REFERENCES
1. Chen YF, Luh F, Ho YS, Yen Y. Exosomes: a review of biologic function, diagnostic and targeted therapy applications, and clinical trials. J Biomed Sci. 2024;31:67.
2. Yan W, Jiang S. Immune cell-derived exosomes in the cancer-immunity cycle. Trends Cancer. 2020;6:506-17.
3. Zhang K, Cheng K. Stem cell-derived exosome versus stem cell therapy. Nat Rev Bioeng. 2023;1:608-9.
4. Doyle LM, Wang MZ. Overview of extracellular vesicles, their origin, composition, purpose, and methods for exosome isolation and analysis. Cells. 2019;8:727.
5. Jeppesen DK, Fenix AM, Franklin JL, et al. Reassessment of exosome composition. Cell. 2019;177:428-45.e18.
6. O’Brien K, Breyne K, Ughetto S, Laurent LC, Breakefield XO. RNA delivery by extracellular vesicles in mammalian cells and its applications. Nat Rev Mol Cell Biol. 2020;21:585-606.
7. Isaac R, Reis FCG, Ying W, Olefsky JM. Exosomes as mediators of intercellular crosstalk in metabolism. Cell Metab. 2021;33:1744-62.
8. Paskeh MDA, Entezari M, Mirzaei S, et al. Emerging role of exosomes in cancer progression and tumor microenvironment remodeling. J Hematol Oncol. 2022;15:83.
9. Melo SA, Luecke LB, Kahlert C, et al. Glypican-1 identifies cancer exosomes and detects early pancreatic cancer. Nature. 2015;523:177-82.
10. Wei P, Wu F, Kang B, et al. Plasma extracellular vesicles detected by single molecule array technology as a liquid biopsy for colorectal cancer. J Extracell Vesicles. 2020;9:1809765.
11. Sahoo S, Adamiak M, Mathiyalagan P, Kenneweg F, Kafert-Kasting S, Thum T. Therapeutic and diagnostic translation of extracellular vesicles in cardiovascular diseases: roadmap to the clinic. Circulation. 2021;143:1426-49.
12. Kugeratski FG, Hodge K, Lilla S, et al. Quantitative proteomics identifies the core proteome of exosomes with syntenin-1 as the highest abundant protein and a putative universal biomarker. Nat Cell Biol. 2021;23:631-41.
13. Chang W, Wang J. Exosomes and their noncoding RNA cargo are emerging as new modulators for diabetes mellitus. Cells. 2019;8:853.
14. Stuendl A, Kunadt M, Kruse N, et al. Induction of α-synuclein aggregate formation by CSF exosomes from patients with Parkinson’s disease and dementia with Lewy bodies. Brain. 2016;139:481-94.
16. Yu W, Hurley J, Roberts D, et al. Exosome-based liquid biopsies in cancer: opportunities and challenges. Ann Oncol. 2021;32:466-77.
17. Yu D, Li Y, Wang M, et al. Exosomes as a new frontier of cancer liquid biopsy. Mol Cancer. 2022;21:56.
18. Hoshino A, Kim HS, Bojmar L, et al. Extracellular vesicle and particle biomarkers define multiple human cancers. Cell. 2020;182:1044-61.e18.
19. Thomou T, Mori MA, Dreyfuss JM, et al. Adipose-derived circulating miRNAs regulate gene expression in other tissues. Nature. 2017;542:450-5.
20. Hoshino A, Costa-Silva B, Shen TL, et al. Tumour exosome integrins determine organotropic metastasis. Nature. 2015;527:329-35.
21. Gurung S, Perocheau D, Touramanidou L, Baruteau J. The exosome journey: from biogenesis to uptake and intracellular signalling. Cell Commun Signal. 2021;19:47.
22. Wang T, Zhang H. Exploring the roles and molecular mechanisms of RNA binding proteins in the sorting of noncoding RNAs into exosomes during tumor progression. J Adv Res. 2024;65:105-23.
23. Garcia-Martin R, Wang G, Brandão BB, et al. MicroRNA sequence codes for small extracellular vesicle release and cellular retention. Nature. 2022;601:446-51.
24. Arya SB, Collie SP, Parent CA. The ins-and-outs of exosome biogenesis, secretion, and internalization. Trends Cell Biol. 2024;34:90-108.
25. Tu H, Wang Z, Yuan Y, et al. The PripA-TbcrA complex-centered Rab GAP cascade facilitates macropinosome maturation in Dictyostelium. Nat Commun. 2022;13:1787.
26. Scott CC, Vacca F, Gruenberg J. Endosome maturation, transport and functions. Semin Cell Dev Biol. 2014;31:2-10.
27. Xu M, Ji J, Jin D, et al. The biogenesis and secretion of exosomes and multivesicular bodies (MVBs): intercellular shuttles and implications in human diseases. Genes Dis. 2023;10:1894-907.
28. Juan T, Fürthauer M. Biogenesis and function of ESCRT-dependent extracellular vesicles. Semin Cell Dev Biol. 2018;74:66-77.
29. Lee YJ, Shin KJ, Jang HJ, et al. GPR143 controls ESCRT-dependent exosome biogenesis and promotes cancer metastasis. Dev Cell. 2023;58:320-34.e8.
30. Vietri M, Radulovic M, Stenmark H. The many functions of ESCRTs. Nat Rev Mol Cell Biol. 2020;21:25-42.
31. Mercier V, Larios J, Molinard G, et al. Endosomal membrane tension regulates ESCRT-III-dependent intra-lumenal vesicle formation. Nat Cell Biol. 2020;22:947-59.
32. Banjade S, Zhu L, Jorgensen JR, Suzuki SW, Emr SD. Recruitment and organization of ESCRT-0 and ubiquitinated cargo via condensation. Sci Adv. 2022;8:eabm5149.
33. Babst M. MVB vesicle formation: ESCRT-dependent, ESCRT-independent and everything in between. Curr Opin Cell Biol. 2011;23:452-7.
34. Li M, Li S, Du C, et al. Exosomes from different cells: characteristics, modifications, and therapeutic applications. Eur J Med Chem. 2020;207:112784.
35. Han QF, Li WJ, Hu KS, et al. Exosome biogenesis: machinery, regulation, and therapeutic implications in cancer. Mol Cancer. 2022;21:207.
36. Song L, Tang S, Han X, et al. KIBRA controls exosome secretion via inhibiting the proteasomal degradation of Rab27a. Nat Commun. 2019;10:1639.
37. Jahn R, Cafiso DC, Tamm LK. Mechanisms of SNARE proteins in membrane fusion. Nat Rev Mol Cell Biol. 2024;25:101-18.
38. Liu H, Dang R, Zhang W, Hong J, Li X. SNARE proteins: core engines of membrane fusion in cancer. Biochim Biophys Acta Rev Cancer. 2024:189148.
39. Messenger SW, Woo SS, Sun Z, Martin TFJ. A Ca2+-stimulated exosome release pathway in cancer cells is regulated by Munc13-4. J Cell Biol. 2018;217:2877-90.
40. Zhang L, Pan J, Wang M, et al. Chronic stress-induced and tumor derived SP1+ exosomes polarizing IL-1β+ neutrophils to increase lung metastasis of breast cancer. Adv Sci. 2025;12:e2310266.
41. Guo W, Qiao T, Dong B, Li T, Liu Q, Xu X. The effect of hypoxia-induced exosomes on anti-tumor immunity and its implication for immunotherapy. Front Immunol. 2022;13:915985.
42. Kumar A, Deep G. Hypoxia in tumor microenvironment regulates exosome biogenesis: molecular mechanisms and translational opportunities. Cancer Lett. 2020;479:23-30.
43. Wang X, Wu R, Zhai P, et al. Hypoxia promotes EV secretion by impairing lysosomal homeostasis in HNSCC through negative regulation of ATP6V1A by HIF-1α. J Extracell Vesicles. 2023;12:e12310.
44. Jiang H, Zhao H, Zhang M, et al. Hypoxia induced changes of exosome cargo and subsequent biological effects. Front Immunol. 2022;13:824188.
45. He G, Peng X, Wei S, et al. Exosomes in the hypoxic TME: from release, uptake and biofunctions to clinical applications. Mol Cancer. 2022;21:19.
46. Kalluri R, LeBleu VS. The biology, function, and biomedical applications of exosomes. Science. 2020;367:eaau6977.
47. Nail HM, Chiu CC, Leung CH, Ahmed MMM, Wang HD. Exosomal miRNA-mediated intercellular communications and immunomodulatory effects in tumor microenvironments. J Biomed Sci. 2023;30:69.
48. Zhang Z, Zou Y, Song C, et al. Advances in the study of exosomes in cardiovascular diseases. J Adv Res. 2024;66:133-53.
49. Essola JM, Zhang M, Yang H, et al. Exosome regulation of immune response mechanism: pros and cons in immunotherapy. Bioact Mater. 2024;32:124-46.
50. Beninson LA, Fleshner M. Exosomes: an emerging factor in stress-induced immunomodulation. Semin Immunol. 2014;26:394-401.
51. Jeon JS, Kim E, Bae YU, et al. microRNA in extracellular vesicles released by damaged podocytes promote apoptosis of renal tubular epithelial cells. Cells. 2020;9:1409.
52. Grange C, Bussolati B. Extracellular vesicles in kidney disease. Nat Rev Nephrol. 2022;18:499-513.
53. Lin SW, Tsai JC, Shyong YJ. Drug delivery of extracellular vesicles: preparation, delivery strategies and applications. Int J Pharm. 2023;642:123185.
54. Lu W, Zhang J, Wu Y, Sun W, Jiang Z, Luo X. Engineered NF-κB siRNA-encapsulating exosomes as a modality for therapy of skin lesions. Front Immunol. 2023;14:1109381.
55. Rong Y, Wang Z, Tang P, et al. Engineered extracellular vesicles for delivery of siRNA promoting targeted repair of traumatic spinal cord injury. Bioact Mater. 2023;23:328-42.
56. Sun Y, Zhao Y, Ni X, et al.
57. Yong T, Wei Z, Gan L, Yang X. Extracellular-vesicle-based drug delivery systems for enhanced antitumor therapies through modulating the cancer-immunity cycle. Adv Mater. 2022;34:e2201054.
58. Whitley JA, Cai H. Engineering extracellular vesicles to deliver CRISPR ribonucleoprotein for gene editing. J Extracell Vesicles. 2023;12:e12343.
59. Liu X, Cao Z, Wang W, et al. Engineered extracellular vesicle-delivered CRISPR/Cas9 for radiotherapy sensitization of glioblastoma. ACS Nano. 2023;17:16432-47.
60. Gebert LFR, MacRae IJ. Regulation of microRNA function in animals. Nat Rev Mol Cell Biol. 2019;20:21-37.
61. Rhim J, Baek W, Seo Y, Kim JH. From molecular mechanisms to therapeutics: understanding microRNA-21 in cancer. Cells. 2022;11:2791.
62. Kim H, Lee YY, Kim VN. The biogenesis and regulation of animal microRNAs. Nat Rev Mol Cell Biol. 2025;26:276-96.
63. Shang R, Lee S, Senavirathne G, Lai EC. microRNAs in action: biogenesis, function and regulation. Nat Rev Genet. 2023;24:816-33.
64. Wu K, He J, Pu W, Peng Y. The role of exportin-5 in microRNA biogenesis and cancer. Genomics Proteomics Bioinformatics. 2018;16:120-6.
65. Iwakawa HO, Tomari Y. Life of RISC: formation, action, and degradation of RNA-induced silencing complex. Mol Cell. 2022;82:30-43.
66. Jiang X, Liu B, Nie Z, et al. The role of m6A modification in the biological functions and diseases. Signal Transduct Target Ther. 2021;6:74.
67. Liu R, Zhong Y, Chen R, et al. m6A reader hnRNPA2B1 drives multiple myeloma osteolytic bone disease. Theranostics. 2022;12:7760-74.
68. Samsonova A, El Hage K, Desforges B, et al. Lin28, a major translation reprogramming factor, gains access to YB-1-packaged mRNA through its cold-shock domain. Commun Biol. 2021;4:359.
69. Davis BN, Hilyard AC, Lagna G, Hata A. SMAD proteins control DROSHA-mediated microRNA maturation. Nature. 2008;454:56-61.
70. Jiang F, Tang X, Tang C, et al. HNRNPA2B1 promotes multiple myeloma progression by increasing AKT3 expression via m6A-dependent stabilization of ILF3 mRNA. J Hematol Oncol. 2021;14:54.
71. Min KW, Jo MH, Song M, et al. Mature microRNA-binding protein QKI promotes microRNA-mediated gene silencing. RNA Biol. 2024;21:1-15.
72. Chen X, Liu Y, Xu C, et al. QKI is a critical pre-mRNA alternative splicing regulator of cardiac myofibrillogenesis and contractile function. Nat Commun. 2021;12:89.
73. Wang F, Song W, Zhao H, et al. The RNA-binding protein QKI5 regulates primary miR-124-1 processing via a distal RNA motif during erythropoiesis. Cell Res. 2017;27:416-39.
74. Jia Y, Wei Y. Modulators of microRNA function in the immune system. Int J Mol Sci. 2020;21:2357.
75. Taganov KD, Boldin MP, Chang KJ, Baltimore D. NF-kappaB-dependent induction of microRNA miR-146, an inhibitor targeted to signaling proteins of innate immune responses. Proc Natl Acad Sci U S A. 2006;103:12481-6.
76. Mahesh G, Biswas R. MicroRNA-155: a master regulator of inflammation. J Interferon Cytokine Res. 2019;39:321-30.
77. Bautista-Sánchez D, Arriaga-Canon C, Pedroza-Torres A, et al. The promising role of miR-21 as a cancer biomarker and its importance in RNA-based therapeutics. Mol Ther Nucleic Acids. 2020;20:409-20.
78. Hashemi M, Mirdamadi MSA, Talebi Y, et al. Pre-clinical and clinical importance of miR-21 in human cancers: tumorigenesis, therapy response, delivery approaches and targeting agents. Pharmacol Res. 2023;187:106568.
79. Gao S, Gao H, Dai L, et al. miR-126 regulates angiogenesis in myocardial ischemia by targeting HIF-1α. Exp Cell Res. 2021;409:112925.
80. Morita S, Horii T, Kimura M, Ochiya T, Tajima S, Hatada I. miR-29 represses the activities of DNA methyltransferases and DNA demethylases. Int J Mol Sci. 2013;14:14647-58.
81. Mańka R, Janas P, Sapoń K, Janas T, Janas T. Role of RNA motifs in RNA interaction with membrane lipid rafts: implications for therapeutic applications of exosomal RNAs. Int J Mol Sci. 2021;22:9416.
82. Pust S, Brech A, Wegner CS, Stenmark H, Haglund K. Vesicle-mediated transport of ALIX and ESCRT-III to the intercellular bridge during cytokinesis. Cell Mol Life Sci. 2023;80:235.
83. Lee H, Li C, Zhang Y, Zhang D, Otterbein LE, Jin Y. Caveolin-1 selectively regulates microRNA sorting into microvesicles after noxious stimuli. J Exp Med. 2019;216:2202-20.
84. Zhuang W, Liu C, Hong Y, et al. Tumor-suppressive miR-4732-3p is sorted into fucosylated exosome by hnRNPK to avoid the inhibition of lung cancer progression. J Exp Clin Cancer Res. 2024;43:123.
85. Li C, Qin F, Wang W, et al. hnRNPA2B1-mediated extracellular vesicles sorting of miR-122-5p potentially promotes lung cancer progression. Int J Mol Sci. 2021;22:12866.
86. Fabbiano F, Corsi J, Gurrieri E, Trevisan C, Notarangelo M, D’Agostino VG. RNA packaging into extracellular vesicles: an orchestra of RNA-binding proteins? J Extracell Vesicles. 2020;10:e12043.
87. Villarroya-Beltri C, Gutiérrez-Vázquez C, Sánchez-Cabo F, et al. Sumoylated hnRNPA2B1 controls the sorting of miRNAs into exosomes through binding to specific motifs. Nat Commun. 2013;4:2980.
88. Liu Z, Xin B, Zhang N, et al. LSD1 modulates the bone metastasis of breast cancer cells through hnRNPA2B1-mediated sorting of exosomal miRNAs. Cell Death Discov. 2024;10:115.
89. He W, Belsham DD. RNA-binding protein motifs predict microRNA secretion and cellular retention in hypothalamic and other cell types. Biomedicines. 2024;12:857.
90. Zhou X, Hong Y, Liu Y, et al. Intervening in hnRNPA2B1-mediated exosomal transfer of tumor-suppressive miR-184-3p for tumor microenvironment regulation and cancer therapy. J Nanobiotechnology. 2023;21:422.
91. Balaguer N, Moreno I, Herrero M, González M, Simón C, Vilella F. Heterogeneous nuclear ribonucleoprotein C1 may control miR-30d levels in endometrial exosomes affecting early embryo implantation. Mol Hum Reprod. 2018;24:411-25.
92. Jiang NX, Li XL. The complicated effects of extracellular vesicles and their cargos on embryo implantation. Front Endocrinol. 2021;12:681266.
93. Liao Z, Tong B, Zhang X, et al. Selective cargo sorting in stem cell-derived small extracellular vesicles: impact on therapeutic efficacy for intervertebral disc degeneration. Clin Transl Med. 2023;13:e1494.
94. Ye T, Zhong L, Ye X, Liu J, Li L, Yi H. miR-221-3p and miR-222-3p regulate the SOCS3/STAT3 signaling pathway to downregulate the expression of NIS and reduce radiosensitivity in thyroid cancer. Exp Ther Med. 2021;21:652.
95. Li S, Cai X, Chen L, et al. Inhibition of hepatocellular carcinoma growth via modulation of the miR-221/SOX11 axis by curcumin and berberine. PeerJ. 2023;11:e16593.
96. Wu X, Wang X, Yin Y, Zhu L, Zhang F, Yang J. Investigation of the role of miR-221 in diabetic peripheral neuropathy and related molecular mechanisms. Adv Clin Exp Med. 2021;30:623-32.
97. Zietzer A, Hosen MR, Wang H, et al. The RNA-binding protein hnRNPU regulates the sorting of microRNA-30c-5p into large extracellular vesicles. J Extracell Vesicles. 2020;9:1786967.
98. Zhao Y, Li F, Zhang X, Liu A, Qi J, Cui H, Zhao P. MicroRNA-194 acts as a prognostic marker and inhibits proliferation in hepatocellular carcinoma by targeting MAP4K4. Int J Clin Exp Pathol. 2015;8:12446-54.
99. Santangelo L, Giurato G, Cicchini C, et al. The RNA-binding protein SYNCRIP is a component of the hepatocyte exosomal machinery controlling microRNA sorting. Cell Rep. 2016;17:799-808.
100. Hobor F, Dallmann A, Ball NJ, et al. A cryptic RNA-binding domain mediates Syncrip recognition and exosomal partitioning of miRNA targets. Nat Commun. 2018;9:831.
101. Robinson H, Ruelcke JE, Lewis A, et al. Caveolin-1-driven membrane remodelling regulates hnRNPK-mediated exosomal microRNA sorting in cancer. Clin Transl Med. 2021;11:e381.
102. Liu D, Liu F, Li Z, et al. HNRNPA1-mediated exosomal sorting of miR-483-5p out of renal tubular epithelial cells promotes the progression of diabetic nephropathy-induced renal interstitial fibrosis. Cell Death Dis. 2021;12:255.
103. Dou R, Liu K, Yang C, et al. EMT-cancer cells-derived exosomal miR-27b-3p promotes circulating tumour cells-mediated metastasis by modulating vascular permeability in colorectal cancer. Clin Transl Med. 2021;11:e595.
104. Qin X, Guo H, Wang X, et al. Exosomal miR-196a derived from cancer-associated fibroblasts confers cisplatin resistance in head and neck cancer through targeting CDKN1B and ING5. Genome Biol. 2019;20:12.
105. Zhang H, Deng T, Liu R, et al. CAF secreted miR-522 suppresses ferroptosis and promotes acquired chemo-resistance in gastric cancer. Mol Cancer. 2020;19:43.
106. Qiu W, Guo X, Li B, et al. Exosomal miR-1246 from glioma patient body fluids drives the differentiation and activation of myeloid-derived suppressor cells. Mol Ther. 2021;29:3449-64.
107. Gao X, Wan Z, Wei M, et al. Chronic myelogenous leukemia cells remodel the bone marrow niche via exosome-mediated transfer of miR-320. Theranostics. 2019;9:5642-56.
108. Ma L, Singh J, Schekman R. Two RNA-binding proteins mediate the sorting of miR223 from mitochondria into exosomes. Elife. 2023;12:e85878.
109. Shurtleff MJ, Temoche-Diaz MM, Karfilis KV, Ri S, Schekman R. Y-box protein 1 is required to sort microRNAs into exosomes in cells and in a cell-free reaction. Elife. 2016;5:e19276.
110. Liu XM, Ma L, Schekman R. Selective sorting of microRNAs into exosomes by phase-separated YBX1 condensates. Elife. 2021;10:e71982.
111. Lin F, Zeng Z, Song Y, et al. YBX-1 mediated sorting of miR-133 into hypoxia/reoxygenation-induced EPC-derived exosomes to increase fibroblast angiogenesis and MEndoT. Stem Cell Res Ther. 2019;10:263.
112. Kossinova OA, Gopanenko AV, Tamkovich SN, et al. Cytosolic YB-1 and NSUN2 are the only proteins recognizing specific motifs present in mRNAs enriched in exosomes. Biochim Biophys Acta Proteins Proteom. 2017;1865:664-73.
113. Guo J, Zhao J, Xu Q, Huang D. MEX3C as a potential target for hepatocellular carcinoma drug and immunity: combined therapy with Lenvatinib. BMC Cancer. 2023;23:967.
114. Lu P, Li H, Li N, et al. MEX3C interacts with adaptor-related protein complex 2 and involves in miR-451a exosomal sorting. PLoS One. 2017;12:e0185992.
115. Majumder M, Chakraborty P, Mohan S, Mehrotra S, Palanisamy V. HuR as a molecular target for cancer therapeutics and immune-related disorders. Adv Drug Deliv Rev. 2022;188:114442.
116. Mukherjee K, Ghoshal B, Ghosh S, et al. Reversible HuR-microRNA binding controls extracellular export of miR-122 and augments stress response. EMBO Rep. 2016;17:1184-203.
117. Shi Y, Wang Z, Zhu X, et al. Exosomal miR-1246 in serum as a potential biomarker for early diagnosis of gastric cancer. Int J Clin Oncol. 2020;25:89-99.
118. Torii C, Maishi N, Kawamoto T, et al. miRNA-1246 in extracellular vesicles secreted from metastatic tumor induces drug resistance in tumor endothelial cells. Sci Rep. 2021;11:13502.
119. Geekiyanage H, Rayatpisheh S, Wohlschlegel JA, Brown R Jr, 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 U S A. 2020;117:24213-23.
120. Marzec M. New insights into the function of mammalian Argonaute2. PLoS Genet. 2020;16:e1009058.
121. Deshmukh P, Markande S, Fandade V, Ramtirtha Y, Madhusudhan MS, Joseph J. The miRISC component AGO2 has multiple binding sites for Nup358 SUMO-interacting motif. Biochem Biophys Res Commun. 2021;556:45-52.
122. Müller M, Fazi F, Ciaudo C. Argonaute proteins: from structure to function in development and pathological cell fate determination. Front Cell Dev Biol. 2019;7:360.
123. Weaver AM, Patton JG. Argonautes in extracellular vesicles: artifact or selected cargo? Cancer Res. 2020;80:379-81.
124. Zhang X, Niu D, Carbonell A, et al. ARGONAUTE PIWI domain and microRNA duplex structure regulate small RNA sorting in Arabidopsis. Nat Commun. 2014;5:5468.
125. Ye Z, Jin H, Qian Q. Argonaute 2: a novel rising star in cancer research. J Cancer. 2015;6:877-82.
126. McKenzie AJ, Hoshino D, Hong NH, et al. KRAS-MEK signaling controls Ago2 sorting into exosomes. Cell Rep. 2016;15:978-87.
127. Li Y, Estep JA, Karginov FV. Transcriptome-wide identification and validation of interactions between the miRNA machinery and HuR on mRNA targets. J Mol Biol. 2018;430:285-96.
128. Tapparo M, Pomatto MAC, Deregibus MC, et al. Serum derived extracellular vesicles mediated delivery of synthetic miRNAs in human endothelial cells. Front Mol Biosci. 2021;8:636587.
129. Pérez-Boza J, Boeckx A, Lion M, Dequiedt F, Struman I. hnRNPA2B1 inhibits the exosomal export of miR-503 in endothelial cells. Cell Mol Life Sci. 2020;77:4413-28.
130. Vanessa L, Cristina PRX, Diana S, et al. ALIX protein analysis: storage temperature may impair results. J Mol Clin Med. 2019;2:29.
131. Teng Y, Ren Y, Hu X, et al. MVP-mediated exosomal sorting of miR-193a promotes colon cancer progression. Nat Commun. 2017;8:14448.
132. Telford BJ, Yahyanejad S, de Gunst T, et al. Multi-modal effects of 1B3, a novel synthetic miR-193a-3p mimic, support strong potential for therapeutic intervention in oncology. Oncotarget. 2021;12:422-39.
133. Wang F, Yuan Y, Yang P, Li X. Extracellular vesicles-mediated transfer of miR-208a/b exaggerate hypoxia/reoxygenation injury in cardiomyocytes by reducing QKI expression. Mol Cell Biochem. 2017;431:187-95.
134. Sharma M, Sharma S, Alawada A. Understanding the binding specificities of mRNA targets by the mammalian Quaking protein. Nucleic Acids Res. 2019;47:10564-79.
135. Zhao Z, Qing Y, Dong L, et al. QKI shuttles internal m7G-modified transcripts into stress granules and modulates mRNA metabolism. Cell. 2023;186:3208-26.e27.
136. Gissi C, Radeghieri A, Antonetti Lamorgese Passeri C, et al. Extracellular vesicles from rat-bone-marrow mesenchymal stromal/stem cells improve tendon repair in rat Achilles tendon injury model in dose-dependent manner: a pilot study. PLoS One. 2020;15:e0229914.
137. Xu C, Xie N, Su Y, et al. HnRNP F/H associate with hTERC and telomerase holoenzyme to modulate telomerase function and promote cell proliferation. Cell Death Differ. 2020;27:1998-2013.
138. Temoche-Diaz MM, Shurtleff MJ, Nottingham RM, et al. Distinct mechanisms of microRNA sorting into cancer cell-derived extracellular vesicle subtypes. Elife. 2019;8:e47544.
139. Xu YF, Xu X, Gin A, et al. SRSF1 regulates exosome microRNA enrichment in human cancer cells. Cell Commun Signal. 2020;18:130.
140. Edbauer D, Neilson JR, Foster KA, et al. Regulation of synaptic structure and function by FMRP-associated microRNAs miR-125b and miR-132. Neuron. 2010;65:373-84.
141. Wozniak AL, Adams A, King KE, et al. The RNA binding protein FMR1 controls selective exosomal miRNA cargo loading during inflammation. J Cell Biol. 2020;219:e201912074.
142. Liu Q, Li D, Pan X, Liang Y. Targeted therapy using engineered extracellular vesicles: principles and strategies for membrane modification. J Nanobiotechnology. 2023;21:334.
143. Zhang M, Hu S, Liu L, et al. Engineered exosomes from different sources for cancer-targeted therapy. Signal Transduct Target Ther. 2023;8:124.
144. Berggreen AH, Petersen JL, Lin L, Benabdellah K, Luo Y. CRISPR delivery with extracellular vesicles: promises and challenges. J Extracell Biol. 2023;2:e111.
145. Kenari A, Cheng L, Hill AF. Methods for loading therapeutics into extracellular vesicles and generating extracellular vesicles mimetic-nanovesicles. Methods. 2020;177:103-13.
146. Erana-Perez Z, Igartua M, Santos-Vizcaino E, Hernandez RM. Genetically engineered loaded extracellular vesicles for drug delivery. Trends Pharmacol Sci. 2024;45:350-65.
147. Simon L, Constanzo J, Terraza-Aguirre C, et al. Surface modification of extracellular vesicles with polyoxazolines to enhance their plasma stability and tumor accumulation. Biomaterials. 2025;313:122748.
148. Suk JS, Xu Q, Kim N, Hanes J, Ensign LM. PEGylation as a strategy for improving nanoparticle-based drug and gene delivery. Adv Drug Deliv Rev. 2016;99:28-51.
149. Hu J, Liu WF, Zhang XY, et al. Synthetic miR-26a mimics delivered by tumor exosomes repress hepatocellular carcinoma through downregulating lymphoid enhancer factor 1. Hepatol Int. 2023;17:1265-78.
150. Reshke R, Taylor JA, Savard A, et al. Reduction of the therapeutic dose of silencing RNA by packaging it in extracellular vesicles via a pre-microRNA backbone. Nat Biomed Eng. 2020;4:52-68.
151. Li Q, Sun T, Liu S, Kou Y, Han N. Overview of biomembrane-derived nanoparticles as emerging drug delivery systems for treating bone aging diseases. Int J Pharm. 2025;681:125832.
152. Xing Y, Cheng Z, Wang R, Lv C, James TD, Yu F. Analysis of extracellular vesicles as emerging theranostic nanoplatforms. Coord Chem Rev. 2020;424:213506.
153. Cheng L, Hill AF. Therapeutically harnessing extracellular vesicles. Nat Rev Drug Discov. 2022;21:379-99.
