavelu","year":"2024","journal-title":"Pathol. Res. Pract."},{"key":"ref_149","doi-asserted-by":"crossref","unstructured":"Wang, Z., Guo, H., Song, Y., Wang, A., Yan, Y., Ma, L., and Liu, B. (2025). Lung cancer tumor immune microenvironment: Analyzing immune escape mechanisms and exploring emerging therapeutic targets. Front. Immunol., 16.","DOI":"10.3389\/fimmu.2025.1597686"},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"693","DOI":"10.4103\/0973-1482.146130","article-title":"Increased expression of microRNA-301a in nonsmall-cell lung cancer and its clinical significance","volume":"12","author":"Shi","year":"2016","journal-title":"J. Cancer Res. Ther."},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"347","DOI":"10.3233\/CBM-201685","article-title":"MicroRNA-629 promotes the tumorigenesis of non-small-cell lung cancer by targeting FOXO1 and activating PI3K\/AKT pathway","volume":"29","author":"Zhu","year":"2020","journal-title":"Cancer Biomark."},{"key":"ref_152","doi-asserted-by":"crossref","unstructured":"Jusufovi\u0107, E., Rijavec, M., Keser, D., Koro\u0161ec, P., Sodja, E., Iljazovi\u0107, E., Radojevi\u0107, Z., and Ko\u0161nik, M. (2012). let-7b and miR-126 are down-regulated in tumor tissue and correlate with microvessel density and survival outcomes in non\u2013small\u2013cell lung cancer. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0045577"},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"4499","DOI":"10.3892\/mmr.2020.11553","article-title":"miR-379-5p inhibits cell proliferation and promotes cell apoptosis in non-small cell lung cancer by targeting \u03b2-arrestin-1","volume":"22","author":"Jiang","year":"2020","journal-title":"Mol. Med. Rep."},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"165","DOI":"10.17305\/bb.2024.11163","article-title":"miR-149-3p targeting TMPRSS4 regulates the sensitivity to cisplatin to inhibit the progression of lung cancer","volume":"25","author":"Qin","year":"2024","journal-title":"Biomol. Biomed."},{"key":"ref_155","first-page":"4113","article-title":"Antitumor effect of miR-27b-3p on lung cancer cells via targeting Fzd7","volume":"21","author":"Sun","year":"2017","journal-title":"Eur. Rev. Med. Pharmacol. Sci."}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/15\/6918\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T13:05:47Z","timestamp":1785762347000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/15\/6918"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,1]]},"references-count":155,"journal-issue":{"issue":"15","published-online":{"date-parts":[[2026,8]]}},"alternative-id":["ijms27156918"],"URL":"https:\/\/doi.org\/10.3390\/ijms27156918","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,1]]}}}