REFERENCES
1. Nigri J, Leca J, Tubiana SS, et al. CD9 mediates the uptake of extracellular vesicles from cancer-associated fibroblasts that promote pancreatic cancer cell aggressiveness. Sci Signal 2022;15:eabg8191.
2. Berumen Sánchez G, Bunn KE, Pua HH, Rafat M. Extracellular vesicles: mediators of intercellular communication in tissue injury and disease. Cell Commun Signal 2021;19:104.
3. Park JE, Dutta B, Tse SW, et al. Hypoxia-induced tumor exosomes promote M2-like macrophage polarization of infiltrating myeloid cells and microRNA-mediated metabolic shift. Oncogene 2019;38:5158-73.
4. Théry C, Witwer KW, Aikawa E, et al. Minimal information for studies of extracellular vesicles 2018 (MISEV2018): a position statement of the international society for extracellular vesicles and update of the MISEV2014 guidelines. J Extracell Vesicles 2018;7:1535750.
5. Teng F, Fussenegger M. Shedding light on extracellular vesicle biogenesis and bioengineering. Adv Sci 2020;8:2003505.
6. van Niel G, D'Angelo G, Raposo G. Shedding light on the cell biology of extracellular vesicles. Nat Rev Mol Cell Biol 2018;19:213-28.
7. Korbei B. Ubiquitination of the ubiquitin-binding machinery: how early ESCRT components are controlled. Essays Biochem 2022;66:169-77.
9. Meister M, Bänfer S, Gärtner U, et al. Regulation of cargo transfer between ESCRT-0 and ESCRT-I complexes by flotillin-1 during endosomal sorting of ubiquitinated cargo. Oncogenesis 2017;6:e344.
10. Wang J, Zhuang X, Greene KS, et al. Cdc42 functions as a regulatory node for tumour-derived microvesicle biogenesis. J Extracell Vesicles 2021;10:e12051.
11. Li B, Antonyak MA, Zhang J, Cerione RA. RhoA triggers a specific signaling pathway that generates transforming microvesicles in cancer cells. Oncogene 2012;31:4740-9.
12. Li M, Liao L, Tian W. Extracellular vesicles derived from apoptotic cells: an essential link between death and regeneration. Front Cell Dev Biol 2020;8:573511.
13. Willms E, Johansson HJ, Mäger I, et al. Cells release subpopulations of exosomes with distinct molecular and biological properties. Sci Rep 2016;6:22519.
14. Colombo M, Moita C, van Niel G, et al. Analysis of ESCRT functions in exosome biogenesis, composition and secretion highlights the heterogeneity of extracellular vesicles. J Cell Sci 2013;126:5553-65.
15. Joshi BS, Ortiz D, Zuhorn IS. Converting extracellular vesicles into nanomedicine: loading and unloading of cargo. Materials Today Nano 2021;16:100148.
16. Kamei N, Nishimura H, Matsumoto A, et al. Comparative study of commercial protocols for high recovery of high-purity mesenchymal stem cell-derived extracellular vesicle isolation and their efficient labeling with fluorescent dyes. Nanomedicine 2021;35:102396.
17. García-Romero N, Madurga R, Rackov G, et al. Polyethylene glycol improves current methods for circulating extracellular vesicle-derived DNA isolation. J Transl Med 2019;17:75.
18. Zhang Q, Jeppesen DK, Higginbotham JN, Franklin JL, Coffey RJ. Comprehensive isolation of extracellular vesicles and nanoparticles. Nat Protoc 2023;18:1462-87.
19. Zhang X, Borg EGF, Liaci AM, Vos HR, Stoorvogel W. A novel three step protocol to isolate extracellular vesicles from plasma or cell culture medium with both high yield and purity. J Extracell Vesicles 2020;9:1791450.
20. Zhang H, Freitas D, Kim HS, et al. Identification of distinct nanoparticles and subsets of extracellular vesicles by asymmetric flow field-flow fractionation. Nat Cell Biol 2018;20:332-43.
21. Gomes PA, Bodo C, Nogueras-Ortiz C, et al. A novel isolation method for spontaneously released extracellular vesicles from brain tissue and its implications for stress-driven brain pathology. Cell Commun Signal 2023;21:35.
22. 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.
23. Ghoshal B, Bertrand E, Bhattacharyya SN. Non-canonical argonaute loading of extracellular vesicle-derived exogenous single-stranded miRNA in recipient cells. J Cell Sci 2021;134:jcs253914.
24. Seras-Franzoso J, Díaz-Riascos ZV, Corchero JL, et al. Extracellular vesicles from recombinant cell factories improve the activity and efficacy of enzymes defective in lysosomal storage disorders. J Extracell Vesicles 2021;10:e12058.
25. Li H, Pinilla-Macua I, Ouyang Y, et al. Internalization of trophoblastic small extracellular vesicles and detection of their miRNA cargo in P-bodies. J Extracell Vesicles 2020;9:1812261.
26. Heusermann W, Hean J, Trojer D, et al. Exosomes surf on filopodia to enter cells at endocytic hot spots, traffic within endosomes, and are targeted to the ER. J Cell Biol 2016;213:173-84.
27. Feng D, Zhao WL, Ye YY, et al. Cellular internalization of exosomes occurs through phagocytosis. Traffic 2010;11:675-87.
28. Nakase I, Kobayashi NB, Takatani-Nakase T, Yoshida T. Active macropinocytosis induction by stimulation of epidermal growth factor receptor and oncogenic Ras expression potentiates cellular uptake efficacy of exosomes. Sci Rep 2015;5:10300.
29. Tian T, Zhu YL, Zhou YY, et al. Exosome uptake through clathrin-mediated endocytosis and macropinocytosis and mediating miR-21 delivery. J Biol Chem 2014;289:22258-67.
30. Escrevente C, Keller S, Altevogt P, Costa J. Interaction and uptake of exosomes by ovarian cancer cells. BMC Cancer 2011;11:108.
31. Svensson KJ, Christianson HC, Wittrup A, et al. Exosome uptake depends on ERK1/2-heat shock protein 27 signaling and lipid Raft-mediated endocytosis negatively regulated by caveolin-1. J Biol Chem 2013;288:17713-24.
32. Ginini L, Billan S, Fridman E, Gil Z. Insight into extracellular vesicle-cell communication: from cell recognition to intracellular fate. Cells 2022;11:1375.
33. Mulcahy LA, Pink RC, Carter DRF. Routes and mechanisms of extracellular vesicle uptake. J Extracell Vesicles 2014;3:24641.
34. Phuyal S, Hessvik NP, Skotland T, Sandvig K, Llorente A. Regulation of exosome release by glycosphingolipids and flotillins. FEBS J 2014;281:2214-27.
35. Muralidharan-Chari V, Clancy J, Plou C, et al. ARF6-regulated shedding of tumor cell-derived plasma membrane microvesicles. Curr Biol 2009;19:1875-85.
36. Schelhaas M, Ewers H, Rajamäki ML, Day PM, Schiller JT, Helenius A. Human papillomavirus type 16 entry: retrograde cell surface transport along actin-rich protrusions. PLoS Pathog 2008;4:e1000148.
37. Hunziker A, Glas I, Pohl MO, Stertz S. Phosphoproteomic profiling of influenza virus entry reveals infection-triggered filopodia induction counteracted by dynamic cortactin phosphorylation. Cell Rep 2022;38:110306.
38. ur Rehman Z, Sjollema KA, Kuipers J, Hoekstra D, Zuhorn IS. Nonviral gene delivery vectors use syndecan-dependent transport mechanisms in filopodia to reach the cell surface. ACS Nano 2012;6:7521-32.
39. Zuhorn IS, Kalicharan R, Hoekstra D. Lipoplex-mediated transfection of mammalian cells occurs through the cholesterol-dependent clathrin-mediated pathway of endocytosis. J Biol Chem 2002;277:18021-8.
40. Nakase I, Noguchi K, Aoki A, Takatani-Nakase T, Fujii I, Futaki S. Arginine-rich cell-penetrating peptide-modified extracellular vesicles for active macropinocytosis induction and efficient intracellular delivery. Sci Rep 2017;7:1991.
41. Nakase I, Ueno N, Matsuzawa M, et al. Environmental pH stress influences cellular secretion and uptake of extracellular vesicles. FEBS Open Bio 2021;11:753-67.
42. Shimoda A, Miura R, Tateno H, et al. Assessment of surface glycan diversity on extracellular vesicles by lectin microarray and glycoengineering strategies for drug delivery applications. Small Methods 2022;6:e2100785.
43. Rejman J, Oberle V, Zuhorn IS, Hoekstra D. Size-dependent internalization of particles via the pathways of clathrin- and caveolae-mediated endocytosis. Biochem J 2004;377:159-69.
44. Peretz V, Motiei M, Sukenik CN, Popovtzer R. The effect of nanoparticle size on cellular binding probability. J AT Mol Opt 2012;2012:1-7.
45. Kowal J, Arras G, Colombo M, et al. Proteomic comparison defines novel markers to characterize heterogeneous populations of extracellular vesicle subtypes. Proc Natl Acad Sci USA 2016;113:E968-77.
46. Mathieu M, Névo N, Jouve M, et al. Specificities of exosome versus small ectosome secretion revealed by live intracellular tracking of CD63 and CD9. Nat Commun 2021;12:4389.
47. Cardeñes B, Clares I, Toribio V, et al. Cellular integrin α5β1 and exosomal ADAM17 mediate the binding and uptake of exosomes produced by colorectal carcinoma cells. Int J Mol Sci 2021;22:9938.
48. Yáñez-Mó M, Barreiro O, Gordon-Alonso M, Sala-Valdés M, Sánchez-Madrid F. Tetraspanin-enriched microdomains: a functional unit in cell plasma membranes. Trends Cell Biol 2009;19:434-46.
49. Hantak MP, Qing E, Earnest JT, Gallagher T. Tetraspanins: architects of viral entry and exit platforms. J Virol 2019;93:e01429-17.
50. Tognoli ML, Dancourt J, Bonsergent E, et al. Lack of involvement of CD63 and CD9 tetraspanins in the extracellular vesicle content delivery process. Commun Biol 2023;6:532.
51. Pang X, He X, Qiu Z, et al. Targeting integrin pathways: mechanisms and advances in therapy. Signal Transduct Target Ther 2023;8:1.
52. Hoshino A, Costa-Silva B, Shen TL, et al. Tumour exosome integrins determine organotropic metastasis. Nature 2015;527:329-35.
53. Altei WF, Pachane BC, Dos Santos PK, et al. Inhibition of αvβ3 integrin impairs adhesion and uptake of tumor-derived small extracellular vesicles. Cell Commun Signal 2020;18:158.
54. Feng Y, Chen Q, Lau SY, et al. The blocking of integrin-mediated interactions with maternal endothelial cells reversed the endothelial cell dysfunction induced by evs, derived from preeclamptic placentae. Int J Mol Sci 2022;23:13115.
55. Ludwig BS, Kessler H, Kossatz S, Reuning U. RGD-binding integrins revisited: how recently discovered functions and novel synthetic ligands (Re-)shape an ever-evolving field. Cancers 2021;13:1711.
56. You Y, Borgmann K, Edara VV, Stacy S, Ghorpade A, Ikezu T. Activated human astrocyte-derived extracellular vesicles modulate neuronal uptake, differentiation and firing. J Extracell Vesicles 2020;9:1706801.
57. Zheng W, He R, Liang X, et al. Cell-specific targeting of extracellular vesicles though engineering the glycocalyx. J Extracell Vesicles 2022;11:e12290.
58. Parolini I, Federici C, Raggi C, et al. Microenvironmental pH is a key factor for exosome traffic in tumor cells. J Biol Chem 2009;284:34211-22.
59. 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.
60. Cui GH, Guo HD, Li H, et al. RVG-modified exosomes derived from mesenchymal stem cells rescue memory deficits by regulating inflammatory responses in a mouse model of Alzheimer's disease. Immun Ageing 2019;16:10.
61. Xue C, Shen Y, Li X, et al. Exosomes derived from hypoxia-treated human adipose mesenchymal stem cells enhance angiogenesis through the PKA signaling pathway. Stem Cells Dev 2018;27:456-65.
62. Pachane BC, Nunes ACC, Cataldi TR, et al. Small extracellular vesicles from hypoxic triple-negative breast cancer cells induce oxygen-dependent cell invasion. Int J Mol Sci 2022;23:12646.
63. Gong C, Zhang X, Shi M, et al. Tumor exosomes reprogrammed by low pH are efficient targeting vehicles for smart drug delivery and personalized therapy against their homologous tumor. Adv Sci 2021;8:2002787.
64. Cerezo-Magaña M, Christianson HC, van Kuppevelt TH, Forsberg-Nilsson K, Belting M. Hypoxic induction of exosome uptake through proteoglycan-dependent endocytosis fuels the lipid droplet phenotype in glioma. Mol Cancer Res 2021;19:528-40.
65. Fukuta T, Nishikawa A, Kogure K. Low level electricity increases the secretion of extracellular vesicles from cultured cells. Biochem Biophys Rep 2020;21:100713.
66. Hisey CL, Artuyants A, Guo G, et al. Investigating the consistency of extracellular vesicle production from breast cancer subtypes using CELLine adherent bioreactors. J of Extracellular Bio 2022;1:e60.
67. Sajidah ES, Lim K, Yamano T, et al. Spatiotemporal tracking of small extracellular vesicle nanotopology in response to physicochemical stresses revealed by HS-AFM. J Extracell Vesicles 2022;11:e12275.
68. Morandi MI, Busko P, Ozer-Partuk E, et al. Extracellular vesicle fusion visualized by cryo-EM. PNAS Nexus 2022;1:pgac156.
69. Yao Z, Qiao Y, Li X, et al. Exosomes exploit the virus entry machinery and pathway to transmit alpha interferon-induced antiviral activity. J Virol 2018;92:e01578-18.
70. Toribio V, Morales S, López-Martín S, Cardeñes B, Cabañas C, Yáñez-Mó M. Development of a quantitative method to measure EV uptake. Sci Rep 2019;9:10522.
71. Ilahibaks NF, Ardisasmita AI, Xie S, et al. TOP-EVs: technology of protein delivery through extracellular vesicles is a versatile platform for intracellular protein delivery. J Control Release 2023;355:579-92.
72. Bui S, Dancourt J, Lavieu G. Virus-free method to control and enhance extracellular vesicle cargo loading and delivery. ACS Appl Bio Mater 2023;6:1081-91.
73. Zhang C, Schekman R. Syncytin-mediated open-ended membrane tubular connections facilitate the intercellular transfer of cargos including Cas9 protein. Elife 2023;12:e84391.
74. Uygur B, Melikov K, Arakelyan A, Margolis LB, Chernomordik LV. Syncytin 1 dependent horizontal transfer of marker genes from retrovirally transduced cells. Sci Rep 2019;9:17637.
75. Somiya M, Kuroda S. Reporter gene assay for membrane fusion of extracellular vesicles. J Extracell Vesicles 2021;10:e12171.
76. Somiya M, Kuroda S. Real-time luminescence assay for cytoplasmic cargo delivery of extracellular vesicles. Anal Chem 2021;93:5612-20.
77. Dennison SM, Greenfield N, Lenard J, Lentz BR. VSV transmembrane domain (TMD) peptide promotes PEG-mediated fusion of liposomes in a conformationally sensitive fashion. Biochemistry 2002;41:14925-34.
78. Schnell U, Kuipers J, Giepmans BN. EpCAM proteolysis: new fragments with distinct functions? Biosci Rep 2013;33:e00030.
79. Kooijmans SA, Vader P, van Dommelen SM, van Solinge WW, Schiffelers RM. Exosome mimetics: a novel class of drug delivery systems. Int J Nanomedicine 2012;7:1525-41.
80. Stratton BS, Warner JM, Wu Z, et al. Cholesterol increases the openness of SNARE-mediated flickering fusion pores. Biophys J 2016;110:1538-50.
81. Kreutzberger AJ, Kiessling V, Tamm LK. High cholesterol obviates a prolonged hemifusion intermediate in fast SNARE-mediated membrane fusion. Biophys J 2015;109:319-29.
82. Bonsergent E, Lavieu G. Content release of extracellular vesicles in a cell-free extract. FEBS Lett 2019;593:1983-92.
83. Lu J, Pan Q, Rong L, He W, Liu SL, Liang C. The IFITM proteins inhibit HIV-1 infection. J Virol 2011;85:2126-37.
84. Tartour K, Appourchaux R, Gaillard J, et al. IFITM proteins are incorporated onto HIV-1 virion particles and negatively imprint their infectivity. Retrovirology 2014;11:103.
85. Weidner JM, Jiang D, Pan XB, Chang J, Block TM, Guo JT. Interferon-induced cell membrane proteins, IFITM3 and tetherin, inhibit vesicular stomatitis virus infection via distinct mechanisms. J Virol 2010;84:12646-57.
86. Buchrieser J, Degrelle SA, Couderc T, et al. IFITM proteins inhibit placental syncytiotrophoblast formation and promote fetal demise. Science 2019;365:176-80.
87. Perrin P, Janssen L, Janssen H, et al. Retrofusion of intralumenal MVB membranes parallels viral infection and coexists with exosome release. Curr Biol 2021;31:3884-3893.e4.
88. Somiya M. Where does the cargo go?: solutions to provide experimental support for the "extracellular vesicle cargo transfer hypothesis". J Cell Commun Signal 2020;14:135-46.
89. Gaudin Y. Reversibility in fusion protein conformational changes the intriguing case of rhabdovirus-induced membrane fusion. Subcell Biochem ;34:379-408.
91. Fontana J, Steven AC. Influenza virus-mediated membrane fusion: structural insights from electron microscopy. Arch Biochem Biophys 2015;581:86-97.
92. Chlanda P, Mekhedov E, Waters H, et al. The hemifusion structure induced by influenza virus haemagglutinin is determined by physical properties of the target membranes. Nat Microbiol 2016;1:16050.
93. Mitsui K, Koshimura Y, Yoshikawa Y, Matsushita M, Kanazawa H. The endosomal Na(+)/H(+) exchanger contributes to multivesicular body formation by regulating the recruitment of ESCRT-0 Vps27p to the endosomal membrane. J Biol Chem 2011;286:37625-38.
94. Boersma AJ, Zuhorn IS, Poolman B. A sensor for quantification of macromolecular crowding in living cells. Nat Methods 2015;12:227-9, 1 p following 229.
95. Wang Z, Chen D, Guan D, et al. Material properties of phase-separated TFEB condensates regulate the autophagy-lysosome pathway. J Cell Biol 2022;221:e202112024.
96. Liu XM, Ma L, Schekman R. Selective sorting of microRNAs into exosomes by phase-separated YBX1 condensates. Elife 2021;10:e71982.
97. de Gassart A, Geminard C, Hoekstra D, Vidal M. Exosome secretion: the art of reutilizing nonrecycled proteins? Traffic 2004;5:896-903.
99. Wang S, Sun H, Tanowitz M, Liang XH, Crooke ST. Intra-endosomal trafficking mediated by lysobisphosphatidic acid contributes to intracellular release of phosphorothioate-modified antisense oligonucleotides. Nucleic Acids Res 2017;45:5309-22.
100. ur Rehman Z, Hoekstra D, Zuhorn IS. Protein kinase A inhibition modulates the intracellular routing of gene delivery vehicles in HeLa cells, leading to productive transfection. J Control Release 2011;156:76-84.
101. Savina A, Fader CM, Damiani MT, Colombo MI. Rab11 promotes docking and fusion of multivesicular bodies in a calcium-dependent manner. Traffic 2005;6:131-43.
102. Schafer JC, Baetz NW, Lapierre LA, McRae RE, Roland JT, Goldenring JR. Rab11-FIP2 interaction with MYO5B regulates movement of Rab11a-containing recycling vesicles. Traffic 2014;15:292-308.
103. Ostrowski M, Carmo NB, Krumeich S, et al. Rab27a and Rab27b control different steps of the exosome secretion pathway. Nat Cell Biol 2010;12:19-30.
104. Hsu C, Morohashi Y, Yoshimura S, et al. Regulation of exosome secretion by Rab35 and its GTPase-activating proteins TBC1D10A-C. J Cell Biol 2010;189:223-32.
105. Joshi BS, Zuhorn IS. Heparan sulfate proteoglycan-mediated dynamin-dependent transport of neural stem cell exosomes in an in vitro blood-brain barrier model. Eur J Neurosci 2021;53:706-19.
106. Singh A, Fedele C, Lu H, Nevalainen MT, Keen JH, Languino LR. Exosome-mediated Transfer of αvβ3 integrin from tumorigenic to nontumorigenic cells promotes a migratory phenotype. Mol Cancer Res 2016;14:1136-46.
107. Reclusa P, Verstraelen P, Taverna S, et al. Improving extracellular vesicles visualization: from static to motion. Sci Rep 2020;10:6494.
108. Simonsen JB. Pitfalls associated with lipophilic fluorophore staining of extracellular vesicles for uptake studies. J Extracell Vesicles 2019;8:1582237.
109. Degors IMS, Wang C, Rehman ZU, Zuhorn IS. Carriers break barriers in drug delivery: endocytosis and endosomal escape of gene delivery vectors. Acc Chem Res 2019;52:1750-60.
110. Rennick JJ, Johnston APR, Parton RG. Key principles and methods for studying the endocytosis of biological and nanoparticle therapeutics. Nat Nanotechnol 2021;16:266-76.
111. Georgieva JV, Kalicharan D, Couraud PO, et al. Surface characteristics of nanoparticles determine their intracellular fate in and processing by human blood-brain barrier endothelial cells in vitro. Mol Ther 2011;19:318-25.
112. Vercauteren D, Vandenbroucke RE, Jones AT, et al. The use of inhibitors to study endocytic pathways of gene carriers: optimization and pitfalls. Mol Ther 2010;18:561-9.
113. Itakura S, Shohji A, Amagai S, et al. Gene knockdown in HaCaT cells by small interfering RNAs entrapped in grapefruit-derived extracellular vesicles using a microfluidic device. Sci Rep 2023;13:3102.
114. 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.
115. Rappa G, Santos MF, Green TM, et al. Nuclear transport of cancer extracellular vesicle-derived biomaterials through nuclear envelope invagination-associated late endosomes. Oncotarget 2017;8:14443-61.
116. Ohki S, Flanagan TD, Hoekstra D. Probe transfer with and without membrane fusion in a fluorescence fusion assay. Biochemistry 1998;37:7496-503.
117. Costafreda MI, Abbasi A, Lu H, Kaplan G. Exosome mimicry by a HAVCR1-NPC1 pathway of endosomal fusion mediates hepatitis A virus infection. Nat Microbiol 2020;5:1096-106.
118. Xia HF, Yu ZL, Zhang LJ, et al. Real-Time dissection of the transportation and miRNA-release dynamics of small extracellular vesicles. Adv Sci 2023;10:e2205566.
119. 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.
120. Hall MP, Unch J, Binkowski BF, et al. Engineered luciferase reporter from a deep sea shrimp utilizing a novel imidazopyrazinone substrate. ACS Chem Biol 2012;7:1848-57.
121. Somiya M, Kuroda SI. Verification of extracellular vesicle-mediated functional mRNA delivery via RNA editing. bioRxiv 2022.
122. Dixon AS, Schwinn MK, Hall MP, et al. NanoLuc complementation reporter optimized for accurate measurement of protein interactions in cells. ACS Chem Biol 2016;11:400-8.
123. Wittrup A, Ai A, Liu X, et al. Visualizing lipid-formulated siRNA release from endosomes and target gene knockdown. Nat Biotechnol 2015;33:870-6.
124. Munson MJ, O'Driscoll G, Silva AM, et al. A high-throughput Galectin-9 imaging assay for quantifying nanoparticle uptake, endosomal escape and functional RNA delivery. Commun Biol 2021;4:211.
125. Du Rietz H, Hedlund H, Wilhelmson S, Nordenfelt P, Wittrup A. Imaging small molecule-induced endosomal escape of siRNA. Nat Commun 2020;11:1809.
126. de Jong OG, Murphy DE, Mäger I, et al. A CRISPR-Cas9-based reporter system for single-cell detection of extracellular vesicle-mediated functional transfer of RNA. Nat Commun 2020;11:1113.
127. Zomer A, Steenbeek SC, Maynard C, van Rheenen J. Studying extracellular vesicle transfer by a Cre-loxP method. Nat Protoc 2016;11:87-101.
128. Zomer A, Maynard C, Verweij FJ, et al. In Vivo imaging reveals extracellular vesicle-mediated phenocopying of metastatic behavior. Cell 2015;161:1046-57.
129. Mateescu B, Kowal EJ, van Balkom BW, et al. Obstacles and opportunities in the functional analysis of extracellular vesicle RNA - an ISEV position paper. J Extracell Vesicles 2017;6:1286095.
130. Gámez-Valero A, Monguió-Tortajada M, Carreras-Planella L, Franquesa Ml, Beyer K, Borràs FE. Size-exclusion chromatography-based isolation minimally alters extracellular vesicles' characteristics compared to precipitating agents. Sci Rep 2016;6:33641.
131. Jones DM, Padilla-Parra S. The beta-lactamase assay: harnessing a FRET Biosensor to analyse viral fusion mechanisms. Sensors 2016;16:950.
132. Squillace DM, Zhao Z, Call GM, Gao J, Yao JQ. Viral Inactivation of human osteochondral grafts with methylene blue and light. Cartilage 2014;5:28-36.
133. Quijada NM, Fongaro G, Barardi CR, Hernández M, Rodríguez-Lázaro D. Propidium monoazide integrated with qPCR enables the detection and enumeration of infectious enteric RNA and DNA viruses in clam and fermented sausages. Front Microbiol 2016;7:2008.
134. 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.
135. Murphy DE, de Jong OG, Evers MJW, Nurazizah M, Schiffelers RM, Vader P. Natural or synthetic RNA delivery: a stoichiometric comparison of extracellular vesicles and synthetic nanoparticles. Nano Lett 2021;21:1888-95.
136. Chevillet JR, Kang Q, Ruf IK, et al. Quantitative and stoichiometric analysis of the microRNA content of exosomes. Proc Natl Acad Sci USA 2014;111:14888-93.
137. He D, Wang H, Ho SL, et al. Total internal reflection-based single-vesicle in situ quantitative and stoichiometric analysis of tumor-derived exosomal microRNAs for diagnosis and treatment monitoring. Theranostics 2019;9:4494-507.
138. Lu M, Zhao X, Xing H, et al. Comparison of exosome-mimicking liposomes with conventional liposomes for intracellular delivery of siRNA. Int J Pharm 2018;550:100-13.
139. Schindler C, Collinson A, Matthews C, et al. Exosomal delivery of doxorubicin enables rapid cell entry and enhanced in vitro potency. PLoS One 2019;14:e0214545.
140. Gilleron J, Querbes W, Zeigerer A, et al. Image-based analysis of lipid nanoparticle-mediated siRNA delivery, intracellular trafficking and endosomal escape. Nat Biotechnol 2013;31:638-46.
141. Lara P, Chan AB, Cruz LJ, Quest AFG, Kogan MJ. Exploiting the natural properties of extracellular vesicles in targeted delivery towards specific cells and tissues. Pharmaceutics 2020;12:1022.
142. Ravichandran R, Bansal S, Rahman M, et al. The role of donor-derived exosomes in lung allograft rejection. Hum Immunol 2019;80:588-94.
143. Meijering BD, Juffermans LJ, van Wamel A, et al. Ultrasound and microbubble-targeted delivery of macromolecules is regulated by induction of endocytosis and pore formation. Circ Res 2009;104:679-87.
144. Nguyen VH, Lee BJ. Protein corona: a new approach for nanomedicine design. Int J Nanomedicine 2017;12:3137-51.
145. Aliyandi A, Zuhorn IS, Salvati A. Disentangling biomolecular corona interactions with cell receptors and implications for targeting of nanomedicines. Front Bioeng Biotechnol 2020;8:599454.
146. Aliyandi A, Reker-Smit C, Bron R, Zuhorn IS, Salvati A. Correlating Corona composition and cell uptake to identify proteins affecting nanoparticle entry into endothelial cells. ACS Biomater Sci Eng 2021;7:5573-84.
147. Tóth EÁ, Turiák L, Visnovitz T, et al. Formation of a protein corona on the surface of extracellular vesicles in blood plasma. J Extracell Vesicles 2021;10:e12140.
