REFERENCES
1. Chopp M, Li Y. Treatment of neural injury with marrow stromal cells. Lancet Neurol. 2002;1:92-100.
2. Haynesworth SE, Baber MA, Caplan AI. Cytokine expression by human marrow-derived mesenchymal progenitor cells in vitro: effects of dexamethasone and IL-1α. J Cell Physiol. 1996;166:585-92.
3. Caplan AI, Dennis JE. Mesenchymal stem cells as trophic mediators. J Cell Biochem. 2006;98:1076-84.
4. Timmers L, Lim SK, Arslan F, et al. Reduction of myocardial infarct size by human mesenchymal stem cell conditioned medium. Stem Cell Res. 2007;1:129-37.
5. Bruno S, Grange C, Deregibus MC, et al. Mesenchymal stem cell-derived microvesicles protect against acute tubular injury. J Am Soc Nephrol. 2009;20:1053-67.
6. Lai RC, Arslan F, Lee MM, et al. Exosome secreted by MSC reduces myocardial ischemia/reperfusion injury. Stem Cell Res. 2010;4:214-22.
7. Bruno S, Tapparo M, Collino F, et al. Renal regenerative potential of different extracellular vesicle populations derived from bone marrow mesenchymal stromal cells. Tissue Eng Part A. 2017;23:1262-73.
8. Witwer KW, Van Balkom BWM, Bruno S, et al. Defining mesenchymal stromal cell (MSC)-derived small extracellular vesicles for therapeutic applications. J Extracell Vesicles. 2019;8:1609206.
9. Doeppner TR, Herz J, Görgens A, et al. Extracellular vesicles improve post-stroke neuroregeneration and prevent postischemic immunosuppression. Stem Cells Transl Med. 2015;4:1131-43.
10. Nawaz M, Fatima F, Vallabhaneni KC, et al. Extracellular vesicles: evolving factors in stem cell biology. Stem Cells Int. 2016;2016:1073140.
11. Baek G, Choi H, Kim Y, Lee HC, Choi C. Mesenchymal stem cell-derived extracellular vesicles as therapeutics and as a drug delivery platform. Stem Cells Transl Med. 2019;8:880-6.
12. Tertel T, Dittrich R, Arsène P, Jensen A, Giebel B. EV products obtained from iPSC-derived MSCs show batch-to-batch variations in their ability to modulate allogeneic immune responses in vitro. Front Cell Dev Biol. 2023;11:1282860.
13. Hollmann J, Brecht J, Goetzke R, et al. Genetic barcoding reveals clonal dominance in iPSC-derived mesenchymal stromal cells. Stem Cell Res Ther. 2020;11:105.
14. Friedenstein AJ, Chailakhjan RK, Lalykina KS. The development of fibroblast colonies in monolayer cultures of guinea-pig bone marrow and spleen cells. Cell Tissue Kinet. 1970;3:393-403.
15. Romanov YA, Darevskaya AN, Merzlikina NV, Buravkova LB. Mesenchymal stem cells from human bone marrow and adipose tissue: isolation, characterization, and differentiation potentialities. Bull Exp Biol Med. 2005;140:138-43.
16. In ’t Anker PS, Scherjon SA, Kleijburg-van der Keur C, et al. Isolation of mesenchymal stem cells of fetal or maternal origin from human placenta. Stem Cells. 2004;22:1338-45.
17. Kestendjieva S, Kyurkchiev D, Tsvetkova G, et al. Characterization of mesenchymal stem cells isolated from the human umbilical cord. Cell Biol Int. 2008;32:724-32.
18. Lian Q, Lye E, Suan Yeo K, et al. Derivation of clinically compliant MSCs from CD105+, CD24- differentiated human ESCs. Stem Cells. 2007;25:425-36.
19. Hynes K, Menicanin D, Mrozik K, Gronthos S, Bartold PM. Generation of functional mesenchymal stem cells from different induced pluripotent stem cell lines. Stem Cells Dev. 2014;23:1084-96.
20. Allan D, Tieu A, Lalu M, Burger D. Mesenchymal stromal cell-derived extracellular vesicles for regenerative therapy and immune modulation: progress and challenges toward clinical application. Stem Cells Transl Med. 2020;9:39-46.
21. Madel RJ, Börger V, Dittrich R, et al. Independent human mesenchymal stromal cell-derived extracellular vesicle preparations differentially attenuate symptoms in an advanced murine graft-versus-host disease model. Cytotherapy. 2023;25:821-36.
22. Selich A, Daudert J, Hass R, et al. Massive clonal selection and transiently contributing clones during expansion of mesenchymal stem cell cultures revealed by lentiviral RGB-barcode technology. Stem Cells Transl Med. 2016;5:591-601.
23. McLeod CM, Mauck RL. On the origin and impact of mesenchymal stem cell heterogeneity: new insights and emerging tools for single cell analysis. Eur Cell Mater. 2017;34:217-31.
24. Chen TS, Arslan F, Yin Y, et al. Enabling a robust scalable manufacturing process for therapeutic exosomes through oncogenic immortalization of human ESC-derived MSCs. J Transl Med. 2011;9:47.
25. Kraskiewicz H, Paprocka M, Bielawska-Pohl A, et al. Can supernatant from immortalized adipose tissue MSC replace cell therapy? Stem Cell Res Ther. 2020;11:29.
26. Labusek N, Mouloud Y, Köster C, et al. Extracellular vesicles from immortalized mesenchymal stromal cells protect against neonatal hypoxic-ischemic brain injury. Inflamm Regen. 2023;43:24.
27. Shekari F, Alibhai FJ, Baharvand H, et al. Cell culture-derived extracellular vesicles: considerations for reporting cell culturing parameters. J Extracell Biol. 2023;2:e115.
28. Patel DB, Gray KM, Santharam Y, Lamichhane TN, Stroka KM, Jay SM. Impact of cell culture parameters on production and vascularization bioactivity of mesenchymal stem cell-derived extracellular vesicles. Bioeng Transl Med. 2017;2:170-9.
29. Almeria C, Kreß S, Weber V, Egger D, Kasper C. Heterogeneity of mesenchymal stem cell-derived extracellular vesicles is highly impacted by the tissue/cell source and culture conditions. Cell Biosci. 2022;12:51.
30. Gimona M, Brizzi MF, Choo ABH, et al. Critical considerations for the development of potency tests for therapeutic applications of mesenchymal stromal cell-derived small extracellular vesicles. Cytotherapy. 2021;23:373-80.
31. Bonsergent E, Grisard E, Buchrieser J, Schwartz O, Théry C, Lavieu G. Quantitative characterization of extracellular vesicle uptake and content delivery within mammalian cells. Nat Commun. 2021;12:1864.
32. O'Brien K, Ughetto S, Mahjoum S, Nair AV, Breakefield XO. Uptake, functionality, and re-release of extracellular vesicle-encapsulated cargo. Cell Rep. 2022;39:110651.
33. Zickler AM, Liang X, Gupta D, et al. Novel endogenous engineering platform for robust loading and delivery of functional mRNA by extracellular vesicles. Adv Sci. 2024;11:e2407619.
34. Liang X, Gupta D, Xie J, et al. Multimodal engineering of extracellular vesicles for efficient intracellular protein delivery. bioRxiv. 2023; bioRxiv:2023.04.30.535834.
35. Jahnke K, Staufer O. Membranes on the move: the functional role of the extracellular vesicle membrane for contact-dependent cellular signalling. J Extracell Vesicles. 2024;13:e12436.
36. Tan TT, Lai RC, Sim WK, Zhang B, Lim SK. Enhancing EV-cell communication through “External Modulation of Cell by EV” (EMCEV). Cytotherapy. 2025;27:1-6.
37. Tan TT, Lim SK. Relevance of RNA to the therapeutic efficacy of mesenchymal stromal/stem cells extracellular vesicles. RNA Biol. 2025;22:1-7.
38. Baglio SR, Rooijers K, Koppers-Lalic D, et al. Human bone marrow- and adipose-mesenchymal stem cells secrete exosomes enriched in distinctive miRNA and tRNA species. Stem Cell Res Ther. 2015;6:127.
39. De Luca L, Trino S, Laurenzana I, et al. MiRNAs and piRNAs from bone marrow mesenchymal stem cell extracellular vesicles induce cell survival and inhibit cell differentiation of cord blood hematopoietic stem cells: a new insight in transplantation. Oncotarget. 2016;7:6676-92.
40. Lai RC, Tan SS, Yeo RW, et al. MSC secretes at least 3 EV types each with a unique permutation of membrane lipid, protein and RNA. J Extracell Vesicles. 2016;5:29828.
41. Kodali M, Madhu LN, Reger RL, et al. Intranasally administered human MSC-derived extracellular vesicles inhibit NLRP3-p38/MAPK signaling after TBI and prevent chronic brain dysfunction. Brain Behav Immun. 2023;108:118-34.
42. Kuchen S, Resch W, Yamane A, et al. Regulation of microRNA expression and abundance during lymphopoiesis. Immunity. 2010;32:828-39.
43. Hanson B, Vorobieva I, Zheng W, et al. EV-mediated promotion of myogenic differentiation is dependent on dose, collection medium, and isolation method. Mol Ther Nucleic Acids. 2023;33:511-28.
44. 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.
45. 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.
46. Bissels U, Wild S, Tomiuk S, et al. Absolute quantification of microRNAs by using a universal reference. RNA. 2009;15:2375-84.
47. Chang J, Nicolas E, Marks D, et al. miR-122, a mammalian liver-specific microRNA, is processed from hcr mRNA and may downregulate the high affinity cationic amino acid transporter CAT-1. RNA Biol. 2004;1:106-13.
48. Denzler R, Agarwal V, Stefano J, Bartel DP, Stoffel M. Assessing the ceRNA hypothesis with quantitative measurements of miRNA and target abundance. Mol Cell. 2014;54:766-76.
49. Ho V, Baker JR, Willison KR, Barnes PJ, Donnelly LE, Klug DR. Single cell quantification of microRNA from small numbers of non-invasively sampled primary human cells. Commun Biol. 2023;6:458.
50. Liang Y, Ridzon D, Wong L, Chen C. Characterization of microRNA expression profiles in normal human tissues. BMC Genomics. 2007;8:166.
51. Valdmanis PN, Kim HK, Chu K, et al. miR-122 removal in the liver activates imprinted microRNAs and enables more effective microRNA-mediated gene repression. Nat Commun. 2018;9:5321.
52. van Balkom BWM, Gremmels H, Giebel B, Lim SK. Proteomic signature of mesenchymal stromal cell-derived small extracellular vesicles. Proteomics. 2019;19:e1800163.







