{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T11:18:55Z","timestamp":1788261535974,"version":"build-2803163510"},"reference-count":159,"publisher":"MDPI AG","issue":"5","license":[{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Non-Coding RNA","ncRNA"],"abstract":"<jats:p>MicroRNAs (miRNAs) are small non-coding RNAs that have emerged as critical regulators of gene expression and key cellular processes in cancer, including tumor cell proliferation, survival, metastasis, and both innate and adaptive immune responses within the tumor microenvironment. Through their ability to modulate immune cell development, differentiation, polarization, antigen presentation, cytokine signaling and immune checkpoint expression, miRNAs play a vital role in shaping immune responses and represent attractive therapeutic candidates for remodeling the tumor immune microenvironment. This review provides a comprehensive overview of the mechanisms through which miRNAs regulate anti-tumor immunity and contribute to immune evasion with particular emphasis on their role in modulating major immune checkpoint pathways, including PD-1\/PD-L1, CTLA-4, LAG-3, TIM-3 and CD28-mediated signaling. We further examine the contribution of miRNAs to therapeutic resistance and discuss their potential integration with immune checkpoint blockade, chemotherapy, PARP inhibitors and targeted therapies to enhance antitumor efficacy and overcome treatment resistance. Recent advances in spatial transcriptomics and computational approaches have expanded our understanding of miRNA-mediated regulatory networks at single-cell and tissue levels and provide new insights into tumor\u2013immune interactions. In addition, we evaluate the current progress in the clinical development of miRNA-based therapeutics, including miRNA mimics and inhibitors and discuss the major challenges associated with efficient delivery, off-target effects, immunogenicity and patient heterogeneity. Collectively, the evidence highlights the growing potential of miRNAs as diagnostic biomarkers, therapeutic targets and therapeutic tools in cancer immunotherapy. Continued advances in RNA therapeutics, delivery technologies and multi-omics approaches are expected to accelerate the clinical translation of miRNA-based strategies for personalized cancer therapy.<\/jats:p>","DOI":"10.3390\/ncrna12050034","type":"journal-article","created":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T10:32:18Z","timestamp":1788258738000},"page":"34","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["microRNAs as Regulators of the Immune Response and Their Potential Therapeutic Applications in Cancer"],"prefix":"10.3390","volume":"12","author":[{"ORCID":"https:\/\/orcid.org\/0009-0002-6338-1341","authenticated-orcid":false,"given":"Ezgi","family":"Biltekin","sequence":"first","affiliation":[{"name":"Department of Nanomedicine, Houston Methodist Research Institute, Houston, TX 77030, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bulent","family":"Ozpolat","sequence":"additional","affiliation":[{"name":"Department of Nanomedicine, Houston Methodist Research Institute, Houston, TX 77030, USA"},{"name":"Stephenson School of Biomedical Engineering, Stephenson Cancer Center, The University of Oklahoma, Norman, OK 73019, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,1]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"100123","DOI":"10.1016\/j.mocell.2024.100123","article-title":"Nobel-Winning MicroRNA, the Micromaestro of Gene Silencing","volume":"47","author":"Kim","year":"2024","journal-title":"Mol. Cells"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"15004","DOI":"10.1038\/sigtrans.2015.4","article-title":"The Role of MicroRNAs in Human Cancer","volume":"1","author":"Peng","year":"2016","journal-title":"Signal Transduct. Target. Ther."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1016\/S0092-8674(04)00045-5","article-title":"MicroRNAs: Genomics, Biogenesis, Mechanism, and Function","volume":"116","author":"Bartel","year":"2004","journal-title":"Cell"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"816","DOI":"10.1038\/s41576-023-00611-y","article-title":"MicroRNAs in Action: Biogenesis, Function and Regulation","volume":"24","author":"Shang","year":"2023","journal-title":"Nat. Rev. Genet."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"5166","DOI":"10.1073\/pnas.0800121105","article-title":"MiR-15a and MiR-16-1 Cluster Functions in Human Leukemia","volume":"105","author":"Calin","year":"2008","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"15524","DOI":"10.1073\/pnas.242606799","article-title":"Frequent Deletions and Down-Regulation of Micro-RNA Genes MiR15 and MiR16 at 13q14 in Chronic Lymphocytic Leukemia","volume":"99","author":"Calin","year":"2002","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"345","DOI":"10.1038\/s41568-021-00347-z","article-title":"Systemic Immunity in Cancer","volume":"21","author":"Allen","year":"2021","journal-title":"Nat. Rev. Cancer"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1016\/S1074-7613(00)80089-8","article-title":"Development of Lupus-like Autoimmune Diseases by Disruption of the PD-1 Gene Encoding an ITIM Motif-Carrying Immunoreceptor","volume":"11","author":"Nishimura","year":"1999","journal-title":"Immunity"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"1734","DOI":"10.1126\/science.271.5256.1734","article-title":"Enhancement of Antitumor Immunity by CTLA-4 Blockade","volume":"271","author":"Leach","year":"1996","journal-title":"Science"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"302","DOI":"10.1016\/j.gendis.2018.10.003","article-title":"2018 Nobel Prize in Medicine Awarded to Cancer Immunotherapy: Immune Checkpoint Blockade\u2014A Personal Account","volume":"5","author":"Zang","year":"2018","journal-title":"Genes Dis."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"e714","DOI":"10.1002\/mco2.714","article-title":"The Interaction of Innate Immune and Adaptive Immune System","volume":"5","author":"Wang","year":"2024","journal-title":"MedComm"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1038\/s41577-022-00732-1","article-title":"Roles of Natural Killer Cells in Immunity to Cancer, and Applications to Immunotherapy","volume":"23","author":"Wolf","year":"2022","journal-title":"Nat. Rev. Immunol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"302","DOI":"10.1038\/s41392-024-02005-w","article-title":"Comprehensive Snapshots of Natural Killer Cells Functions, Signaling, Molecular Mechanisms and Clinical Utilization","volume":"9","author":"Chen","year":"2024","journal-title":"Signal Transduct. Target. Ther."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"173090","DOI":"10.1016\/j.ejphar.2020.173090","article-title":"Macrophage M1\/M2 Polarization","volume":"877","author":"Yunna","year":"2020","journal-title":"Eur. J. Pharmacol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1038\/s41571-024-00859-1","article-title":"Dendritic Cells as Orchestrators of Anticancer Immunity and Immunotherapy","volume":"21","author":"Wculek","year":"2024","journal-title":"Nat. Rev. Clin. Oncol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"587","DOI":"10.1038\/s41423-020-0465-0","article-title":"Antigen Presentation by Dendritic Cells and Their Instruction of CD4+ T Helper Cell Responses","volume":"17","author":"Hilligan","year":"2020","journal-title":"Cell. Mol. Immunol."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Nanbakhsh, A., and Malarkannan, S. (2021). The Role of MicroRNAs in NK Cell Development and Function. Cells, 10.","DOI":"10.3390\/cells10082020"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"775","DOI":"10.1038\/ncb1613","article-title":"MicroRNAs 17-5p-20a-106a Control Monocytopoiesis through AML1 Targeting and M-CSF Receptor Upregulation","volume":"9","author":"Fontana","year":"2007","journal-title":"Nat. Cell Biol."},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Self-Fordham, J.B., Naqvi, A.R., Uttamani, J.R., Kulkarni, V., and Nares, S. (2017). MicroRNA: Dynamic Regulators of Macrophage Polarization and Plasticity. Front. Immunol., 8.","DOI":"10.3389\/fimmu.2017.01062"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"350","DOI":"10.1016\/j.it.2013.02.003","article-title":"MicroRNA-Mediated Control of Macrophages and Its Implications for Cancer","volume":"34","author":"Squadrito","year":"2013","journal-title":"Trends Immunol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"2806","DOI":"10.4049\/jimmunol.1500949","article-title":"MicroRNA-15\/16 Antagonizes Myb to Control NK Cell Maturation","volume":"195","author":"Sullivan","year":"2015","journal-title":"J. Immunol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"2275","DOI":"10.1182\/blood-2011-02-335141","article-title":"Macrophage Development from HSCs Requires PU.1-Coordinated MicroRNA Expression","volume":"118","author":"Ghani","year":"2011","journal-title":"Blood"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"2892","DOI":"10.1161\/CIRCULATIONAHA.111.087817","article-title":"A Novel Regulator of Macrophage Activation: MiR-223 in Obesity-Associated Adipose Tissue Inflammation","volume":"125","author":"Zhuang","year":"2012","journal-title":"Circulation"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"4149","DOI":"10.1172\/JCI81656","article-title":"MicroRNA-223 Is a Crucial Mediator of PPAR\u03b3-Regulated Alternative Macrophage Activation","volume":"125","author":"Ying","year":"2015","journal-title":"J. Clin. Investig."},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Wang, Z., Brandt, S., Medeiros, A., Wang, S., Wu, H., Dent, A., and Serezani, C.H. (2015). MicroRNA 21 Is a Homeostatic Regulator of Macrophage Polarization and Prevents Prostaglandin E2-Mediated M2 Generation. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0115855"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"2661","DOI":"10.2147\/OTT.S197682","article-title":"Cell-Free MiR-17-5p as a Diagnostic Biomarker for Gastric Cancer Inhibits Dendritic Cell Maturation","volume":"12","author":"Cui","year":"2019","journal-title":"OncoTargets Ther."},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Javaid, N., and Choi, S. (2020). Toll-like Receptors from the Perspective of Cancer Treatment. Cancers, 12.","DOI":"10.3390\/cancers12020297"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"2500","DOI":"10.4049\/jimmunol.1601778","article-title":"Serum MiR-29a Is Upregulated in Acute Graft-versus-Host Disease and Activates Dendritic Cells through TLR Binding","volume":"198","author":"Ranganathan","year":"2017","journal-title":"J. Immunol."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Hong, Y., Wu, J., Zhao, J., Wang, H., Liu, Y., Chen, T., Kan, X., Tao, Q., Shen, X., and Yan, K. (2013). MiR-29b and MiR-29c Are Involved in Toll-Like Receptor Control of Glucocorticoid-Induced Apoptosis in Human Plasmacytoid Dendritic Cells. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0069926"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"E2110","DOI":"10.1073\/pnas.1209414109","article-title":"MicroRNAs Bind to Toll-like Receptors to Induce Prometastatic Inflammatory Response","volume":"109","author":"Fabbri","year":"2012","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"298","DOI":"10.1038\/s41568-021-00339-z","article-title":"Antigen Presentation in Cancer: Insights into Tumour Immunogenicity and Immune Evasion","volume":"21","author":"Jhunjhunwala","year":"2021","journal-title":"Nat. Rev. Cancer"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"e001111","DOI":"10.1136\/jitc-2020-001111","article-title":"Antigen Processing and Presentation in Cancer Immunotherapy","volume":"8","author":"Lee","year":"2020","journal-title":"J. Immunother. Cancer"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"610","DOI":"10.1016\/j.bbrc.2012.12.097","article-title":"MiR-9 Modulates the Expression of Interferon-Regulated Genes and MHC Class I Molecules in Human Nasopharyngeal Carcinoma Cells","volume":"431","author":"Gao","year":"2013","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"784","DOI":"10.1053\/j.gastro.2018.06.030","article-title":"MicroRNA 125a Regulates MHC-I Expression on Esophageal Adenocarcinoma Cells, Associated With Suppression of Antitumor Immune Response and Poor Outcomes of Patients","volume":"155","author":"Mari","year":"2018","journal-title":"Gastroenterology"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"e2120","DOI":"10.1038\/cddis.2016.28","article-title":"Proteomic Screening Identifies Calreticulin as a MiR-27a Direct Target Repressing MHC Class I Cell Surface Exposure in Colorectal Cancer","volume":"7","author":"Colangelo","year":"2016","journal-title":"Cell Death Dis."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"2052","DOI":"10.1016\/j.cell.2024.03.037","article-title":"Principles and Therapeutic Applications of Adaptive Immunity","volume":"187","author":"Chi","year":"2024","journal-title":"Cell"},{"key":"ref_37","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_38","doi-asserted-by":"crossref","first-page":"6431","DOI":"10.1038\/ncomms7431","article-title":"ERK Phosphorylation and MiR-181a Expression Modulate Activation of Human Memory TH17 Cells","volume":"6","author":"Mele","year":"2015","journal-title":"Nat. Commun."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1016\/j.immuni.2011.07.009","article-title":"MicroRNA-29 Regulates T-Box Transcription Factors and Interferon-\u03b3 Production in Helper T Cells","volume":"35","author":"Steiner","year":"2011","journal-title":"Immunity"},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Skinner, J.P.J., Keown, A.A., and Chong, M.M.W. (2014). The MiR-17\u223c92a Cluster of MicroRNAs Is Required for the Fitness of Foxp3+ Regulatory T Cells. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0088997"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"153145","DOI":"10.1016\/j.prp.2020.153145","article-title":"Higher Expression of MiR-150-5p Promotes Tumorigenesis by Suppressing LKB1 in Non-Small Cell Lung Cancer","volume":"216","author":"Wu","year":"2020","journal-title":"Pathol. Res. Pract."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"16399","DOI":"10.1038\/srep16399","article-title":"MiR-150 Regulates Differentiation and Cytolytic Effector Function in CD8+ T Cells","volume":"5","author":"Smith","year":"2015","journal-title":"Sci. Rep."},{"key":"ref_43","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_44","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_45","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_46","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1038\/ni.3313","article-title":"Cancer Mediates Effector T Cell Dysfunction by Targeting MicroRNAs and EZH2 via Glycolysis Restriction","volume":"17","author":"Zhao","year":"2015","journal-title":"Nat. Immunol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"10662","DOI":"10.1073\/pnas.1100994108","article-title":"B Regulatory Cells and the Tumor-Promoting Actions of TNF-\u03b1 during Squamous Carcinogenesis","volume":"108","author":"Schioppa","year":"2011","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Tan, R., Nie, M., and Long, W. (2022). The Role of B Cells in Cancer Development. Front. Oncol., 12.","DOI":"10.3389\/fonc.2022.958756"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1374","DOI":"10.1182\/blood-2009-05-220814","article-title":"Src Homology 2 Domain-Containing Inositol-5-Phosphatase and CCAAT Enhancer-Binding Protein \u03b2 Are Targeted by MiR-155 in B Cells of E\u03bc-MiR-155 Transgenic Mice","volume":"114","author":"Costinean","year":"2009","journal-title":"Blood"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"2408","DOI":"10.1182\/blood-2016-08-731166","article-title":"MiR-28 Regulates the Germinal Center Reaction and Blocks Tumor Growth in Preclinical Models of Non-Hodgkin Lymphoma","volume":"129","author":"Mur","year":"2017","journal-title":"Blood"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"43076","DOI":"10.18632\/oncotarget.9246","article-title":"Epithelial Ovarian Cancer-Secreted Exosomal MiR-222-3p Induces Polarization of Tumor-Associated Macrophages","volume":"7","author":"Ying","year":"2016","journal-title":"Oncotarget"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"151875","DOI":"10.1016\/j.imbio.2019.11.010","article-title":"Promising Approaches in Cancer Immunotherapy","volume":"225","author":"Taefehshokr","year":"2020","journal-title":"Immunobiology"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"660","DOI":"10.1038\/s41422-020-0343-4","article-title":"Immune Checkpoint Signaling and Cancer Immunotherapy","volume":"30","author":"He","year":"2020","journal-title":"Cell Res."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1560","DOI":"10.1038\/bjc.2013.117","article-title":"Exploiting CTLA-4, PD-1 and PD-L1 to Reactivate the Host Immune Response against Cancer","volume":"108","author":"Quezada","year":"2013","journal-title":"Br. J. Cancer"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"852","DOI":"10.1038\/nri3108","article-title":"The Emerging Role of CTLA4 as a Cell-Extrinsic Regulator of T Cell Responses","volume":"11","author":"Walker","year":"2011","journal-title":"Nat. Rev. Immunol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"541","DOI":"10.1016\/1074-7613(95)90125-6","article-title":"Loss of CTLA-4 Leads to Massive Lymphoproliferation and Fatal Multiorgan Tissue Destruction, Revealing a Critical Negative Regulatory Role of CTLA-4","volume":"3","author":"Tivol","year":"1995","journal-title":"Immunity"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"535","DOI":"10.1016\/S1074-7613(00)80480-X","article-title":"Intracellular Trafficking of CTLA-4 and Focal Localization towards Sites of TCR Engagement","volume":"4","author":"Linsley","year":"1996","journal-title":"Immunity"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"878","DOI":"10.1038\/s41577-024-01061-1","article-title":"CD28 Co-Stimulation: Novel Insights and Applications in Cancer Immunotherapy","volume":"24","author":"Lotze","year":"2024","journal-title":"Nat. Rev. Immunol."},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Vaddi, P.K., Osborne, D.G., Nicklawsky, A., Williams, N.K., Menon, D.R., Smith, D., Mayer, J., Reid, A., Domenico, J., and Nguyen, G.H. (2023). CTLA4 MRNA Is Downregulated by MiR-155 in Regulatory T Cells, and Reduced Blood CTLA4 Levels Are Associated with Poor Prognosis in Metastatic Melanoma Patients. Front. Immunol., 14.","DOI":"10.3389\/fimmu.2023.1173035"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"10662","DOI":"10.1038\/s41598-019-47234-0","article-title":"Low Expression of MiR-424-3p Is Highly Correlated with Clinical Failure in Prostate Cancer","volume":"9","author":"Richardsen","year":"2019","journal-title":"Sci. Rep."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1177\/0004563220901564","article-title":"Clinical Value of CTLA4-Associated MicroRNAs Combined with Inflammatory Factors in the Diagnosis of Non-Small Cell Lung Cancer","volume":"57","author":"Liu","year":"2020","journal-title":"Ann. Clin. Biochem."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"8891","DOI":"10.1073\/pnas.92.19.8891","article-title":"P56Lck and P59Fyn Regulate CD28 Binding to Phosphatidylinositol 3-Kinase, Growth Factor Receptor-Bound Protein GRB-2, and T Cell-Specific Protein-Tyrosine Kinase ITK: Implications for T-Cell Costimulation","volume":"92","author":"Raab","year":"1995","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"860","DOI":"10.1038\/s41573-020-0081-9","article-title":"Targeting Co-Stimulatory Molecules in Autoimmune Disease","volume":"19","author":"Edner","year":"2020","journal-title":"Nat. Rev. Drug Discov."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"385","DOI":"10.1016\/j.cell.2017.08.018","article-title":"Mitochondrial Priming by CD28","volume":"171","author":"Geltink","year":"2017","journal-title":"Cell"},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Tang, Q., Chen, Y., Li, X., Long, S., Shi, Y., Yu, Y., Wu, W., Han, L., and Wang, S. (2022). The Role of PD-1\/PD-L1 and Application of Immune-Checkpoint Inhibitors in Human Cancers. Front. Immunol., 13.","DOI":"10.3389\/fimmu.2022.964442"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"201","DOI":"10.1007\/978-981-15-3266-5_9","article-title":"Functions of Immune Checkpoint Molecules Beyond Immune Evasion","volume":"Volume 1248","author":"Zhang","year":"2020","journal-title":"Regulation of Cancer Immune Checkpoints"},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Chen, R.Y., Zhu, Y., Shen, Y.Y., Xu, Q.Y., Tang, H.Y., Cui, N.X., Jiang, L., Dai, X.M., Chen, W.Q., and Lin, Q. (2023). The Role of PD-1 Signaling in Health and Immune-Related Diseases. Front. Immunol., 14.","DOI":"10.3389\/fimmu.2023.1163633"},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Song, N., Li, P., Song, P., Li, Y., Zhou, S., Su, Q., Li, X., Yu, Y., Li, P., and Feng, M. (2020). MicroRNA-138-5p Suppresses Non-Small Cell Lung Cancer Cells by Targeting PD-L1\/PD-1 to Regulate Tumor Microenvironment. Front. Cell Dev. Biol., 8, Correction in Front. Cell Dev. Biol. 2020, 8, 746. https:\/\/doi.org\/10.3389\/fcell.2020.00746.","DOI":"10.3389\/fcell.2020.00746"},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Zhang, M., Gao, D., Shi, Y., Wang, Y., Joshi, R., Yu, Q., Liu, D., Alotaibi, F., Zhang, Y., and Wang, H. (2019). MiR-149-3p Reverses CD8+ T-Cell Exhaustion by Reducing Inhibitory Receptors and Promoting Cytokine Secretion in Breast Cancer Cells. Open Biol., 9.","DOI":"10.1098\/rsob.190061"},{"key":"ref_70","doi-asserted-by":"crossref","unstructured":"Parvez, A., Choudhary, F., Mudgal, P., Khan, R., Qureshi, K.A., Farooqi, H., and Aspatwar, A. (2023). PD-1 and PD-L1: Architects of Immune Symphony and Immunotherapy Breakthroughs in Cancer Treatment. Front. Immunol., 14.","DOI":"10.3389\/fimmu.2023.1296341"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"1325","DOI":"10.4049\/jimmunol.182.3.1325","article-title":"MicroRNA-513 Regulates B7-H1 Translation and Is Involved in IFN-\u03b3-Induced B7-H1 Expression in Cholangiocytes","volume":"182","author":"Gong","year":"2009","journal-title":"J. Immunol."},{"key":"ref_72","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_73","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1007\/s11033-022-08047-z","article-title":"P53 Downregulates PD-L1 Expression via MiR-34a to Inhibit the Growth of Triple-Negative Breast Cancer Cells: A Potential Clinical Immunotherapeutic Target","volume":"50","author":"Deng","year":"2023","journal-title":"Mol. Biol. Rep."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"102211","DOI":"10.1016\/j.genrep.2025.102211","article-title":"MicroRNA-34a Enhances Immune Response and Suppresses PI3K\/AKT Signaling through Targeting PD-L1 in Triple-Negative Breast Cancer","volume":"40","author":"Tavakolpour","year":"2025","journal-title":"Gene Rep."},{"key":"ref_75","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_76","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_77","doi-asserted-by":"crossref","first-page":"11406","DOI":"10.1038\/ncomms11406","article-title":"MiR-424(322) Reverses Chemoresistance via T-Cell Immune Response Activation by Blocking the PD-L1 Immune Checkpoint","volume":"7","author":"Xu","year":"2016","journal-title":"Nat. Commun."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"1268","DOI":"10.1007\/s12033-021-00379-8","article-title":"MicroRNA-4458 Regulates PD-L1 Expression to Enhance Anti-Tumor Immunity in NSCLC via Targeting STAT3","volume":"63","author":"Liu","year":"2021","journal-title":"Mol. Biotechnol."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"8239","DOI":"10.1038\/s41598-025-92537-0","article-title":"MicroRNA 320a-3p up-Regulation Reduces PD-L1 Expression in Gastric Cancer Cells: An Experimental and Bioinformatic Study","volume":"15","author":"Asghariazar","year":"2025","journal-title":"Sci. Rep."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1016\/j.ebiom.2019.02.034","article-title":"MiR-873\/PD-L1 Axis Regulates the Stemness of Breast Cancer Cells","volume":"41","author":"Gao","year":"2019","journal-title":"EBioMedicine"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"2522","DOI":"10.21037\/tlcr-2025-117","article-title":"MiR-200a Regulates PD-L1 and Predicts Response to Immune Checkpoint Inhibitors in Advanced Non-Small Cell Lung Cancer","volume":"14","author":"Kaneko","year":"2025","journal-title":"Transl. Lung Cancer Res."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"93","DOI":"10.4149\/neo_2019_190202N106","article-title":"Mir-34a-5p\/Pd-L1 Axis Regulates Cisplatin Chemoresistance of Ovarian Cancer Cells","volume":"67","author":"Zuo","year":"2020","journal-title":"Neoplasma"},{"key":"ref_83","doi-asserted-by":"crossref","unstructured":"Wei, S., Wang, K., Huang, X., Zhao, Z., and Zhao, Z. (2019). Lncrna Malat1 Contributes to Non-Small Cell Lung Cancer Progression via Modulating Mir-200a-3p\/Programmed Death-Ligand 1 Axis. Int. J. Immunopathol. Pharmacol., 33.","DOI":"10.1177\/2058738419859699"},{"key":"ref_84","doi-asserted-by":"crossref","unstructured":"Anastasiadou, E., Messina, E., Sanavia, T., Mundo, L., Farinella, F., Lazzi, S., Megiorni, F., Ceccarelli, S., Pontecorvi, P., and Marampon, F. (2021). Mir-200c-3p Contrasts Pd-L1 Induction by Combinatorial Therapies and Slows Proliferation of Epithelial Ovarian Cancer through Downregulation of \u03b2-Catenin and c-Myc. Cells, 10.","DOI":"10.3390\/cells10030519"},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"639","DOI":"10.1093\/neuonc\/nov292","article-title":"MiR-138 Exerts Anti-Glioma Efficacy by Targeting Immune Checkpoints","volume":"18","author":"Wei","year":"2016","journal-title":"Neuro-Oncol."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"122352","DOI":"10.1016\/j.ijpharm.2022.122352","article-title":"Nanobubbles Containing PD-L1 Ab and MiR-424 Mediated PD-L1 Blockade, and Its Expression Inhibition to Enable and Potentiate Hepatocellular Carcinoma Immunotherapy in Mice","volume":"629","author":"Liu","year":"2022","journal-title":"Int. J. Pharm."},{"key":"ref_87","doi-asserted-by":"crossref","unstructured":"Kalkusova, K., Taborska, P., Stakheev, D., and Smrz, D. (2022). The Role of MiR-155 in Antitumor Immunity. Cancers, 14.","DOI":"10.3390\/cancers14215414"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"2348","DOI":"10.1261\/rna.1034808","article-title":"MicroRNA MiR-21 Overexpression in Human Breast Cancer Is Associated with Advanced Clinical Stage, Lymph Node Metastasis and Patient Poor Prognosis","volume":"14","author":"Yan","year":"2008","journal-title":"RNA"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"3777","DOI":"10.7150\/thno.97755","article-title":"Targeted Delivery of Anti-MiRNA21 Sensitizes PD-L1high Tumor to Immunotherapy by Promoting Immunogenic Cell Death","volume":"14","author":"Kim","year":"2024","journal-title":"Theranostics"},{"key":"ref_90","doi-asserted-by":"crossref","unstructured":"Chocarro, L., Blanco, E., Zuazo, M., Arasanz, H., Bocanegra, A., Fern\u00e1ndez-Rubio, L., Morente, P., Fern\u00e1ndez-Hinojal, G., Echaide, M., and Garnica, M. (2021). Understanding Lag-3 Signaling. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22105282"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"101305","DOI":"10.1016\/j.smim.2019.101305","article-title":"Lymphocyte-Activation Gene 3 (LAG3): The next Immune Checkpoint Receptor","volume":"42","author":"Ruffo","year":"2019","journal-title":"Semin. Immunol."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1186\/s12957-022-02810-z","article-title":"High Co-Expression of Immune Checkpoint Receptors PD-1, CTLA-4, LAG-3, TIM-3, and TIGIT on Tumor-Infiltrating Lymphocytes in Early-Stage Breast Cancer","volume":"20","author":"Mollavelioglu","year":"2022","journal-title":"World J. Surg. Oncol."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"814","DOI":"10.1016\/j.jtho.2017.01.019","article-title":"LAG-3 Protein Expression in Non\u2013Small Cell Lung Cancer and Its Relationship with PD-1\/PD-L1 and Tumor-Infiltrating Lymphocytes","volume":"12","author":"He","year":"2017","journal-title":"J. Thorac. Oncol."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"2082","DOI":"10.1002\/ijc.30912","article-title":"MiR-15a\/16 Deficiency Enhances Anti-Tumor Immunity of Glioma-Infiltrating CD8+ T Cells through Targeting MTOR","volume":"141","author":"Yang","year":"2017","journal-title":"Int. J. Cancer"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"389","DOI":"10.1007\/s12079-020-00546-7","article-title":"Identification of Extracellular Vesicles-Transported MiRNAs in Erlotinib-Resistant Head and Neck Squamous Cell Carcinoma","volume":"14","author":"Zheng","year":"2020","journal-title":"J. Cell Commun. Signal."},{"key":"ref_96","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_97","doi-asserted-by":"crossref","first-page":"61425","DOI":"10.18632\/oncotarget.18251","article-title":"Expression of LAG-3 Defines Exhaustion of Intratumoral PD-1+ T Cells and Correlates with Poor Outcome in Follicular Lymphoma","volume":"8","author":"Yang","year":"2017","journal-title":"Oncotarget"},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"e2419888122","DOI":"10.1073\/pnas.2419888122","article-title":"Triple Checkpoint Blockade of PD-1, Tim-3, and Lag-3 Enhances Adoptive T Cell Immunotherapy in a Mouse Model of Ovarian Cancer","volume":"122","author":"Alencar","year":"2025","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_99","doi-asserted-by":"crossref","unstructured":"Tao, J., Shen, X., Qian, H., Ding, Q., and Wang, L. (2024). TIM Proteins and MicroRNAs: Distinct Impact and Promising Interactions on Transplantation Immunity. Front. Immunol., 15.","DOI":"10.3389\/fimmu.2024.1500228"},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"1482","DOI":"10.1016\/j.prp.2018.07.019","article-title":"Evaluation of the Effect of TIM-3 Suppression by MiR-498 and Its Effect on Apoptosis and Proliferation Rate of HL-60 Cell Line","volume":"214","author":"Moghaddam","year":"2018","journal-title":"Pathol.-Res. Pract."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"e1211219","DOI":"10.1080\/2162402X.2016.1211219","article-title":"Tim-3 Promotes Tumor-Promoting M2 Macrophage Polarization by Binding to STAT1 and Suppressing the STAT1-MiR-155 Signaling Axis","volume":"5","author":"Jiang","year":"2016","journal-title":"Oncoimmunology"},{"key":"ref_102","doi-asserted-by":"crossref","unstructured":"Ciesielska-Figlon, K., and Lisowska, K.A. (2024). The Role of the CD28 Family Receptors in T-Cell Immunomodulation. Int. J. Mol. Sci., 25.","DOI":"10.3390\/ijms25021274"},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"a002436","DOI":"10.1101\/cshperspect.a002436","article-title":"An Enigmatic Tail of CD28 Signaling","volume":"2","author":"Boomer","year":"2010","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"ref_104","doi-asserted-by":"crossref","unstructured":"Raychaudhuri, K., Rangu, R., Ma, A., Alvinez, N., Tran, A.D., Pallikkuth, S., McIntire, K.M., Garvey, J.A., Yi, J., and Samelson, L.E. (2024). CD28 Shapes T Cell Receptor Signaling by Regulating Lck Dynamics and ZAP70 Activation. Front. Immunol., 15.","DOI":"10.3389\/fimmu.2024.1503018"},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"935","DOI":"10.1016\/S1074-7613(01)00244-8","article-title":"CD28 as a Molecular Amplifier Extending TCR Ligation and Signaling Capabilities","volume":"15","author":"Michel","year":"2001","journal-title":"Immunity"},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"2241","DOI":"10.1073\/pnas.1315606111","article-title":"CD28 and CD3 Have Complementary Roles in T-Cell Traction Forces","volume":"111","author":"Bashour","year":"2014","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_107","doi-asserted-by":"crossref","unstructured":"Teteloshvili, N., Dekkema, G., Boots, A.M., Heeringa, P., Jellema, P., de Jong, D., Terpstra, M., Brouwer, E., Pawelec, G., and Kok, K. (2018). Involvement of MicroRNAs in the Aging-Related Decline of CD28 Expression by Human T Cells. Front. Immunol., 9.","DOI":"10.3389\/fimmu.2018.01400"},{"key":"ref_108","doi-asserted-by":"crossref","unstructured":"Rodr\u00edguez-Gal\u00e1n, A., Fern\u00e1ndez-Messina, L., and S\u00e1nchez-Madrid, F. (2018). Control of Immunoregulatory Molecules by miRNAs in T Cell Activation. Front. Immunol., 9.","DOI":"10.3389\/fimmu.2018.02148"},{"key":"ref_109","doi-asserted-by":"crossref","unstructured":"Garc\u00eda-Gim\u00e9nez, J.L., Saadi, W., Ortega, A.L., Lahoz, A., Suay, G., Carretero, J., Pereda, J., Fatmi, A., Pallard\u00f3, F.V., and Mena-Molla, S. (2024). MiRNAs Related to Immune Checkpoint Inhibitor Response: A Systematic Review. Int. J. Mol. Sci., 25.","DOI":"10.3390\/ijms25031737"},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"560","DOI":"10.1053\/j.gastro.2021.04.036","article-title":"Tumor-Secreted Extracellular Vesicles Regulate T-Cell Costimulation and Can Be Manipulated To Induce Tumor-Specific T-Cell Responses","volume":"161","author":"Zhao","year":"2021","journal-title":"Gastroenterology"},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"37626","DOI":"10.18632\/oncotarget.5285","article-title":"High Intensity Focused Ultrasound Enhances Anti-Tumor Immunity by Inhibiting the Negative Regulatory Effect of MiR-134 on CD86 in a Murine Melanoma Model","volume":"6","author":"Yuan","year":"2015","journal-title":"Oncotarget"},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"1287","DOI":"10.1007\/s11481-013-9498-9","article-title":"Identification of Novel MicroRNA Signatures Linked to Experimental Autoimmune Myasthenia Gravis Pathogenesis: Down-Regulated Mir-145 Promotes Pathogenetic Th17 Cell Response","volume":"8","author":"Wang","year":"2013","journal-title":"J. Neuroimmune Pharmacol."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1186\/s12943-019-0975-5","article-title":"Exosomes in the Tumor Microenvironment as Mediators of Cancer Therapy Resistance","volume":"18","author":"Li","year":"2019","journal-title":"Mol. Cancer"},{"key":"ref_114","doi-asserted-by":"crossref","unstructured":"Wang, S., Jin, S., Zhang, J., and Wang, X. (2025). Exosomal MiRNAs: Key Regulators of the Tumor Microenvironment and Cancer Stem Cells. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms26199323"},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"e009774","DOI":"10.1136\/jitc-2024-009774","article-title":"Immune Modulatory MicroRNAs in Tumors, Their Clinical Relevance in Diagnosis and Therapy","volume":"12","author":"Vaxevanis","year":"2024","journal-title":"J. Immunother. Cancer"},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1186\/s12951-019-0563-2","article-title":"Engineered Exosomes for Targeted Co-Delivery of MiR-21 Inhibitor and Chemotherapeutics to Reverse Drug Resistance in Colon Cancer","volume":"18","author":"Liang","year":"2020","journal-title":"J. Nanobiotechnology"},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"4203","DOI":"10.7314\/APJCP.2015.16.10.4203","article-title":"Macrophage-Secreted Exosomes Delivering MiRNA-21 Inhibitor Can Regulate BGC-823 Cell Proliferation","volume":"16","author":"Wang","year":"2015","journal-title":"Asian Pac. J. Cancer Prev."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"619","DOI":"10.1186\/s12951-025-03697-0","article-title":"Engineered Exosomes: A Promising Approach for Overcoming Challenges in Pancreatic Cancer Therapy","volume":"23","author":"Sha","year":"2025","journal-title":"J. Nanobiotechnology"},{"key":"ref_119","doi-asserted-by":"crossref","unstructured":"Biltekin, E., Dilmac, S., Kahraman, N., Gul, O.A., Akay, Y.M., Wang, Z., Akay, M., and Ozpolat, B. (2026). Tumor-Suppressive MicroRNA Therapy Inhibits Growth of Glioblastoma Multiforme Xenografts. Cancers, 18.","DOI":"10.3390\/cancers18091479"},{"key":"ref_120","doi-asserted-by":"crossref","unstructured":"Mendieta, M., Bandegi, M., Biltekin, E., Akay, Y.M., Ozpolat, B., and Akay, M. (2025). MiR 329\/449 Suppresses Cell Proliferation, Migration and Synergistically Sensitizes GBM to TMZ by Inhibiting Src\/FAK, NF-KB, and Cyclin D1 Activity. Int. J. Mol. Sci., 26.","DOI":"10.20944\/preprints202504.1687.v1"},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"146","DOI":"10.1109\/OJEMB.2026.3678868","article-title":"MicroRNA-873 Suppresses Viability and Invasion of Colorectal Cancer through KRAS\/MAPK Signaling and Sensitizes Tumor Spheroids to 5-Fluorouracil in a 3D Microwell Model","volume":"7","author":"Bandegi","year":"2026","journal-title":"IEEE Open J. Eng. Med. Biol."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"126","DOI":"10.1186\/s13046-025-03391-x","article-title":"MicroRNA in Cancer Therapy: Breakthroughs and Challenges in Early Clinical Applications","volume":"44","author":"Tagliaferri","year":"2025","journal-title":"J. Exp. Clin. Cancer Res."},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"62","DOI":"10.1016\/j.jconrel.2022.02.031","article-title":"PDL1-Binding Peptide\/Anti-MiRNA21 Conjugate as a Therapeutic Modality for PD-L1high Tumors and TAMs","volume":"345","author":"Kim","year":"2022","journal-title":"J. Control. Release"},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"108","DOI":"10.1186\/s13048-021-00856-3","article-title":"GATA1-Induced Upregulation of LINC01503 Promotes Carboplatin Resistance in Ovarian Carcinoma by Upregulating PD-L1 via Sponging MiR-766-5p","volume":"14","author":"Li","year":"2021","journal-title":"J. Ovarian Res."},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1111\/cei.13406","article-title":"Cisplatin-Mediated down-Regulation of MiR-145 Contributes to up-Regulation of PD-L1 via the c-Myc Transcription Factor in Cisplatin-Resistant Ovarian Carcinoma Cells","volume":"200","author":"Sheng","year":"2020","journal-title":"Clin. Exp. Immunol."},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"748","DOI":"10.1038\/s41419-021-04033-8","article-title":"MiR-526b-3p Inhibits Lung Cancer Cisplatin-Resistance and Metastasis by Inhibiting STAT3-Promoted PD-L1","volume":"12","author":"Chen","year":"2021","journal-title":"Cell Death Dis."},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"6988923","DOI":"10.1155\/2022\/6988923","article-title":"Inhibition of PARP Potentiates Immune Checkpoint Therapy through MiR-513\/PD-L1 Pathway in Hepatocellular Carcinoma","volume":"2022","author":"Sun","year":"2022","journal-title":"J. Oncol."},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"558","DOI":"10.1007\/s12026-021-09237-w","article-title":"Circular RNA Hsa_circ_0001598 Promotes Programmed Death-Ligand-1-Mediated Immune Escape and Trastuzumab Resistance via Sponging MiR-1184 in Breast Cancer Cells","volume":"69","author":"Huang","year":"2021","journal-title":"Immunol. Res."},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"1281","DOI":"10.2217\/epi-2021-0130","article-title":"LncRNA HCG18 Suppresses CD8+T Cells to Confer Resistance to Cetuximab in Colorectal Cancer via MiR-20b-5p\/PD-L1 Axis","volume":"13","author":"Xu","year":"2021","journal-title":"Epigenomics"},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1038\/nrd.2016.246","article-title":"MicroRNA Therapeutics: Towards a New Era for the Management of Cancer and Other Diseases","volume":"16","author":"Rupaimoole","year":"2017","journal-title":"Nat. Rev. Drug Discov."},{"key":"ref_131","doi-asserted-by":"crossref","unstructured":"Raue, R., Frank, A.C., Syed, S.N., and Br\u00fcne, B. (2021). Therapeutic Targeting of Micrornas in the Tumor Microenvironment. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22042210"},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"470","DOI":"10.18632\/oncotarget.28608","article-title":"INT-1B3, an LNP Formulated MiR-193a-3p Mimic, Promotes Anti-Tumor Immunity by Enhancing T Cell Mediated Immune Responses via Modulation of the Tumor Microenvironment and Induction of Immunogenic Cell Death","volume":"15","author":"Duurland","year":"2024","journal-title":"Oncotarget"},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"1898","DOI":"10.1111\/jcmm.12882","article-title":"MiR-16 Regulates Mouse Peritoneal Macrophage Polarization and Affects T-Cell Activation","volume":"20","author":"Jia","year":"2016","journal-title":"J. Cell. Mol. Med."},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"1079","DOI":"10.2217\/epi-2016-0035","article-title":"Clinical Development of TargomiRs, a MiRNA Mimic-Based Treatment for Patients with Recurrent Thoracic Cancer","volume":"8","author":"Reid","year":"2016","journal-title":"Epigenomics"},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1007\/s10637-016-0407-y","article-title":"Phase I Study of MRX34, a Liposomal MiR-34a Mimic, Administered Twice Weekly in Patients with Advanced Solid Tumors","volume":"35","author":"Beg","year":"2017","journal-title":"Investig. New Drugs"},{"key":"ref_136","doi-asserted-by":"crossref","unstructured":"Bartolucci, D., Pession, A., Hrelia, P., and Tonelli, R. (2022). Precision Anti-Cancer Medicines by Oligonucleotide Therapeutics in Clinical Research Targeting Undruggable Proteins and Non-Coding RNAs. Pharmaceutics, 14.","DOI":"10.3390\/pharmaceutics14071453"},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"428","DOI":"10.1111\/bjh.15547","article-title":"Cobomarsen, an Oligonucleotide Inhibitor of MiR-155, Co-Ordinately Regulates Multiple Survival Pathways to Reduce Cellular Proliferation and Survival in Cutaneous T-Cell Lymphoma","volume":"183","author":"Seto","year":"2018","journal-title":"Br. J. Haematol."},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"S379","DOI":"10.1016\/j.annonc.2024.08.331","article-title":"383P Clinical Experience with TTX-MC138: A First-in-Class Therapy against Metastatic Cancer","volume":"35","author":"Varkaris","year":"2024","journal-title":"Ann. Oncol."},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1016\/j.jconrel.2015.08.011","article-title":"Small Molecules Targeting MicroRNA for Cancer Therapy: Promises and Obstacles","volume":"219","author":"Wen","year":"2015","journal-title":"J. Control. Release"},{"key":"ref_140","doi-asserted-by":"crossref","first-page":"1111","DOI":"10.1038\/gt.2011.100","article-title":"Chemical Modification and Design of Anti-MiRNA Oligonucleotides","volume":"18","author":"Lennox","year":"2011","journal-title":"Gene Ther."},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"591","DOI":"10.18632\/oncotarget.28641","article-title":"Inhibition of MiR-10b Treats Metastatic Breast Cancer by Targeting Stem Cell-like Properties","volume":"15","author":"Halim","year":"2024","journal-title":"Oncotarget"},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"15330338251339256","DOI":"10.1177\/15330338251339256","article-title":"MiR-10b as a Clinical Marker and a Therapeutic Target for Metastatic Breast Cancer","volume":"24","author":"Halim","year":"2025","journal-title":"Technol. Cancer Res. Treat."},{"key":"ref_143","doi-asserted-by":"crossref","unstructured":"Chen, M., Medarova, Z., Rogers, L.R., and Moore, A. (2026). The Multifaceted Role of MicroRNA-10b (MiR-10b) in Glioblastoma: From Oncogenic Driver to Therapeutic Target. Cells, 15.","DOI":"10.3390\/cells15090784"},{"key":"ref_144","doi-asserted-by":"crossref","unstructured":"Kara, G., Arun, B., Calin, G.A., and Ozpolat, B. (2022). MiRacle of MicroRNA-Driven Cancer Nanotherapeutics. Cancers, 14.","DOI":"10.3390\/cancers14153818"},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1007\/s00253-025-13620-0","article-title":"MiRNA-Based Drugs: Challenges and Delivery Strategies","volume":"109","author":"Wang","year":"2025","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"e1903296","DOI":"10.1002\/smll.201903296","article-title":"PH-Responsive PEG-Shedding and Targeting Peptide-Modified Nanoparticles for Dual-Delivery of Irinotecan and MicroRNA to Enhance Tumor-Specific Therapy","volume":"15","author":"Juang","year":"2019","journal-title":"Small"},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1186\/s12951-019-0526-7","article-title":"Functional Exosome-Mediated Co-Delivery of Doxorubicin and Hydrophobically Modified MicroRNA 159 for Triple-Negative Breast Cancer Therapy","volume":"17","author":"Gong","year":"2019","journal-title":"J. Nanobiotechnology"},{"key":"ref_148","doi-asserted-by":"crossref","unstructured":"Waheed, I., Ali, A., Tabassum, H., Khatoon, N., Lai, W.-F., and Zhou, X. (2024). Lipid-Based Nanoparticles as Drug Delivery Carriers for Cancer Therapy. Front. Oncol., 14.","DOI":"10.3389\/fonc.2024.1296091"},{"key":"ref_149","doi-asserted-by":"crossref","unstructured":"Savenkova, D.A., Makarova, A.L.A., Shalik, I.K., and Yudkin, D.V. (2022). MiRNA Pathway Alteration in Response to Non-Coding RNA Delivery in Viral Vector-Based Gene Therapy. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms232314954"},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"zcab030","DOI":"10.1093\/narcan\/zcab030","article-title":"Ligand-Mediated Delivery of RNAi-Based Therapeutics for the Treatment of Oncological Diseases","volume":"3","author":"Abdelaal","year":"2021","journal-title":"NAR Cancer"},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"S1","DOI":"10.21037\/tlcr.2017.12.01","article-title":"An Innovative Mesothelioma Treatment Based on MiR-16 Mimic Loaded EGFR Targeted Minicells (TargomiRs)","volume":"7","author":"Viteri","year":"2018","journal-title":"Transl. Lung Cancer Res."},{"key":"ref_152","doi-asserted-by":"crossref","unstructured":"Kim, B.D., Chen, M., Mondal, S.K., Kenyon, E., Mallett, C.L., deCarvalho, A., Medarova, Z., and Moore, A. (2025). Nanotherapy Targeting MiR-10b Improves Survival in Orthotopic Glioblastoma Models. J. Funct. Biomater., 17.","DOI":"10.3390\/jfb17010015"},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"96","DOI":"10.1186\/s12929-022-00879-y","article-title":"Spatial Multi-Omics Analyses of the Tumor Immune Microenvironment","volume":"29","author":"Hsieh","year":"2022","journal-title":"J. Biomed. Sci."},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"1131","DOI":"10.1016\/j.tig.2025.06.005","article-title":"Spatially Resolved MicroRNA Expression in Tissues: Technologies, Challenges, and Opportunities","volume":"41","author":"Zou","year":"2025","journal-title":"Trends Genet."},{"key":"ref_155","doi-asserted-by":"crossref","unstructured":"Su, R., Du, Y., Tian, P., Ma, W., Hui, Y., and Yang, S. (2025). Single-Cell and Spatial Transcriptomics Reveal Correlation between RNA Methylation-Related MiRNA Risk Model and Immune Infiltration in Hepatocellular Carcinoma. Front. Oncol., 15.","DOI":"10.3389\/fonc.2025.1553239"},{"key":"ref_156","doi-asserted-by":"crossref","unstructured":"Yuan, J., Xu, P., Ye, Z., and Liu, W. (2025). STmiR: A Novel XGBoost-Based Framework for Spatially Resolved MiRNA Activity Prediction in Cancer Transcriptomics. PLoS ONE, 20.","DOI":"10.1101\/2025.03.18.644021"},{"key":"ref_157","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1038\/s42003-025-07454-9","article-title":"Inferring Single-Cell and Spatial MicroRNA Activity from Transcriptomics Data","volume":"8","author":"Herbst","year":"2025","journal-title":"Commun. Biol."},{"key":"ref_158","doi-asserted-by":"crossref","unstructured":"Seyhan, A.A. (2024). Trials and Tribulations of MicroRNA Therapeutics. Int. J. Mol. Sci., 25.","DOI":"10.3390\/ijms25031469"},{"key":"ref_159","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1186\/s40164-022-00277-y","article-title":"Heterogeneity of the Tumor Immune Microenvironment and Its Clinical Relevance","volume":"11","author":"Jia","year":"2022","journal-title":"Exp. Hematol. Oncol."}],"container-title":["Non-Coding RNA"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2311-553X\/12\/5\/34\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T10:54:15Z","timestamp":1788260055000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2311-553X\/12\/5\/34"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,1]]},"references-count":159,"journal-issue":{"issue":"5","published-online":{"date-parts":[[2026,10]]}},"alternative-id":["ncrna12050034"],"URL":"https:\/\/doi.org\/10.3390\/ncrna12050034","relation":{},"ISSN":["2311-553X"],"issn-type":[{"value":"2311-553X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,1]]}}}