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Review Open Access

MicroRNA regulation of macrophage polarization in lung cancer: Regulatory networks and therapeutic potential (Review)

  • Authors:
    • Zhao Yiming
    • Li Jie
  • View Affiliations / Copyright

    Affiliations: Clinical Medical College of Beijing University of Chinese Medicine, Beijing 100029, P.R. China
    Copyright: © Yiming et al. This is an open access article distributed under the terms of Creative Commons Attribution License.
  • Article Number: 121
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    Published online on: September 2, 2026
       https://doi.org/10.3892/ijo.2026.5934
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Abstract

Lung cancer is among the most prevalent and fatal cancers worldwide, with its progression heavily shaped by the tumor microenvironment (TME). As key immune cell populations within the TME of lung cancer, macrophages are involved in the entire process of tumor initiation and development. Macrophage polarization serves as a critical link between inflammation and tumorigenesis and is broadly divided into classically activated M1 and alternatively activated M2 phenotypes. These phenotypes exert tumor‑suppressive and tumor‑promoting effects, respectively, by secreting distinct cytokine profiles. MicroRNAs (miRNAs/miRs) are a class of endogenous small non‑coding RNAs that play broad roles in the malignant progression of lung cancer by regulating various cellular processes, including proliferation, resistance to apoptosis, and metabolic reprogramming. Emerging evidence suggests that miRNAs such as miR‑335‑5p, miR‑106a‑5p and miR‑99b regulate M1/M2 macrophage polarization by targeting key pathways in lung cancer. Through these regulatory events, miRNAs exert synergistic effects on multiple tumor hallmarks, including proliferation, invasion, migration, apoptosis, angiogenesis, cell cycle progression, stemness maintenance, and epithelial‑mesenchymal transition. Collectively, these events drive lung cancer progression and influence therapeutic efficacy. The present review synthesizes recent discoveries to explore the relationship between macrophage polarization and lung cancer, with a specific focus on miRNA‑mediated regulatory mechanisms in lung cancer therapy. Ultimately, the present review aimed to provide new insight and a reference point to inform future prevention and treatment strategies for lung cancer.
View Figures

Figure 1

Mechanisms of macrophage
polarization. The image presents the complete regulatory network
for macrophage polarization. Under the guidance of Th1 and Th2
cells and GM-CSF and other signals, macrophages can polarize into
the pro-inflammatory M1 type (induced by TLR ligands, expressing
CD80/CD86/iNOS, secreting IL-1β, TNF-α, etc., with anti-pathogen
and anti-tumor functions), as well as four M2 subtypes: M2a
(induced by IL-4/IL-13, involved in type 2 immunity and allergy),
M2b (induced by immune complexes, having both immunoregulatory and
pro-inflammatory effects), M2c (induced by glucocorticoids/IL-10,
mediating immunosuppression and tissue repair), and M2d (induced by
TLR ligands/adenosine, promoting angiogenesis). The figure
systematically reveals the functional diversity of macrophages in
inflammation, immune regulation and tissue remodeling. GM-CSF,
granulocyte-macrophage colony-stimulating factor; TLR, Toll-like
receptor; iNOS, inducible nitric oxide synthase; MHC-II, major
histocompatibility complex class II.

Figure 2

Mechanisms underlying the regulation
of lung cancer by macrophage polarization. The image illustrates
that macrophage polarization profoundly affects the occurrence and
development of lung cancer by regulating multiple key biological
processe,s such as the proliferation, invasion and migration,
apoptosis, angiogenesis, cell cycle progression, stem cell
maintenance and the epithelial-mesenchymal transition of lung
cancer cells.

Figure 3

Mechanisms of miRNA-mediated
regulation of macrophage polarization in lung cancer therapy. The
image illustrates that miR-99b, miR-let-7b-5p, as well as the
upregulation of miR-135a-5p, miR-770 and miR-613 expression can
induce M1 polarization and inhibit M2 polarization, thereby
synergistically inhibiting the proliferation, invasion and
migration of lung cancer cells, inhibiting epithelial-mesenchymal
transition, and promoting cell apoptosis, thus significantly
delaying the growth and metastasis of lung cancer. MHC-II, major
histocompatibility complex class II; Arg-1, arginase-1; MR, mannose
receptor.

Figure 4

Mechanisms by which miRNAs regulate
macrophage polarization to promote lung cancer progression. The
image illustrates shows that the upregulation of miR-106a-5p,
miR-21-5p, miR-3153 and miR-146a can inhibit M1 polarization,
promote M2 polarization, and promote the proliferation, invasion
and migration of lung cancer cells, induce EMT, and inhibit lung
cancer cell apoptosis, while promoting lung cancer angiogenesis. In
addition, downregulation of miR-335-5p, miR-4319, miR-103a,
miR-155, and miR-132-3p can inhibit M1 polarization, promote M2
polarization, and synergistically enhance the maintenance of lung
cancer cell stemness, cell cycle progression and EMT, ultimately
accelerating tumor growth and metastasis. EMT,
epithelial-mesenchymal transition; IGF2BP3, insulin-like growth
factor 2 mRNA-binding protein 3; VAPA, vesicle-associated membrane
protein-associated protein A; NECAB3, N-terminal EF-hand type
calcium-binding protein 3; PTEN, phosphatase and tensin homolog;
MINK1, misshapen-like kinase 1; TREM2, triggering receptor
expressed on myeloid cells 2; TRAF6, TNF receptor-associated factor
6; IRAK1, interleukin-1 receptor-associated kinase 1.
View References

1 

Ji Y, Zhang Y, Liu S, Li J, Jin Q, Wu J, Duan H, Liu X, Yang L and Huang Y: The epidemiological landscape of lung cancer: Current status, temporal trend and future projections based on the latest estimates from GLOBOCAN 2022. J Natl Cancer Cent. 5:278–286. 2025.PubMed/NCBI

2 

Passaro A, Brahmer J, Antonia S, Mok T and Peters S: Managing resistance to immune checkpoint inhibitors in lung cancer: Treatment and novel strategies. J Clin Oncol. 40:598–610. 2022. View Article : Google Scholar : PubMed/NCBI

3 

Xiao M, He J, Yin L, Chen X, Zu X and Shen Y: Tumor-associated macrophages: Critical players in drug resistance of breast cancer. Front Immunol. 12:7994282021. View Article : Google Scholar

4 

Peng H, Xian D, Liu J, Pan S, Tang R and Zhong J: Regulating the polarization of macrophages: A promising approach to vascular dermatosis. J Immunol Res. 2020:81482722020. View Article : Google Scholar : PubMed/NCBI

5 

Hu Y, Tang J, Sun H, Li Y, Yu F, Zhang G, Chen J, Xu H, Zhong Z, Huang C, et al: Artesunate modulates the tumor microenvironment via STAT1/IRF1-mediated TAM repolarization and T cell activation in non-small cell lung cancer. Phytomedicine. 146:1570852025. View Article : Google Scholar : PubMed/NCBI

6 

Yu H, Zhang J, Liu Q, Liu L, Le Y, Cen H, Peng W, Wei J, Liu S, Qin A, et al: Biomimetic proteolipid vesicles delivering small activating RNA to activate the macrophage immunotherapy for the treatment of lung cancer. J Nanobiotechnology. 24:5082026. View Article : Google Scholar : PubMed/NCBI

7 

Chengzhi W, Songwei L, Yifan L, Mengmeng D and Huan L: The role of microRNAs in primary Sjögren's disease: Deciphering regulatory networks and assessing current therapeutic perspectives. Front Immunol. 16:16693822025. View Article : Google Scholar

8 

Wang Y, Chen W, Yu B, Wang L and Tang W: Exosomal miR-93-5p modulates macrophage polarization to enhance prostate cancer progression. Arch Biochem Biophys. 779:1107532026. View Article : Google Scholar : PubMed/NCBI

9 

Bartoszewska E, Misiąg P, Czapla M, Rakoczy K, Tomecka P, Filipski M, Wawrzyniak-Dzierżek E and Choromańska A: The role of microRNAs in lung cancer: Mechanisms, diagnostics and therapeutic potential. Int J Mol Sci. 26:37362025. View Article : Google Scholar : PubMed/NCBI

10 

Huang Q, Chu X, Yang C, Huai Y, He C, Ma X, Pei J, Gao J, Liu Z, Jiang S, et al: Bio-engineered microRNA-7 effectively interferes with the Akt3/p53 axis to suppress human non-small cell lung cancer. Cancer Cell Int. 25:2502025. View Article : Google Scholar : PubMed/NCBI

11 

Deng J and Wu M: COX10-AS1-mediated miR-361-5p regulated cell invasion and migration by targeting SPRY1 in oral squamous cell carcinoma. Am J Transl Res. 15:2191–2206. 2023.PubMed/NCBI

12 

Steadman K, You S, Srinivas DV, Mouakkad L, Yan Y, Kim M, Venugopal SV, Tanaka H and Freeman MR: Autonomous action and cooperativity between the ONECUT2 transcription factor and its 3'untranslated region. Front Cell Dev Biol. 11:12062592023. View Article : Google Scholar

13 

Song K and Artibani M: The role of DNA methylation in ovarian cancer chemoresistance: A narrative review. Health Sci Rep. 6:e12352023. View Article : Google Scholar : PubMed/NCBI

14 

Lu TX and Rothenberg ME: MicroRNA. J Allergy Clin Immunol. 141:1202–1207. 2018. View Article : Google Scholar

15 

El Founini Y, Chaoui I, Dehbi H, El Mzibri M, Abounader R and Guessous F: MicroRNAs: Key regulators in lung cancer. Microrna. 10:109–122. 2021. View Article : Google Scholar : PubMed/NCBI

16 

Geekiyanage H, Rayatpisheh S, Wohlschlegel JA, Brown R and Ambros V: Extracellular microRNAs in human circulation are associated with miRISC complexes that are accessible to anti-AGO2 antibody and can bind target mimic oligonucleotides. Proc Natl Acad Sci USA. 117:24213–24223. 2020. View Article : Google Scholar : PubMed/NCBI

17 

Shanehbandi D, Asadi M, Seyedrezazadeh E, Zafari V, Shekari N, Akbari M, Rahbarnia L and Zarredar H: MicroRNA-based biomarkers in lung cancer: Recent advances and potential applications. Curr Mol Med. 23:648–667. 2023. View Article : Google Scholar

18 

Liu J, Wang W, Wang K, Liu W, Zhao Y, Han X, Wang L and Jiang BH: HDAC1 and FOXK1 mediate EGFR-TKI resistance of non-small cell lung cancer through miR-33a silencing. J Transl Med. 22:7932024. View Article : Google Scholar : PubMed/NCBI

19 

Shintani T, Shun YT, Toyozumi Y, Ikemura K, Shiroyama T, Nagatomo I, Jingushi K, Takeda Y, Kumanogoh A and Okuda M: MicroRNA-130a-3p regulates osimertinib resistance by targeting runt-related transcription factor 3 in lung adenocarcinoma. Sci Rep. 14:244292024. View Article : Google Scholar : PubMed/NCBI

20 

Ni ZZ, He JK, Tang X, Tao Z, Zhang Y and Xie B: Identification of ELAVL1 gene and miRNA-139-3p involved in the aggressiveness of NSCLC. Eur Rev Med Pharmacol Sci. 24:9453–9464. 2020.PubMed/NCBI

21 

Locati M, Curtale G and Mantovani A: Diversity, mechanisms, and significance of macrophage plasticity. Annu Rev Pathol. 15:123–147. 2020. View Article : Google Scholar

22 

Peng Y, Zhou M, Yang H, Qu R, Qiu Y, Hao J, Bi H and Guo D: Regulatory mechanism of M1/M2 macrophage polarization in the development of autoimmune diseases. Mediators Inflamm. 2023:88216102023. View Article : Google Scholar : PubMed/NCBI

23 

Li SJ, Wang XH, Li LR, Chen L and Sun ZJ: Leveraging macrophage plasticity for precision-targeted tumor immunotherapy. Biochim Biophys Acta Mol Basis Dis. 1872:1681762026. View Article : Google Scholar : PubMed/NCBI

24 

Zhang J, Zhou X and Hao H: Macrophage phenotype-switching in cancer. Eur J Pharmacol. 931:1752292022. View Article : Google Scholar : PubMed/NCBI

25 

Zou Z, Lin H, Li M and Lin B: Tumor-associated macrophage polarization in the inflammatory tumor microenvironment. Front Oncol. 13:11031492023. View Article : Google Scholar : PubMed/NCBI

26 

Goswami KK, Bose A and Baral R: Macrophages in tumor: An inflammatory perspective. Clin Immunol. 232:1088752021. View Article : Google Scholar : PubMed/NCBI

27 

Gao J, Liang Y and Wang L: Shaping polarization of tumor-associated macrophages In cancer immunotherapy. Front Immunol. 13:8887132022. View Article : Google Scholar : PubMed/NCBI

28 

Liang X, Bai R, Sun Y, Ma P, Lu X, Han T and Guan L: TAMs: Guardians or foes? Overcoming heterogeneity and delivery barriers for precision immunotherapy. Int Immunopharmacol. 163:1151952025. View Article : Google Scholar : PubMed/NCBI

29 

Yang Y, Li S, To KKW, Zhu S, Wang F and Fu L: Tumor-associated macrophages remodel the suppressive tumor immune microenvironment and targeted therapy for immunotherapy. J Exp Clin Cancer Res. 44:1452025. View Article : Google Scholar : PubMed/NCBI

30 

Saeed AF: Tumor-associated macrophages: Polarization, immunoregulation, and immunotherapy. Cells. 14:7412025. View Article : Google Scholar : PubMed/NCBI

31 

Xu W, Zhao X, Daha MR and van Kooten C: Reversible differentiation of pro- and anti-inflammatory macrophages. Mol Immunol. 53:179–186. 2013. View Article : Google Scholar

32 

Yang L, Shao Y, Zhang Z, Li Y, Wang F and Yu H: Metabolic insights into TAMs and the tumor immune microenvironment: Regulatory mechanisms and therapeutic interventions. Biochim Biophys Acta Rev Cancer. 1880:1894112025. View Article : Google Scholar : PubMed/NCBI

33 

Ni Z, Zhou S, Zhang Y, Wang X, Yang H, Zhou H and Tao Z: THOC3 interacts with epithelial-to-mesenchymal transition to promote non-small cell lung cancer carcinoma progression through STAT3 signaling pathway. Transl Oncol. 67:1027412026. View Article : Google Scholar : PubMed/NCBI

34 

Li X, Chen Z, Ni Y, Bian C, Huang J, Chen L, Xie X and Wang J: Tumor-associated macrophages secret exosomal miR-155 and miR-196a-5p to promote metastasis of non-small-cell lung cancer. Transl Lung Cancer Res. 10:1338–1354. 2021. View Article : Google Scholar : PubMed/NCBI

35 

Mi L, Xu L, Chen Z and Zheng Y: The influence of CXCL9 on M2 macrophages in lung cancer development. Transl Cancer Res. 14:8965–8978. 2025. View Article : Google Scholar

36 

Lu CS, Shiau AL, Su BH, Hsu TS, Wang CT, Su YC, Tsai MS, Feng YH, Tseng YL, Yen YT, et al: Oct4 promotes M2 macrophage polarization through upregulation of macrophage colony-stimulating factor in lung cancer. J Hematol Oncol. 13:622020. View Article : Google Scholar : PubMed/NCBI

37 

Ahmad A, Tiwari RK, Almeleebia TM, Al Fayi MS, Alshahrani MY, Ahmad I, Abohassan MS, Saeed M and Ansari IA: Swertia chirayita suppresses the growth of non-small cell lung cancer A549 cells and concomitantly induces apoptosis via downregulation of JAK1/STAT3 pathway. Saudi J Biol Sci. 28:6279–6288. 2021. View Article : Google Scholar : PubMed/NCBI

38 

Hsiao YJ, Hsieh MS, Chang GC, Hsu YC, Wang CY, Chen YM, Chen YL, Yang PC and Yu SL: Tp53 determines the spatial dynamics of M1/M2 tumor-associated macrophages and M1-driven tumoricidal effects. Cell Death Dis. 16:382025. View Article : Google Scholar : PubMed/NCBI

39 

Kiran BK, Siddesh BM, Sherapura A, Thirusangu P, Vijay Avin BR, Vigneshwaran V, Kumaraswamy HM, Suchetha Kumari N, Pramod SN and Prabhakar BT: Allium sativum lectin as an immunotherapeutic agent: Eliciting IFN-γ production via 'M1' macrophage polarization to trigger JAK1/STAT1 and DDR mediated lung cancer cell death. Int J Biol Macromol. 330:1480762025. View Article : Google Scholar

40 

Hao D and Chen S: Targeting tumor-associated macrophages in non-small cell lung cancer: Mechanisms, prognosis, and therapeutic opportunities. Front Immunol. 16:16795372025. View Article : Google Scholar : PubMed/NCBI

41 

Yang FR, Li HL, Hu XW, Fu R, Li XR and Li HJ: Chinese herbal compound xiaoliu pingyi recipe inhibits the growth of lung adenocarcinoma by regulating the tumor vascular microenvironment. Integr Cancer Ther. 23:153473542412739622024. View Article : Google Scholar : PubMed/NCBI

42 

Zhou Y, Ren D, Bi H, Yi B, Zhang C, Wang H and Sun J: Tumor-associated macrophage: Emerging targets for modulating the tumor microenvironment. Zhongguo Fei AI Za Zhi. 27:231–240. 2024.In Chinese. PubMed/NCBI

43 

Brown JM, Recht L and Strober S: The promise of targeting macrophages in cancer therapy. Clin Cancer Res. 23:3241–3250. 2017. View Article : Google Scholar : PubMed/NCBI

44 

Orlichenko LS and Radisky DC: Matrix metalloproteinases stimulate epithelial-mesenchymal transition during tumor development. Clin Exp Metastasis. 25:593–600. 2008. View Article : Google Scholar : PubMed/NCBI

45 

Wei K, Ma Z, Yang F, Zhao X, Jiang W, Pan C, Li Z, Pan X, He Z, Xu J, et al: M2 macrophage-derived exosomes promote lung adenocarcinoma progression by delivering miR-942. Cancer Lett. 526:205–216. 2022. View Article : Google Scholar

46 

Gao W, Wang R, Yang S, Shi Y, Cui M, Jiang R, Li R, Yu Y, Jia D and Che D: MS4A1 regulates M1-polarized tumor-associated macrophage infiltration, angiogenesis, and cancer progression through the HIPPO pathway in lung adenocarcinoma. Cancer Immunol Immunother. 74:3562025. View Article : Google Scholar : PubMed/NCBI

47 

Dhayalan S, Ramarajyam G, Mohanprasanth A, Muruhesan D, Rajendiran S and Arjunan S: Trigonelline suppresses the tumor progression via STAT2 signalling pathway in non-small cell lung carcinoma. Med Oncol. 43:1742026. View Article : Google Scholar : PubMed/NCBI

48 

Ma WP, Hu SM, Xu YL, Li HH, Ma XQ, Wei BH, Li FY, Guan HS, Yu GL, Liu M and Liu HB: Haimufang decoction, a chinese medicine formula for lung cancer, arrests cell cycle, stimulates apoptosis in NCI-H1975 cells, and induces M1 polarization in RAW 264.7 macrophage cells. BMC Complement Med Ther. 20:2432020. View Article : Google Scholar : PubMed/NCBI

49 

Zhu C, Zhang F, Li X, Xing D, Cai H, Wang X, Qu H, Li L and Zheng X: Isoginkgetin inhibits non-small cell lung cancer by inducing oxidative stress and regulating M1 macrophage polarization. Phytomedicine. 152:1578692026. View Article : Google Scholar : PubMed/NCBI

50 

Cui L, Zhao S, Lu G, Zhang S, Zhang R, Zhou J and Yao Y: ACTL6A accelerates the progression of NSCLC through hippo/YAP signaling axis and TAMs-mediated immune regulation. Int Immunopharmacol. 168:1158282026. View Article : Google Scholar

51 

Liu M, Wang T, Fu J, Cui T, Li N, Sun T, Liu J, Su C and Song B: Pinellia exosomal vesicles remodulate tumor-associated macrophage polarization via the serine synthesis/JAK/STAT signaling pathway to inhibit lung cancer growth. Phytomedicine. 151:1577592026. View Article : Google Scholar : PubMed/NCBI

52 

Wang Q, Zhang Q, Wang X, Luo H, Du T, Wu L, Tan M, Chen Y, Wu X, Sun S, et al: TGM2-mediated autophagy contributes to the radio-resistance of non-small cell lung cancer stem-like cells. Biomedicines. 12:22312024. View Article : Google Scholar : PubMed/NCBI

53 

Zhang X, Zhu M, Hong Z and Chen C: Co-culturing polarized M2 thp-1-derived macrophages enhance stemness of lung adenocarcinoma A549 cells. Ann Transl Med. 9:7092021. View Article : Google Scholar : PubMed/NCBI

54 

Liu Y and Sun Y: TIPE2 inhibits the stemness of lung cancer cells by regulating the phenotypic polarization of tumor-associated macrophages. Xi Bao Yu Fen Zi Mian Yi Xue Za Zhi. 41:681–686. 2025.In Chinese.

55 

Huang WC, Chan ML, Chen MJ, Tsai TH and Chen YJ: Modulation of macrophage polarization and lung cancer cell stemness by MUC1 and development of a related small-molecule inhibitor pterostilbene. Oncotarget. 7:39363–39375. 2016. View Article : Google Scholar : PubMed/NCBI

56 

Guo M, Li H, Zhao Z, Wang Y, Tang J, Zhong B and Zhao Y: Suppressive role of SCN4B in the epithelial-mesenchymal transition of lung adenocarcinoma. Oncol Rep. 55:682026. View Article : Google Scholar

57 

Engur-Ozturk S and Dikmen M: Proteasome inhibitor immunotherapy for the epithelial to mesenchymal transition: Assessing the A549 lung cancer cell microenvironment and the role of M1, M2a and M2c 'hydrocortisone-polarised' macrophages. Mol Biol Rep. 49:4777–4793. 2022. View Article : Google Scholar : PubMed/NCBI

58 

Kawaguchi Y, Ohshio Y, Watanabe A, Shiratori T, Okamoto K, Ueda K, Kataoka Y, Suzuki T and Hanaoka J: Depletion of tumor-associated macrophages inhibits lung cancer growth and enhances the antitumor effect of cisplatin. Cancer Sci. 114:751–763. 2023. View Article : Google Scholar

59 

Liao L, Wang YX, Fan SS, Hu YY, Wang XC and Zhang X: The role and clinical significance of tumor-associated macrophages in the epithelial-mesenchymal transition of lung cancer. Front Oncol. 15:15715832025. View Article : Google Scholar : PubMed/NCBI

60 

Hou Y, Li X, Zeng L and Zhang Y: SNHG16-loaded extracellular vesicles from hypoxic NSCLC cells drive M2 macrophage polarization to enhance cancer aggressiveness. Mol Immunol. 187:66–79. 2025. View Article : Google Scholar : PubMed/NCBI

61 

Yang J, Wang X, Huang B, Liu R, Xiong H, Ye F, Zeng C, Fu X and Li L: An IFNγ/STAT1/JMJD3 axis induces ZEB1 expression and promotes aggressiveness in lung adenocarcinoma. Mol Cancer Res. 19:1234–1246. 2021. View Article : Google Scholar : PubMed/NCBI

62 

Curtale G, Rubino M and Locati M: MicroRNAs as molecular switches in macrophage activation. Front Immunol. 10:7992019. View Article : Google Scholar : PubMed/NCBI

63 

Saric A, Hipolito VEB, Kay JG, Canton J, Antonescu CN and Botelho RJ: mTOR controls lysosome tubulation and antigen presentation in macrophages and dendritic cells. Mol Biol Cell. 27:321–333. 2016. View Article : Google Scholar :

64 

Wang L, Hu YY, Zhao JL, Huang F, Liang SQ, Dong L, Chen Y, Yu HC, Bai J, Yang JM, et al: Targeted delivery of miR-99b reprograms tumor-associated macrophage phenotype leading to tumor regression. J Immunother Cancer. 8:e0005172020. View Article : Google Scholar : PubMed/NCBI

65 

Chirshev E, Oberg KC, Ioffe YJ and Unternaehrer JJ: Let-7 as biomarker, prognostic indicator, and therapy for precision medicine in cancer. Clin Transl Med. 8:242019. View Article : Google Scholar : PubMed/NCBI

66 

Rong J, Xu L, Hu Y, Liu F, Yu Y, Guo H, Ni X, Huang Y, Zhao L and Wang Z: Inhibition of let-7b-5p contributes to an anti-tumorigenic macrophage phenotype through the SOCS1/STAT pathway in prostate cancer. Cancer Cell Int. 20:4702020. View Article : Google Scholar : PubMed/NCBI

67 

Jeong J, Hwang YE, Lee M, Keum S, Song S, Kim JW, Choi JH and Rhee S: Downregulation of AP1S1 causes the lysosomal degradation of EGFR in non-small cell lung cancer. J Cell Physiol. 238:2335–2347. 2023. View Article : Google Scholar : PubMed/NCBI

68 

Liu L, Kou Z, Wang T, Shang Q, Zhang Q, Liu G, Ai J, Zhao Y and Sun C: Mechanism of non-small cell lung cancer-derived extracellular vesicle miRNA hsa-let-7b-5p targeting AP1S1 to regulate M2 macrophage polarization. Int J Genomics. 2026:82204782026. View Article : Google Scholar : PubMed/NCBI

69 

Shi JH, Liu LN, Song DD, Liu WW, Ling C, Wu FX, Wang TT, Liu B, Cui NP, Qin Y and Ni ZY: TRAF3/STAT6 axis regulates macrophage polarization and tumor progression. Cell Death Differ. 30:2005–2016. 2023. View Article : Google Scholar : PubMed/NCBI

70 

Diao H, Xu X, Zhao B and Yang G: miR-135a-5p inhibits tumor invasion by targeting ANGPT2 in gallbladder cancer. Mol Med Rep. 24:5282021. View Article : Google Scholar

71 

Zhang H, Wang M, Lang Z, Liu H, Liu J and Ma L: MiR-135a-5p suppresses breast cancer cell proliferation, migration, and invasion by regulating BAG3. Clinics (Sao Paulo). 77:1001152022. View Article : Google Scholar : PubMed/NCBI

72 

Gao HX, Liu MH, Fan M, Zhou JJ, Li AQ and Chen MW: MiR-135a-5p/STAT6-mediated EMT regulates IL-4 secretion in non-small cell lung cancer to affect M2-like TAM polarization. Int Immunopharmacol. 155:1146232025. View Article : Google Scholar : PubMed/NCBI

73 

Zhang Z, Yang Y and Zhang X: MiR-770 inhibits tumorigenesis and EMT by targeting JMJD6 and regulating WNT/β-catenin pathway in non-small cell lung cancer. Life Sci. 188:163–171. 2017. View Article : Google Scholar : PubMed/NCBI

74 

Haupt M, Zechmeister B, Bosche B, Lieschke S, Zheng X, Zhang L, Venkataramani V, Jin F, Hein K, Weber MS, et al: Lithium enhances post-stroke blood-brain barrier integrity, activates the MAPK/ERK1/2 pathway and alters immune cell migration in mice. Neuropharmacology. 181:1083572020. View Article : Google Scholar : PubMed/NCBI

75 

Liu J, Luo R, Wang J, Luan X, Wu D, Chen H, Hou Q, Mao G and Li X: Tumor cell-derived exosomal miR-770 inhibits M2 macrophage polarization via targeting MAP3K1 to inhibit the invasion of non-small cell lung cancer cells. Front Cell Dev Biol. 9:6796582021. View Article : Google Scholar : PubMed/NCBI

76 

Ying H, Jin Y, Guo Y, Li Q, Ruan M, Zhu W, Yang C, Li Q and Zheng L: Long non-coding RNA NUT family member 2A-antisense RNA 1 sponges microRNA-613 to increase the resistance of gastric cancer cells to matrine through regulating oxidative stress and vascular endothelial growth factor A. Aging (Albany NY). 14:5153–5162. 2022. View Article : Google Scholar : PubMed/NCBI

77 

Yang M, Zhou W, Xu M, Han X, Shi Y, Shi M and Wang Z: Tumor suppressor miR-613 alleviates non-small cell lung cancer cell via repressing M2 macrophage polarization. J Oncol. 2023:23112312023. View Article : Google Scholar : PubMed/NCBI

78 

Almendros I, Khalyfa A, Trzepizur W, Gileles-Hillel A, Huang L, Akbarpour M, Andrade J, Farré R and Gozal D: Tumor cell malignant properties are enhanced by circulating exosomes in sleep apnea. Chest. 150:1031–1041. 2016. View Article : Google Scholar

79 

Almendros I, Wang Y, Becker L, Lennon FE, Zheng J, Coats BR, Schoenfelt KS, Carreras A, Hakim F, Zhang SX, et al: Intermittent hypoxia-induced changes in tumor-associated macrophages and tumor malignancy in a mouse model of sleep apnea. Am J Respir Crit Care Med. 189:593–601. 2014. View Article : Google Scholar : PubMed/NCBI

80 

Shan X, Zhang H, Zhang L, Zhou X, Wang T, Zhang J, Shu Y, Zhu W, Wen W and Liu P: Identification of four plasma microRNAs as potential biomarkers in the diagnosis of Male lung squamous cell carcinoma patients in China. Cancer Med. 7:2371–2381. 2018. View Article : Google Scholar

81 

Ren J, Jin Z and Huang Y: Exosomal miR-106a-5p derived from intermittently hypoxic non-small-cell lung cancer increases tumor malignancy. Physiol Rep. 12:e161572024. View Article : Google Scholar : PubMed/NCBI

82 

He G, Peng X, Wei S, Yang S, Li X, Huang M, Tang S, Jin H, Liu J, Zhang S, et al: Exosomes in the hypoxic TME: From release, uptake and biofunctions to clinical applications. Mol Cancer. 21:192022. View Article : Google Scholar : PubMed/NCBI

83 

He B, You L, Uematsu K, Zang K, Xu Z, Lee AY, Costello JF, McCormick F and Jablons DM: SOCS-3 is frequently silenced by hypermethylation and suppresses cell growth in human lung cancer. Proc Natl Acad Sci USA. 100:14133–14138. 2003. View Article : Google Scholar : PubMed/NCBI

84 

Gu J, Yang S, Wang X, Wu Y, Wei J and Xu J: Hypoxic lung adenocarcinoma-derived exosomal miR-1290 induces M2 macrophage polarization by targeting SOCS3. Cancer Med. 12:12639–12652. 2023. View Article : Google Scholar : PubMed/NCBI

85 

Zhu X, Guo Q, Zou J, Wang B, Zhang Z, Wei R, Zhao L, Zhang Y, Chu C, Fu X and Li X: MiR-19a-3p suppresses M1 macrophage polarization by inhibiting STAT1/IRF1 pathway. Front Pharmacol. 12:6140442021. View Article : Google Scholar : PubMed/NCBI

86 

Dai L, Chen F, Zheng Y, Zhang D, Qian B, Ji H, Long F and Cretoiu D: miR-21 regulates growth and EMT in lung cancer cells via PTEN/akt/GSK3β signaling. Front Biosci. 24:1426–1439. 2019. View Article : Google Scholar

87 

Jin J and Yu G: Hypoxic lung cancer cell-derived exosomal miR-21 mediates macrophage M2 polarization and promotes cancer cell proliferation through targeting IRF1. World J Surg Oncol. 20:2412022. View Article : Google Scholar : PubMed/NCBI

88 

You G, Yang Q, Li X and Chen L: TMEM33, an oncogene regulated by miR-214-3p, promotes the progression of lung adenocarcinoma through the wnt/β-catenin signaling pathway. Oncol Res. 33:905–917. 2025. View Article : Google Scholar

89 

Zhu S and Liao B: Mechanism of hypoxia-induced exosome circ_0051799 regulating the progression of lung adenocarcinoma. Biol Chem. 405:143–160. 2024. View Article : Google Scholar

90 

Lee J, Park HY, Kim WW, Lee SJ, Jeong JH, Kang SH, Jung JH and Chae YS: Biological function of long noncoding RNA snaR in HER2-positive breast cancer cells. Tumour Biol J. 39:10104283177073742017.

91 

Nakaoka HJ, Hara T, Yoshino S, Kanamori A, Matsui Y, Shimamura T, Sato H, Murakami Y, Seiki M and Sakamoto T: NECAB3 promotes activation of hypoxia-inducible factor-1 during normoxia and enhances tumourigenicity of cancer cells. Sci Rep. 6:227842016. View Article : Google Scholar : PubMed/NCBI

92 

Li Z, Feng C, Guo J, Hu X and Xie D: GNAS-AS1/miR-4319/NECAB3 axis promotes migration and invasion of non-small cell lung cancer cells by altering macrophage polarization. Funct Integr Genomics. 20:17–28. 2020. View Article : Google Scholar

93 

Bernardo ME and Fibbe WE: Mesenchymal stromal cells: Sensors and switchers of inflammation. Cell Stem Cell. 13:392–402. 2013. View Article : Google Scholar : PubMed/NCBI

94 

Poggi A, Varesano S and Zocchi MR: How to hit mesenchymal stromal cells and make the tumor microenvironment immunostimulant rather than immunosuppressive. Front Immunol. 9:2622018. View Article : Google Scholar : PubMed/NCBI

95 

Yang M, Shen H, Qiu C, Ni Y, Wang L, Dong W, Liao Y and Du J: High expression of miR-21 and miR-155 predicts recurrence and unfavourable survival in non-small cell lung cancer. Eur J Cancer. 49:604–615. 2013. View Article : Google Scholar

96 

Xi J, Huang Q, Wang L, Ma X, Deng Q, Kumar M, Zhou Z, Li L, Zeng Z, Young KH, et al: miR-21 depletion in macrophages promotes tumoricidal polarization and enhances PD-1 immunotherapy. Oncogene. 37:3151–3165. 2018. View Article : Google Scholar : PubMed/NCBI

97 

Ren W, Hou J, Yang C, Wang H, Wu S, Wu Y, Zhao X and Lu C: Extracellular vesicles secreted by hypoxia pre-challenged mesenchymal stem cells promote non-small cell lung cancer cell growth and mobility as well as macrophage M2 polarization via miR-21-5p delivery. J Exp Clin Cancer Res. 38:622019. View Article : Google Scholar : PubMed/NCBI

98 

Pitt JM, Kroemer G and Zitvogel L: Extracellular vesicles: Masters of intercellular communication and potential clinical interventions. J Clin Invest. 126:1139–1143. 2016. View Article : Google Scholar : PubMed/NCBI

99 

Yang Q, Diamond MP and Al-Hendy A: The emerging role of extracellular vesicle-derived miRNAs: Implication in cancer progression and stem cell related diseases. J Clin Epigenetics. 2:132016.

100 

Triner D and Shah YM: Hypoxia-inducible factors: A central link between inflammation and cancer. J Clin Invest. 126:3689–3698. 2016. View Article : Google Scholar : PubMed/NCBI

101 

Akers JC, Ramakrishnan V, Kim R, Phillips S, Kaimal V, Mao Y, Hua W, Yang I, Fu CC, Nolan J, et al: miRNA contents of cerebrospinal fluid extracellular vesicles in glioblastoma patients. J Neurooncol. 123:205–216. 2015. View Article : Google Scholar : PubMed/NCBI

102 

Hsu YL, Hung JY, Chang WA, Jian SF, Lin YS, Pan YC, Wu CY and Kuo PL: Hypoxic lung-cancer-derived extracellular vesicle MicroRNA-103a increases the oncogenic effects of macrophages by targeting PTEN. Mol Ther. 26:568–581. 2018. View Article : Google Scholar : PubMed/NCBI

103 

Xu P, Xiao H, Yang Q, Hu R, Jiang L, Bi R, Jiang X, Wang L, Mei J, Ding F and Huang J: The USP21/YY1/SNHG16 axis contributes to tumor proliferation, migration, and invasion of non-small-cell lung cancer. Exp Mol Med. 52:41–55. 2020. View Article : Google Scholar : PubMed/NCBI

104 

Dong Z, Liu H and Zhao G: Long noncoding RNA SNHG6 promotes proliferation and inhibits apoptosis in non-small cell lung cancer cells by regulating miR-490-3p/RSF1 axis. Cancer Biother Radiopharm. 35:351–361. 2020.PubMed/NCBI

105 

Brown S, Banfill K, Aznar MC, Whitehurst P and Faivre Finn C: The evolving role of radiotherapy in non-small cell lung cancer. Br J Radiol. 92:201905242019. View Article : Google Scholar : PubMed/NCBI

106 

Abdolvand M, Chermahini ZM, Bahaloo S, Emami MH, Fahim A, Rahimi H, Amjadi E, Maghool F, Rohani F, Dadkhah M, et al: New long noncoding RNA biomarkers and ceRNA networks on miR-616-3p in colorectal cancer: Bioinformatics-based study. J Res Med Sci. 29:102024. View Article : Google Scholar : PubMed/NCBI

107 

Song S, Zhao Y, Fu T, Fan Y, Tang J, Wang X, Liu C and Chen X: ELANE promotes M2 macrophage polarization by down-regulating PTEN and participates in the lung cancer progression. Immunol Invest. 52:21–34. 2023. View Article : Google Scholar

108 

Chen T, Liu Y, Li C, Xu C, Ding C, Chen J and Zhao J: Tumor-derived exosomal circFARSA mediates M2 macrophage polarization via the PTEN/PI3K/AKT pathway to promote non-small cell lung cancer metastasis. Cancer Treat Res Commun. 28:1004122021.PubMed/NCBI

109 

Yang L, Zhang Z, Zhang Y, Wang L, Zheng S, Li Y, Du D, Yuan C, Li H, Zeng Z, et al: Radiotherapy promotes M2 polarization of macrophages through the regulation of the PTEN/PI3K/AKT signaling pathway through miR-616-3p in lung cancer cell-derived exosomes. In Vitro Cell Dev Biol Anim. 61:1202–1217. 2025. View Article : Google Scholar : PubMed/NCBI

110 

Baba Y, Nomoto D, Okadome K, Ishimoto T, Iwatsuki M, Miyamoto Y, Yoshida N and Baba H: Tumor immune microenvironment and immune checkpoint inhibitors in esophageal squamous cell carcinoma. Cancer Sci. 111:3132–3141. 2020. View Article : Google Scholar : PubMed/NCBI

111 

Wang NH, Lei Z, Yang HN, Tang Z, Yang MQ, Wang Y, Sui JD and Wu YZ: Radiation-induced PD-L1 expression in tumor and its microenvironment facilitates cancer-immune escape: A narrative review. Ann Transl Med. 10:14062022. View Article : Google Scholar

112 

Xu L, Li K, Li J, Xu F, Liang S, Kong Y and Chen B: The crosstalk between lung adenocarcinoma cells and M2 macrophages promotes cancer cell development via the SFRS1/miR-708-5p/PD-L1 axis. Life Sci. 371:1235992025. View Article : Google Scholar : PubMed/NCBI

113 

Hu JL, Wang W, Lan XL, Zeng ZC, Liang YS, Yan YR, Song FY, Wang FF, Zhu XH, Liao WJ, et al: CAFs secreted exosomes promote metastasis and chemotherapy resistance by enhancing cell stemness and epithelial-mesenchymal transition in colorectal cancer. Mol Cancer. 18:912019. View Article : Google Scholar : PubMed/NCBI

114 

Zhang D and Yang N: MiR-335-5p inhibits cell proliferation, migration and invasion in colorectal cancer through downregulating LDHB. J BUON. 24:1128–1136. 2019.PubMed/NCBI

115 

Yu Y and Ren K: Five long non-coding RNAs establish a prognostic nomogram and construct a competing endogenous RNA network in the progression of non-small cell lung cancer. BMC Cancer. 21:4572021. View Article : Google Scholar : PubMed/NCBI

116 

Skehel PA, Fabian-Fine R and Kandel ER: Mouse VAP33 is associated with the endoplasmic reticulum and microtubules. Proc Natl Acad Sci USA. 97:1101–1106. 2000. View Article : Google Scholar : PubMed/NCBI

117 

Chen J, Sun JJ, Ma YW, Zhu MQ, Hu J, Lu QJ and Cai ZG: Cancer-associated fibroblasts derived exosomal LINC01833 promotes the occurrence of non-small cell lung cancer through miR-335-5p-VAPA axis. J Biochem Mol Toxicol. 38:e237692024. View Article : Google Scholar

118 

Zhai Y, Liu Y, Wang Z, Wang W, Zhou J and Lu J: Long non-coding RNA LINC00313 accelerates cervical carcinoma progression by miR-4677-3p/CDK6 axis. OncoTargets Ther. 14:2213–2226. 2021. View Article : Google Scholar

119 

Chen H, Wahafu P, Wang L and Chen X: LncRNA LINC00313 knockdown inhibits tumorigenesis and metastasis in human osteosarcoma by upregulating FOSL2 through sponging miR-342-3p. Yonsei Med J. 61:359–370. 2020. View Article : Google Scholar : PubMed/NCBI

120 

Dang QQ, Li PH, Wang J, Zhao JY, Zhai SN, Zheng YJ and Yang DK: CircMAN1A2 contributes to nasopharyngeal carcinoma progression via enhancing the ubiquitination of ATMIN through miR-135a-3p/UBR5 axis. Hum Cell. 36:657–675. 2023. View Article : Google Scholar : PubMed/NCBI

121 

Kong W, Zhang L, Chen Y, Yu Z and Zhao Z: Cancer cell-derived exosomal LINC00313 induces M2 macrophage differentiation in non-small cell lung cancer. Clin Transl Oncol. 24:2395–2408. 2022. View Article : Google Scholar : PubMed/NCBI

122 

Ding J, Wang X, Yang H, Zhang L, Ying Y, Pi W, Deng G and Zhu Y: IGF2BP3 Triggers STAT3 pathway by stabilizing SRC RNA in an m6A-Dependent manner to promote lymphatic metastasis in LUAD. Cancer Sci. 116:936–950. 2025. View Article : Google Scholar : PubMed/NCBI

123 

Chen J, Zhang K, Zhi Y, Wu Y, Chen B, Bai J and Wang X: Tumor-derived exosomal miR-19b-3p facilitates M2 macrophage polarization and exosomal LINC00273 secretion to promote lung adenocarcinoma metastasis via hippo pathway. Clin Transl Med. 11:e4782021. View Article : Google Scholar : PubMed/NCBI

124 

Lu GJ, Cui J, Qian Q, Hou ZB, Xie HY, Hu W, Hao KK, Xia N and Zhang Y: Overexpression of hsa_circ_0001715 is a potential diagnostic and prognostic biomarker in lung adenocarcinoma. Onco Targets Ther. 13:10775–10783. 2020. View Article : Google Scholar : PubMed/NCBI

125 

Liao J, Chen Z, Luo X, Su Y, Huang T, Xu H, Lin K, Zheng Q, Zhang L, Lin G and Lin X: Hsa_circ_0006692 promotes lung cancer progression via miR-205-5p/CDK19 axis. Genes (Basel). 13:8462022. View Article : Google Scholar : PubMed/NCBI

126 

Chen M, Cao C and Ma J: Tumor-related exosomal circ_0001715 promotes lung adenocarcinoma cell proliferation and metastasis via enhancing M2 macrophage polarization by regulating triggering receptor expressed on myeloid cells-2. Thorac Cancer. 15:227–238. 2024. View Article : Google Scholar :

127 

Stawowczyk M, Wellenstein MD, Lee SB, Yomtoubian S, Durrans A, Choi H, Narula N, Altorki NK, Gao D and Mittal V: Matrix metalloproteinase 14 promotes lung cancer by cleavage of Heparin-Binding EGF-like Growth Factor. Neoplasia. 19:55–64. 2017. View Article : Google Scholar :

128 

Ji F, Du R, Chen T, Zhang M, Zhu Y, Luo X and Ding Y: Circular RNA circSLC26A4 accelerates cervical cancer progression via miR-1287-5p/HOXA7 axis. Mol Ther Nucleic Acids. 19:413–420. 2020. View Article : Google Scholar : PubMed/NCBI

129 

Qian J, Li J, Ma H and Ji W: Exosomal circ-ADRM1 promotes lung adenocarcinoma progression and induces macrophage M2 polarization through regulating MMP14 mRNA and protein. Anticancer Drugs. 34:333–343. 2023. View Article : Google Scholar

130 

Jin M, Li L, Xu C, Wen Y and Zhao M: Estrogenic activities of two synthetic pyrethroids and their metabolites. J Environ Sci (China). 22:291–296. 2010. View Article : Google Scholar

131 

Cai X, Yin Y, Li N, Zhu D, Zhang J, Zhang CY and Zen K: Re-polarization of tumor-associated macrophages to pro-inflammatory M1 macrophages by microRNA-155. J Mol Cell Biol. 4:341–343. 2012. View Article : Google Scholar : PubMed/NCBI

132 

Huang F, Chen Z, Chen H, Lu W, Xie S, Meng QH, Wu Y and Xia D: Cypermethrin promotes lung cancer metastasis via modulation of macrophage polarization by targeting MicroRNA-155/Bcl6. Toxicol Sci. 163:454–465. 2018. View Article : Google Scholar : PubMed/NCBI

133 

Larhammar M, Huntwork-Rodriguez S, Rudhard Y, Sengupta-Ghosh A and Lewcock JW: The Ste20 family kinases MAP4K4, MINK1, and TNIK converge to regulate stress-induced JNK signaling in neurons. J Neurosci. 37:11074–11084. 2017. View Article : Google Scholar : PubMed/NCBI

134 

Hao J, Hu Y, Li Y, Zhou Q and Lv X: Involvement of JNK signaling in IL4-induced M2 macrophage polarization. Exp Cell Res. 357:155–162. 2017. View Article : Google Scholar : PubMed/NCBI

135 

Ma Y, Pan X, Xu P, Mi Y, Wang W, Wu X, He Q, Liu X, Tang W and An HX: Plasma microRNA alterations between EGFR-activating mutational NSCLC patients with and without primary resistance to TKI. Oncotarget. 8:88529–88536. 2017. View Article : Google Scholar : PubMed/NCBI

136 

Xu L, Wang L, Yang R, Li T and Zhu X: Lung adenocarcinoma cell-derived exosomes promote M2 macrophage polarization through transmission of miR-3153 to activate the JNK signaling pathway. Hum Mol Genet. 32:2162–2176. 2023. View Article : Google Scholar : PubMed/NCBI

137 

Li Y, Li Y, Liu J, Fan Y, Li X, Dong M, Liu H and Chen J: Expression levels of microRNA-145 and microRNA-10b are associated with metastasis in non-small cell lung cancer. Cancer Biol Ther. 17:272–279. 2016. View Article : Google Scholar : PubMed/NCBI

138 

Yuan Y, Guo L and Guo S: Exosomal miR-10b promotes invasion and epithelial-mesenchymal transformation of lung adenocarcinoma A549 cells by regulating macrophage M2 polarization. Zhongguo Fei Ai Za Zhi. 25:835–842. 2022.In Chinese.

139 

Li Y, Zhao L, Shi B, Ma S, Xu Z, Ge Y, Liu Y, Zheng D and Shi J: Functions of miR-146a and miR-222 in tumor-associated macrophages in breast cancer. Sci Rep. 5:186482015. View Article : Google Scholar : PubMed/NCBI

140 

Yin C, Han Q, Xu D, Zheng B, Zhao X and Zhang J: SALL4-mediated upregulation of exosomal miR-146a-5p drives T-cell exhaustion by M2 tumor-associated macrophages in HCC. Oncoimmunology. 8:16014792019. View Article : Google Scholar : PubMed/NCBI

141 

Yan Z, Wen JX, Cao XS, Zhao W, Han YL, Wen XH, Yan L, Zhang M, Wang YF, Hai L, et al: Tumor cell-derived exosomal microRNA-146a promotes non-small cell lung cancer cell invasion and proliferation by inhibiting M1 macrophage polarization. Ann Transl Med. 10:13072022. View Article : Google Scholar

142 

Jiang K, Yang J, Guo S, Zhao G, Wu H and Deng G: Peripheral circulating exosome-mediated delivery of miR-155 as a novel mechanism for acute lung inflammation. Mol Ther J. 27:1758–1771. 2019. View Article : Google Scholar

143 

Lu L, McCurdy S, Huang S, Zhu X, Peplowska K, Tiirikainen M, Boisvert WA and Garmire LX: Time series miRNA-mRNA integrated analysis reveals critical miRNAs and targets in macrophage polarization. Sci Rep. 6:374462016. View Article : Google Scholar : PubMed/NCBI

144 

Hsin JP, Lu Y, Loeb GB, Leslie CS and Rudensky AY: The effect of cellular context on miR-155-mediated gene regulation in four major immune cell types. Nat Immunol. 19:1137–1145. 2018. View Article : Google Scholar : PubMed/NCBI

145 

Liu Y, Lu M, Liu F, Xu G, Feng C, Chen Y, Cai D, Sun H, Zeng Y, Xie J, et al: Extracellular vesicles obtained from lung adenocarcinoma cells cultured under intermittent hypoxia induce M2 macrophage polarization via miR-20a-5p delivery. Technol Cancer Res Treat. 23:153303382312194152024. View Article : Google Scholar : PubMed/NCBI

146 

Lin G, Lin L, Chen X, Chen L, Yang J, Chen Y, Qian D, Zeng Y and Xu Y: PPAR-γ/NF-kB/AQP3 axis in M2 macrophage orchestrates lung adenocarcinoma progression by upregulating IL-6. Cell Death Dis. 15:5322024. View Article : Google Scholar

147 

Courau T, Jaszczak RG, Samad B, Flynn E, Chew NW, Reeder GC, Tsui J, Teklu S, Pass LF, Edwards AW, et al: Differential assembly of mouse and human tumor microenvironments. Nat Immunol. 27:1282–1293. 2026. View Article : Google Scholar :

148 

Raffo-Romero A, Ziane-Chaouche L, Salomé-Desnoulez S, Hajjaji N, Fournier I, Salzet M and Duhamel M: A co-culture system of macrophages with breast cancer tumoroids to study cell interactions and therapeutic responses. Cell Rep Methods. 4:1007922024. View Article : Google Scholar : PubMed/NCBI

149 

de la Rosa C, Kendirli A, Baygün S, Bauernschmitt F, Thomann AS, Kisioglu I, Beckmann D, Carpentier Solorio Y, Pfaffenstaller V, Tai YH, et al: In vivo CRISPR screen reveals regulation of macrophage states in neuroinflammation. Nat Neurosci. 29:493–509. 2026. View Article : Google Scholar :

150 

Liu H, Shi X, Wang D, Zhang H, Xu Z and Tan Z: Three-dimensional-printed In vitro model of colorectal cancer with immune microenvironment and reprogramming capabilities. ACS Biomater Sci Eng. 11:5991–6003. 2025. View Article : Google Scholar : PubMed/NCBI

151 

Shang W, Lai C, Luo S and Chen L: Murine models of lung adenocarcinoma: Valuable tools for preclinical investigation. Gene. 971:1498022025. View Article : Google Scholar : PubMed/NCBI

152 

Li J, Dhilipkannah P, Holden VK, Sachdeva A, Todd NW and Jiang F: Dysregulation of lncRNA MALAT1 contributes to lung cancer in African americans by modulating the tumor immune microenvironment. Cancers (Basel). 16:18762024. View Article : Google Scholar : PubMed/NCBI

153 

Alimohammadi M, Amani D, Adcock IM and Mortaz E: Dual role of mir-146a in non-small cell lung cancer progression: Molecular mechanisms and clinical potential. Cell Signal. 136:1121152025. View Article : Google Scholar : PubMed/NCBI

154 

Boucher A, Klopfenstein N, Hallas WM, Skibbe J, Appert A, Jang SH, Pulakanti K, Rao S, Cowden Dahl KD and Dahl R: The miR-23a~27a~24-2 microRNA cluster promotes inflammatory polarization of macrophages. J Immunol. 206:541–553. 2021. View Article : Google Scholar

155 

Zheng G and Chen Y, Wang Y, Huang Z, Chen W and Chen Y: circHIF1A/miR-486-5p/GRHL2 axis induces macrophage M2 polarization to accelerate lung adenocarcinoma progression. Int Immunopharmacol. 180:1166702026. View Article : Google Scholar : PubMed/NCBI

156 

Shang M, Ni L, Shan X, Cui Y, Hu P, Ji Z, Shen L, Zhang Y, Zhou J, Bing Chen, et al: MTHFD2 reprograms macrophage polarization by inhibiting PTEN. Cell Rep. 42:1124812023. View Article : Google Scholar : PubMed/NCBI

157 

Zhang Y, Jiang WL, Yang JY, Huang J, Kang G, Hu HB and Xie S: Downregulation of lysyl oxidase-like 4 LOXL4 by miR-135a-5p promotes lung cancer progression in vitro and in vivo. J Cell Physiol. 234:18679–18687. 2019. View Article : Google Scholar : PubMed/NCBI

158 

Shi H, Ji Y, Zhang D, Liu Y and Fang P: MiR-135a inhibits migration and invasion and regulates EMT-related marker genes by targeting KLF8 in lung cancer cells. Biochem Biophys Res Commun. 465:125–130. 2015. View Article : Google Scholar : PubMed/NCBI

159 

Abdelaal AM, Sohal IS, Iyer SG, Sudarshan K, Orellana EA, Ozcan KE, Dos Santos AP, Low PS and Kasinski AL: Selective targeting of chemically modified miR-34a to prostate cancer using a small molecule ligand and an endosomal escape agent. Mol Ther Nucleic Acids. 35:1021932024. View Article : Google Scholar : PubMed/NCBI

160 

Pandey S and Yadav P: Exploring the therapeutic potential of microRNAs: Targeted gene regulation strategies for enhanced cancer therapy. J Genet Eng Biotechnol. 23:1005562025. View Article : Google Scholar : PubMed/NCBI

161 

Gaponova S, Patutina O, Sen'kova A, Burakova E, Savin I, Markov A, Shmendel E, Maslov M, Stetsenko D, Vlassov V and Zenkova M: Single shot vs. Cocktail: A comparison of mono- and combinative application of miRNA-targeted mesyl oligonucleotides for efficient antitumor therapy. Cancers (Basel). 14:43962022. View Article : Google Scholar : PubMed/NCBI

162 

Sato Y, Matsui H, Yamamoto N, Sato R, Munakata T, Kohara M and Harashima H: Highly specific delivery of siRNA to hepatocytes circumvents endothelial cell-mediated lipid nanoparticle-associated toxicity leading to the safe and efficacious decrease in the hepatitis B virus. J Control Release. 266:216–225. 2017. View Article : Google Scholar : PubMed/NCBI

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Copy and paste a formatted citation
Spandidos Publications style
Yiming Z and Jie L: MicroRNA regulation of macrophage polarization in lung cancer: Regulatory networks and therapeutic potential (Review). Int J Oncol 69: 121, 2026.
APA
Yiming, Z., & Jie, L. (2026). MicroRNA regulation of macrophage polarization in lung cancer: Regulatory networks and therapeutic potential (Review). International Journal of Oncology, 69, 121. https://doi.org/10.3892/ijo.2026.5934
MLA
Yiming, Z., Jie, L."MicroRNA regulation of macrophage polarization in lung cancer: Regulatory networks and therapeutic potential (Review)". International Journal of Oncology 69.4 (2026): 121.
Chicago
Yiming, Z., Jie, L."MicroRNA regulation of macrophage polarization in lung cancer: Regulatory networks and therapeutic potential (Review)". International Journal of Oncology 69, no. 4 (2026): 121. https://doi.org/10.3892/ijo.2026.5934
Copy and paste a formatted citation
x
Spandidos Publications style
Yiming Z and Jie L: MicroRNA regulation of macrophage polarization in lung cancer: Regulatory networks and therapeutic potential (Review). Int J Oncol 69: 121, 2026.
APA
Yiming, Z., & Jie, L. (2026). MicroRNA regulation of macrophage polarization in lung cancer: Regulatory networks and therapeutic potential (Review). International Journal of Oncology, 69, 121. https://doi.org/10.3892/ijo.2026.5934
MLA
Yiming, Z., Jie, L."MicroRNA regulation of macrophage polarization in lung cancer: Regulatory networks and therapeutic potential (Review)". International Journal of Oncology 69.4 (2026): 121.
Chicago
Yiming, Z., Jie, L."MicroRNA regulation of macrophage polarization in lung cancer: Regulatory networks and therapeutic potential (Review)". International Journal of Oncology 69, no. 4 (2026): 121. https://doi.org/10.3892/ijo.2026.5934
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