REFERENCES
1. Deng W, Zhu X, Li H, Hu P, Qian K, Liu F. Lung tissue extracellular vesicles-mediated delivery of miR-128-3p as a novel mechanism of acute lung inflammation. Int J Nanomedicine. 2025;20:4831-48.
2. Askenase PW. Exosomes provide unappreciated carrier effects that assist transfers of their miRNAs to targeted cells; I. They are ‘The Elephant in the Room’. RNA Biol. 2021;18:2038-53.
3. De Rosa G, Russo C, Garavelli S, et al. MicroRNA-142-3p shuttling in extracellular vesicles marks regulatory T cell dysfunction in multiple sclerosis. Sci Transl Med. 2025;17:eadl1698.
4. Cuomo-Haymour N, Bergamini G, Russo G, et al. Differential expression of serum extracellular vesicle miRNAs in multiple sclerosis: disease-stage specificity and relevance to pathophysiology. Int J Mol Sci. 2022;23:1664.
5. Ebrahimkhani S, Beadnall HN, Wang C, et al. Serum exosome microRNAs predict multiple sclerosis disease activity after fingolimod treatment. Mol Neurobiol. 2020;57:1245-58.
6. Ebrahimkhani S, Vafaee F, Young PE, et al. Exosomal microRNA signatures in multiple sclerosis reflect disease status. Sci Rep. 2017;7:14293.
7. Manna I, Iaccino E, Dattilo V, et al. Exosome-associated miRNA profile as a prognostic tool for therapy response monitoring in multiple sclerosis patients. FASEB J. 2018;32:4241-6.
8. Mohammadinasr M, Montazersaheb S, Hosseini V, et al. Epstein-Barr virus-encoded BART9 and BART15 miRNAs are elevated in exosomes of cerebrospinal fluid from relapsing-remitting multiple sclerosis patients. Cytokine. 2024;179:156624.
9. Mohammadinasr M, Montazersaheb S, Molavi O, et al. Multiplex analysis of cerebrospinal fluid and serum exosomes microRNAs of untreated relapsing remitting multiple sclerosis (RRMS) and proposing noninvasive diagnostic biomarkers. Neuromolecular Med. 2023;25:402-14.
10. Scaroni F, Visconte C, Serpente M, et al. miR-150-5p and let-7b-5p in blood myeloid extracellular vesicles track cognitive symptoms in patients with multiple sclerosis. Cells. 2022;11:1551.
11. Shi Z, Sun H, Tian X, et al. Extracellular vesicles containing miR-181a-5p as a novel therapy for experimental autoimmune encephalomyelitis-induced demyelination. Int Immunopharmacol. 2024;135:112326.
12. Selmaj I, Cichalewska M, Namiecinska M, et al. Global exosome transcriptome profiling reveals biomarkers for multiple sclerosis. Ann Neurol. 2017;81:703-17.
13. Torres-Iglesias G, López-Molina M, Ayala-Suárez R, et al. Extracellular vesicle-derived microRNAs as a biomarker for therapeutic response in multiple sclerosis. Neurol Neuroimmunol Neuroinflamm. 2025;12:e200420.
14. Kimura K, Hohjoh H, Fukuoka M, et al. Circulating exosomes suppress the induction of regulatory T cells via let-7i in multiple sclerosis. Nat Commun. 2018;9:17.
15. Hvalkof VH, Hansen MB, El Mahdaoui S, et al. Cerebrospinal fluid exosomal Epstein-Barr virus microRNAs in multiple sclerosis and effects of disease modifying therapies. Clin Immunol. 2025;281:110608.
16. Hvalkof VH, Olsson AGS, Gustavsen S, et al. Profiling of Epstein-Barr virus microRNAs in whole blood and exosomes in multiple sclerosis. Immunol Invest. 2025;54:1524-41.
17. Wasilewska K, Dziedzic A, Anandan S, et al. Extracellular vesicle-derived miR-760 as a novel promising candidate biomarker differentiating stable RRMS from SPMS. Sci Rep. 2026;16:5208.
18. Dendrou CA, Fugger L, Friese MA. Immunopathology of multiple sclerosis. Nat Rev Immunol. 2015;15:545-58.
19. Balusu S, Van Wonterghem E, De Rycke R, et al. Identification of a novel mechanism of blood-brain communication during peripheral inflammation via choroid plexus-derived extracellular vesicles. EMBO Mol Med. 2016;8:1162-83.
20. Matsumoto J, Stewart T, Sheng L, et al. Transmission of α-synuclein-containing erythrocyte-derived extracellular vesicles across the blood-brain barrier via adsorptive mediated transcytosis: another mechanism for initiation and progression of Parkinson’s disease? Acta Neuropathol Commun. 2017;5:71.
21. Albanese M, Chen YA, Hüls C, et al. MicroRNAs are minor constituents of extracellular vesicles that are rarely delivered to target cells. PLoS Genet. 2021;17:e1009951.
22. Dalvi P, Sun B, Tang N, Pulliam L. Immune activated monocyte exosomes alter microRNAs in brain endothelial cells and initiate an inflammatory response through the TLR4/MyD88 pathway. Sci Rep. 2017;7:9954.
23. Chaudhuri AD, Dastgheyb RM, Yoo SW, et al. TNFα and IL-1β modify the miRNA cargo of astrocyte shed extracellular vesicles to regulate neurotrophic signaling in neurons. Cell Death Dis. 2018;9:363.
24. Fernández-Messina L, Rodríguez-Galán A, de Yébenes VG, et al. Transfer of extracellular vesicle-microRNA controls germinal center reaction and antibody production. EMBO Rep. 2020;21:e48925.
25. Welsh JA, Goberdhan DCI, O’Driscoll L, et al.; MISEV Consortium. Minimal information for studies of extracellular vesicles (MISEV2023): from basic to advanced approaches. J Extracell Vesicles. 2024;13:e12404.
26. Valadi H, Ekström K, Bossios A, Sjöstrand M, Lee JJ, Lötvall JO. Exosome-mediated transfer of mRNAs and microRNAs is a novel mechanism of genetic exchange between cells. Nat Cell Biol. 2007;9:654-9.
27. Pegtel DM, Cosmopoulos K, Thorley-Lawson DA, et al. Functional delivery of viral miRNAs via exosomes. Proc Natl Acad Sci U S A. 2010;107:6328-33.
28. Chevillet JR, Kang Q, Ruf IK, et al. Quantitative and stoichiometric analysis of the microRNA content of exosomes. Proc Natl Acad Sci U S A. 2014;111:14888-93.
29. Liu XM, Halushka MK. Beyond the bubble: a debate on microRNA sorting into extracellular vesicles. Lab Invest. 2025;105:102206.
30. Groot M, Lee H. Sorting mechanisms for microRNAs into extracellular vesicles and their associated diseases. Cells. 2020;9:1044.
31. Qu M, Zhang J, Yu X, Jiang L, Zhu H, Zhang X. The RNA binding protein hnRNP A/B controls exosomal miR-27a-5p loading during Salmonella infection. Poult Sci. 2025;104:105482.
32. 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.
33. Lino M, Garcia-Martin R, Muñoz VR, et al. Multi-step regulation of microRNA expression and secretion into small extracellular vesicles by insulin. Cell Rep. 2024;43:114491.
34. 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.
35. Wu F, Pang H, Li F, Hua M, Song C, Tang J. Progress in cancer research on the regulator of phagocytosis CD47, which determines the fate of tumor cells (Review). Oncol Lett. 2024;27:256.
36. Kamerkar S, LeBleu VS, Sugimoto H, et al. Exosomes facilitate therapeutic targeting of oncogenic KRAS in pancreatic cancer. Nature. 2017;546:498-503.
37. Matsumoto A, Takahashi Y, Nishikawa M, et al. Role of phosphatidylserine-derived negative surface charges in the recognition and uptake of intravenously injected B16BL6-derived exosomes by macrophages. J Pharm Sci. 2017;106:168-75.
38. Wang YZ, Castillon CCM, Gebis KK, et al. Notch receptor-ligand binding facilitates extracellular vesicle-mediated neuron-to-neuron communication. Cell Rep. 2024;43:113680.
39. Purushothaman A, Bandari SK, Liu J, Mobley JA, Brown EE, Sanderson RD. Fibronectin on the surface of myeloma cell-derived exosomes mediates exosome-cell interactions. J Biol Chem. 2016;291:1652-63.
40. Elsharkasy OM, de Voogt WS, Tognoli ML, et al. Integrin beta 1 and fibronectin mediate extracellular vesicle uptake and functional RNA delivery. J Biol Chem. 2025;301:108305.
41. Joshi BS, de Beer MA, Giepmans BNG, Zuhorn IS. Endocytosis of extracellular vesicles and release of their cargo from endosomes. ACS Nano. 2020;14:4444-55.
42. Ribovski L, Joshi B, Gao J, Zuhorn I. Breaking free: endocytosis and endosomal escape of extracellular vesicles. Extracell Vesicles Circ Nucl Acids. 2023;4:283-305.
43. Squadrito ML, Baer C, Burdet F, et al. Endogenous RNAs modulate microRNA sorting to exosomes and transfer to acceptor cells. Cell Rep. 2014;8:1432-46.
44. Solvik TA, Nguyen TA, Tony Lin YH, et al. Secretory autophagy maintains proteostasis upon lysosome inhibition. J Cell Biol. 2022;221:e202110151.
45. Takahashi A, Okada R, Nagao K, et al. Exosomes maintain cellular homeostasis by excreting harmful DNA from cells. Nat Commun. 2017;8:15287.
46. Drysch M, Fiedler A, Schmidt SV, et al. Systemic downregulation of EV-associated MiRNAs following remote ischemic preconditioning. Sci Rep. 2025;15:43657.
47. Marocco F, Garbo S, Montaldo C, et al. Negative regulation of miRNA sorting into EVs is mediated by the capacity of RBP PCBP2 to impair the SYNCRIP-dependent miRNA loading. Elife. 2025;14:RP105017.
48. 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.
49. Lerussi G, Villagrasa-Araya V, Moltó-Abad M, et al. Extracellular vesicles as tools for crossing the blood-brain barrier to treat lysosomal storage diseases. Life. 2025;15:70.
50. Denis HL, de Rus Jacquet A, Alpaugh M, et al. Erythrocyte-derived extracellular vesicles transcytose across the blood-brain barrier to induce Parkinson's disease-like neurodegeneration. Fluids Barriers CNS. 2025;22:38.
51. Morad G, Carman CV, Hagedorn EJ, et al. Tumor-derived extracellular vesicles breach the intact blood-brain barrier via transcytosis. ACS Nano. 2019;13:13853-65.
52. Chen CC, Liu L, Ma F, et al. Elucidation of exosome migration across the blood-brain barrier model in vitro. Cell Mol Bioeng. 2016;9:509-29.
53. Gayen M, Bhomia M, Balakathiresan N, Knollmann-Ritschel B. Exosomal microRNAs released by activated astrocytes as potential neuroinflammatory biomarkers. Int J Mol Sci. 2020;21:2312.
54. Hollingworth BYA, Pallier PN, Jenkins SI, Chen R. Hypoxic neuroinflammation in the pathogenesis of multiple sclerosis. Brain Sci. 2025;15:248.
55. Cui JG, Li YY, Zhao Y, Bhattacharjee S, Lukiw WJ. Differential regulation of interleukin-1 receptor-associated kinase-1 (IRAK-1) and IRAK-2 by microRNA-146a and NF-kappaB in stressed human astroglial cells and in Alzheimer disease. J Biol Chem. 2010;285:38951-60.
56. Pegoretti V, Swanson KA, Bethea JR, Probert L, Eisel ULM, Fischer R. Inflammation and oxidative stress in multiple sclerosis: consequences for therapy development. Oxid Med Cell Longev. 2020;2020:7191080.
57. Ayeldeen G, Shaker OG, Gomaa M, et al. Association of epistatic effects of lncRNA GAS5, miR-146a, IRAK-1, and miR-155 genetic variants with multiple sclerosis risk and severity. Mol Neurobiol. 2025;62:10742-64.
58. Kaskow BJ, Baecher-Allan C. Effector T cells in multiple sclerosis. Cold Spring Harb Perspect Med. 2018;8:a029025.
59. Morris G, Reschke CR, Henshall DC. Targeting microRNA-134 for seizure control and disease modification in epilepsy. EBioMedicine. 2019;45:646-54.
60. Frendo ME, da Silva A, Phan KD, Riche S, Butler SJ. The cofilin/limk1 pathway controls the growth rate of both developing and regenerating motor axons. J Neurosci. 2019;39:9316-27.
61. Baby N, Alagappan N, Dheen ST, Sajikumar S. MicroRNA-134-5p inhibition rescues long-term plasticity and synaptic tagging/capture in an Aβ(1-42)-induced model of Alzheimer’s disease. Aging Cell. 2020;19:e13046.
62. Kreth S, Ledderose C, Schütz S, et al. MicroRNA-150 inhibits expression of adiponectin receptor 2 and is a potential therapeutic target in patients with chronic heart failure. J Heart Lung Transplant. 2014;33:252-60.
63. Choi HM, Doss HM, Kim KS. Multifaceted physiological roles of adiponectin in inflammation and diseases. Int J Mol Sci. 2020;21:1219.
64. Signoriello E, Mallardo M, Nigro E, et al. Adiponectin in cerebrospinal fluid from patients affected by multiple sclerosis is correlated with the progression and severity of disease. Mol Neurobiol. 2021;58:2663-70.
65. Signoriello E, Lus G, Polito R, et al. Adiponectin profile at baseline is correlated to progression and severity of multiple sclerosis. Eur J Neurol. 2019;26:348-55.
66. Liu N, Jiang N, Guo R, et al. MiR-451 inhibits cell growth and invasion by targeting MIF and is associated with survival in nasopharyngeal carcinoma. Mol Cancer. 2013;12:123.
67. Bai S, Zhang G, Chen S, et al. MicroRNA-451 regulates angiogenesis in intracerebral hemorrhage by targeting macrophage migration inhibitory factor. Mol Neurobiol. 2024;61:10481-99.
68. Mamoori A, Gopalan V, Lu CT, et al. Expression pattern of miR-451 and its target MIF (macrophage migration inhibitory factor) in colorectal cancer. J Clin Pathol. 2017;70:308-12.
69. Wang Y, An R, Umanah GK, et al. A nuclease that mediates cell death induced by DNA damage and poly(ADP-ribose) polymerase-1. Science. 2016;354:aad6872.
70. Cox GM, Kithcart AP, Pitt D, et al. Macrophage migration inhibitory factor potentiates autoimmune-mediated neuroinflammation. J Immunol. 2013;191:1043-54.
71. Fagone P, Mazzon E, Cavalli E, et al. Contribution of the macrophage migration inhibitory factor superfamily of cytokines in the pathogenesis of preclinical and human multiple sclerosis: in silico and in vivo evidences. J Neuroimmunol. 2018;322:46-56.
72. Schneider A, Long SA, Cerosaletti K, et al. In active relapsing-remitting multiple sclerosis, effector T cell resistance to adaptive Tregs involves IL-6-mediated signaling. Sci Transl Med. 2013;5:170ra15.
73. Sun Y, Peng YB, Ye LL, Ma LX, Zou MY, Cheng ZG. Propofol inhibits proliferation and cisplatin resistance in ovarian cancer cells through regulating the microRNA-374a/forkhead box O1 signaling axis. Mol Med Rep. 2020;21:1471-80.
74. Wang C, Su K, Lin H, Cen B, Zheng S, Xu X. Identification and verification of a novel MAGI2-AS3/miRNA-374-5p/FOXO1 network associated with HBV-related HCC. Cells. 2022;11:3466.
75. Kraus EE, Kakuk-Atkins L, Farinas MF, Jeffers M, Lovett-Racke AE, Yang Y. Regulation of autoreactive CD4 T cells by FoxO1 signaling in CNS autoimmunity. J Neuroimmunol. 2021;359:577675.
76. Luo CT, Liao W, Dadi S, Toure A, Li MO. Graded Foxo1 activity in Treg cells differentiates tumour immunity from spontaneous autoimmunity. Nature. 2016;529:532-6.
77. Tsai WC, Hsu PW, Lai TC, et al. MicroRNA-122, a tumor suppressor microRNA that regulates intrahepatic metastasis of hepatocellular carcinoma. Hepatology. 2009;49:1571-82.
78. Kieseier BC, Pischel H, Neuen-Jacob E, Tourtellotte WW, Hartung HP. ADAM-10 and ADAM-17 in the inflamed human CNS. Glia. 2003;42:398-405.
79. Plumb J, McQuaid S, Cross AK, et al. Upregulation of ADAM-17 expression in active lesions in multiple sclerosis. Mult Scler. 2006;12:375-85.
80. Iliopoulos D, Hirsch HA, Struhl K. An epigenetic switch involving NF-kappaB, Lin28, Let-7 microRNA, and IL6 links inflammation to cell transformation. Cell. 2009;139:693-706.
82. Petković F, Castellano B. The role of interleukin-6 in central nervous system demyelination. Neural Regen Res. 2016;11:1922-3.
83. Schönrock LM, Gawlowski G, Brück W. Interleukin-6 expression in human multiple sclerosis lesions. Neurosci Lett. 2000;294:45-8.
84. Tanaka T, Narazaki M, Kishimoto T. IL-6 in inflammation, immunity, and disease. Cold Spring Harb Perspect Biol. 2014;6:a016295.
85. Xu B, Zhang Y, Du XF, et al. Neurons secrete miR-132-containing exosomes to regulate brain vascular integrity. Cell Res. 2017;27:882-97.
86. Qu R, Peng Y, Zhou M, et al. MiR-223-3p attenuates M1 macrophage polarization via suppressing the Notch signaling pathway and NLRP3-mediated pyroptosis in experimental autoimmune uveitis. Eur J Pharmacol. 2023;960:176139.
87. Henkel T, Ling PD, Hayward SD, Peterson MG. Mediation of Epstein-Barr virus EBNA2 transactivation by recombination signal-binding protein J kappa. Science. 1994;265:92-5.
88. Ricigliano VA, Handel AE, Sandve GK, et al. EBNA2 binds to genomic intervals associated with multiple sclerosis and overlaps with vitamin D receptor occupancy. PLoS One. 2015;10:e0119605.
89. Ramakrishnaiah V, Thumann C, Fofana I, et al. Exosome-mediated transmission of hepatitis C virus between human hepatoma Huh7.5 cells. Proc Natl Acad Sci U S A. 2013;110:13109-13.
91. 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.
92. Chen Y, Fachko DN, Ivanov NS, Skalsky RL. B cell receptor-responsive miR-141 enhances Epstein-Barr virus lytic cycle via FOXO3 inhibition. mSphere. 2021;6:e00093-21.
93. Almulla AF, Vojdani A, Zhang Y, Vojdani E, Maes M. HHV-6 and EBV reactivation in relapsing remitting multiple sclerosis: disability, progression, and inflammation links. iScience. 2025;28:113048.
94. Skalsky RL, Kang D, Linnstaedt SD, Cullen BR. Evolutionary conservation of primate lymphocryptovirus microRNA targets. J Virol. 2014;88:1617-35.
95. Rahimi SB, Hashemi SMA, Poursadeghfard M, et al. EBV viral load and miR-BART10-3p expression in multiple sclerosis: a case-control study. Mult Scler Relat Disord. 2026;110:107108.
96. Park J, Kim H, Roh YH, Ko J. Advances in single extracellular vesicle characterization and multiplexed profiling. TrAC Trends in Analytical Chemistry. 2026;195:118588.
98. Cohen A, Zinger A, Tiberti N, Grau GER, Combes V. Differential plasma microvesicle and brain profiles of microRNA in experimental cerebral malaria. Malar J. 2018;17:192.
99. El-Assaad F, Hempel C, Combes V, et al. Differential microRNA expression in experimental cerebral and noncerebral malaria. Infect Immun. 2011;79:2379-84.
