REFERENCES
1. Kalluri R, Lebleu VS. The biology, function, and biomedical applications of exosomes. Science. 2020;367:eaau6977.
2. Toyofuku M, Nomura N, Eberl L. Types and origins of bacterial membrane vesicles. Nat Rev Microbiol. 2018;17:13-24.
3. Díaz‐Garrido N, Badia J, Baldomà L. Microbiota‐derived extracellular vesicles in interkingdom communication in the gut. J Extracell Vesicle. 2021;10:e12161.
4. Zakharzhevskaya NB, Vanyushkina AA, Altukhov IA, et al. Outer membrane vesicles secreted by pathogenic and nonpathogenic Bacteroides fragilis represent different metabolic activities. Sci Rep. 2017;7:5008.
5. Hendrix A, Lippens L, Pinheiro C, et al. Extracellular vesicle analysis. Nat Rev Methods Primers. 2023;3:56.
6. 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 Vesicle. 2018;7:1535750.
7. György B, Szabó TG, Pásztói M, et al. Membrane vesicles, current state-of-the-art: emerging role of extracellular vesicles. Cell Mol Life Sci. 2011;68:2667-88.
8. Lötvall J, Hill AF, Hochberg F, et al. Minimal experimental requirements for definition of extracellular vesicles and their functions: a position statement from the International Society for Extracellular Vesicles. J Extracell Vesicle. 2014;3:26913.
9. R. Belak Z, Harkness T, H. Eskiw C. A rapid, high-throughput method for determining chronological lifespan in budding yeast. JBM. 2018;5:1.
10. Kim JH, Lee J, Park J, Gho YS. Gram-negative and Gram-positive bacterial extracellular vesicles. Semin Cell Dev Biol. 2015;40:97-104.
11. Li P, Kaslan M, Lee SH, Yao J, Gao Z. Progress in exosome isolation techniques. Theranostics. 2017;7:789-804.
12. Balhuizen MD, Veldhuizen EJA, Haagsman HP. Outer membrane vesicle induction and isolation for vaccine development. Front. Microbiol. 2021;12:629090.
13. Gardiner C, Di Vizio D, Sahoo S, et al. Techniques used for the isolation and characterization of extracellular vesicles: results of a worldwide survey. J Extracell Vesicle. 2016;5:32945.
16. Onódi Z, Pelyhe C, Terézia Nagy C, et al. Isolation of high-purity extracellular vesicles by the combination of iodixanol density gradient ultracentrifugation and bind-elute chromatography from blood plasma. Front. Physiol. 2018;9:1479.
17. Zhang M, Viennois E, Prasad M, et al. Edible ginger-derived nanoparticles: a novel therapeutic approach for the prevention and treatment of inflammatory bowel disease and colitis-associated cancer. Biomaterials. 2016;101:321-40.
18. He L, Zhu D, Wang J, Wu X. A highly efficient method for isolating urinary exosomes. Int J Mol Med. 2018.
19. Klimentová J, Stulík J. Methods of isolation and purification of outer membrane vesicles from gram-negative bacteria. Microbiol Res. 2015;170:1-9.
20. Northrop‐Albrecht EJ, Taylor WR, Huang BQ, Kisiel JB, Lucien F. Assessment of extracellular vesicle isolation methods from human stool supernatant. J Extracell Vesicle. 2022;11:e12208.
21. Taylor DD, Shah S. Methods of isolating extracellular vesicles impact down-stream analyses of their cargoes. Methods. 2015;87:3-10.
22. Xu K, Jin Y, Li Y, Huang Y, Zhao R. Recent progress of exosome isolation and peptide recognition-guided strategies for exosome research. Front. Chem. 2022;10:844124.
23. Shao H, Im H, Castro CM, Breakefield X, Weissleder R, Lee H. New technologies for analysis of extracellular vesicles. Chem Rev. 2018;118:1917-50.
24. Castillo‐Romero KF, Santacruz A, González‐Valdez J. Production and purification of bacterial membrane vesicles for biotechnology applications: challenges and opportunities. Electrophoresis. 2022;44:107-24.
25. Aytar Çelik P, Derkuş B, Erdoğan K, et al. Bacterial membrane vesicle functions, laboratory methods, and applications. Biotechnol Adv. 2022;54:107869.
26. Wei S, Jiao D, Xing W. A rapid method for isolation of bacterial extracellular vesicles from culture media using epsilon-poly-L-lysine that enables immunological function research. Front. Immunol. 2022;13:930510.
27. Alves NJ, Turner KB, Divito KA, Daniele MA, Walper SA. Affinity purification of bacterial outer membrane vesicles (OMVs) utilizing a His-tag mutant. Res Microbiol. 2017;168:139-46.
28. De Sousa KP, Rossi I, Abdullahi M, Ramirez MI, Stratton D, Inal JM. Isolation and characterization of extracellular vesicles and future directions in diagnosis and therapy. WIREs Nanomed Nanobiotechnol. 2022;15:e1835.
29. 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.
30. Li Z, Yan J, Li X, et al. Advancements in extracellular vesicles biomanufacturing: a comprehensive overview of large-scale production and clinical research. Front. Bioeng. Biotechnol. 2025;13:1487627.
31. Shin H, Han C, Labuz JM, et al. High-yield isolation of extracellular vesicles using aqueous two-phase system. Sci Rep. 2015;5:13103.
32. Davies RT, Kim J, Jang SC, Choi E, Gho YS, Park J. Microfluidic filtration system to isolate extracellular vesicles from blood. Lab Chip. 2012;12:5202.
33. Kanwar SS, Dunlay CJ, Simeone DM, Nagrath S. Microfluidic device (ExoChip) for on-chip isolation, quantification and characterization of circulating exosomes. Lab Chip. 2014;14:1891-900.
34. Le Gall L, Ouandaogo ZG, Anakor E, et al. Optimized method for extraction of exosomes from human primary muscle cells. Skeletal Muscle. 2020;10:20.
35. Tauro BJ, Greening DW, Mathias RA, et al. Comparison of ultracentrifugation, density gradient separation, and immunoaffinity capture methods for isolating human colon cancer cell line LIM1863-derived exosomes. Methods. 2012;56:293-304.
36. Paolini L, Zendrini A, Noto GD, et al. Residual matrix from different separation techniques impacts exosome biological activity. Sci Rep. 2016;6:23550.
37. Chen Y, Zhu Q, Cheng L, et al. Exosome detection via the ultrafast-isolation system: EXODUS. Nat Methods. 2021;18:212-8.
38. Li W, Chen H, Tong M, Niu J, Zhu X, Lin L. Comparison of the yield and purity of plasma exosomes extracted by ultracentrifugation, precipitation, and membrane-based approaches. Open Chem. 2022;20:182-91.
39. Yang D, Zhang W, Zhang H, et al. Progress, opportunity, and perspective on exosome isolation - efforts for efficient exosome-based theranostics. Theranostics. 2020;10:3684-707.
40. Elsharkasy OM, Nordin JZ, Hagey DW, et al. Extracellular vesicles as drug delivery systems: Why and how? Adv Drug Delivery Rev. 2020;159:332-43.
41. Witwer KW, Wolfram J. Extracellular vesicles versus synthetic nanoparticles for drug delivery. Nat Rev Mater. 2021;6:103-6.
42. Fang X, Chen C, Liu B, et al. A magnetic bead-mediated selective adsorption strategy for extracellular vesicle separation and purification. Acta Biomater. 2021;124:336-47.
43. Zarovni N, Corrado A, Guazzi P, et al. Integrated isolation and quantitative analysis of exosome shuttled proteins and nucleic acids using immunocapture approaches. Methods. 2015;87:46-58.
44. Nordin JZ, Lee Y, Vader P, et al. Ultrafiltration with size-exclusion liquid chromatography for high yield isolation of extracellular vesicles preserving intact biophysical and functional properties. Nanomed-Nanotechnol. 2015;11:879-83.
45. Théry C, Amigorena S, Raposo G, Clayton A. Isolation and characterization of exosomes from cell culture supernatants and biological fluids. CP Cell Biology. 2006;30.
46. Drożdż A, Kamińska A, Surman M, et al. Low-vacuum filtration as an alternative extracellular vesicle concentration method: a comparison with ultracentrifugation and differential centrifugation. Pharmaceutics. 2020;12:872.
47. 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.
48. Pauwels MJ, Vandendriessche C, Vandenbroucke RE. Special delEVery: extracellular vesicles as promising delivery platform to the brain. Biomedicines. 2021;9:1734.
49. Kelwick RJR, Webb AJ, Heliot A, Segura CT, Freemont PS. Opportunities to accelerate extracellular vesicle research with cell‐free synthetic biology. J of Extracellular Bio. 2023;2:e90.
50. Jangam TC, Desai SA, Patel VP, Pagare NB, Raut ND. Exosomes as therapeutic and diagnostic tools: advances, challenges, and future directions. Cell Biochem Biophys. 2025;83:2805-29.
51. Casas A, Bultelle M, Motraghi C, Kitney R. Removing the bottleneck: introducing cmatch - a lightweight tool for construct-matching in synthetic biology. Front. Bioeng. Biotechnol. 2022;9:785131.
52. Plahar HA, Rich TN, Lane SD, et al. BioParts - a biological parts search portal and updates to the ICE parts registry software platform. ACS Synth. Biol. 2021;10:2649-60.
53. Haines MC, Carling B, Marshall J, et al. Basicsynbio and the BASIC SEVA collection: software and vectors for an established DNA assembly method. Synth Biol. 2022;7:ysac023.
54. Kelwick R, Macdonald JT, Webb AJ, Freemont P. Developments in the tools and methodologies of synthetic biology. Front. Bioeng. Biotechnol. 2014;2:60.
55. Komor AC, Kim YB, Packer MS, Zuris JA, Liu DR. Programmable editing of a target base in genomic DNA without double-stranded DNA cleavage. Nature. 2016;533:420-4.
56. Anzalone AV, Randolph PB, Davis JR, et al. Search-and-replace genome editing without double-strand breaks or donor DNA. Nature. 2019;576:149-57.
57. Abudayyeh OO, Gootenberg JS, Konermann S, et al. C2c2 is a single-component programmable RNA-guided RNA-targeting CRISPR effector. Science. 2016;353:aaf5573.
58. Campa CC, Weisbach NR, Santinha AJ, Incarnato D, Platt RJ. Multiplexed genome engineering by Cas12a and CRISPR arrays encoded on single transcripts. Nat Methods. 2019;16:887-93.
59. Zhang Y, Wang J, Wang Z, et al. A gRNA-tRNA array for CRISPR-Cas9 based rapid multiplexed genome editing in Saccharomyces cerevisiae. Nat Commun. 2019;10:1053.
60. Zanella I, König E, Tomasi M, et al. Proteome‐minimized outer membrane vesicles from Escherichia coli as a generalized vaccine platform. J Extracell Vesicle. 2021;10:e12066.
61. Rumah BL, Claxton Stevens BH, Yeboah JE, et al. In vivo genome editing in type I and II methanotrophs using a CRISPR/Cas9 system. ACS Synth. Biol. 2023;12:544-54.
62. Xu Z, Qiao S, Wang Z, et al. PMA1-containing extracellular vesicles of Candida albicans triggers immune responses and colitis progression. Gut Microbes. 2025;17:2455508.
63. Jia B, Wu Y, Li B, et al. Precise control of SCRaMbLE in synthetic haploid and diploid yeast. Nat Commun. 2018;9:1933.
64. Zhou S, Wu Y, Zhao Y, et al. Dynamics of synthetic yeast chromosome evolution shaped by hierarchical chromatin organization. Natl Sci Rev. 2023;10:nwad073.
65. Sawabe T, Ojima Y, Nakagawa M, et al. Construction and characterization of a hypervesiculation strain of Escherichia coli Nissle 1917. PLoS ONE. 2024;19:e0301613.
66. Schetters ST, Jong WS, Horrevorts SK, et al. Outer membrane vesicles engineered to express membrane-bound antigen program dendritic cells for cross-presentation to CD8+ T cells. Acta Biomater. 2019;91:248-57.
67. Rappazzo CG, Watkins HC, Guarino CM, et al. Recombinant M2e outer membrane vesicle vaccines protect against lethal influenza A challenge in BALB/c mice. Vaccine. 2016;34:1252-8.
68. Zhou J, Li M, Chen Q, et al. Programmable probiotics modulate inflammation and gut microbiota for inflammatory bowel disease treatment after effective oral delivery. Nat Commun. 2022;13:3432.
69. Cao Z, Wang X, Pang Y, Cheng S, Liu J. Biointerfacial self-assembly generates lipid membrane coated bacteria for enhanced oral delivery and treatment. Nat Commun. 2019;10:5783.
70. Cui M, Sun T, Li S, et al. NIR light-responsive bacteria with live bio-glue coatings for precise colonization in the gut. Cell Rep. 2021;36:109690.
71. Pan H, Sun T, Cui M, et al. Light-sensitive Lactococcus lactis for microbe-gut-brain axis regulating via upconversion optogenetic micro-nano system. ACS Nano. 2022;16:6049-63.
72. Cui M, Ling W, Zhang L, et al. Smartphone bioelectronic drug with visual colorimetric sensor and bulk nanoencapsulation optogenetic bacteria for chronic kidney disease theragnostics. Chem Eng J. 2023;451:138812.
73. Zhang X, Ma N, Ling W, et al. A micro-nano optogenetic system based on probiotics for in situ host metabolism regulation. Nano Res. 2022;16:2829-39.
74. Brown L, Wolf JM, Prados-rosales R, Casadevall A. Through the wall: extracellular vesicles in Gram-positive bacteria, mycobacteria and fungi. Nat Rev Microbiol. 2015;13:620-30.
75. Zhao K, Bleackley M, Chisanga D, et al. Extracellular vesicles secreted by Saccharomyces cerevisiae are involved in cell wall remodelling. Commun Biol. 2019;2:305.
76. Chen J, Liu M, Chen S, et al. Engineered therapeutic bacteria with high-yield membrane vesicle production inspired by eukaryotic membrane curvature for treating inflammatory bowel disease. ACS Nano. 2025;19:2405-18.
77. Wang X, Singh AK, Zhang X, Sun W. Induction of protective antiplague immune responses by self-adjuvanting bionanoparticles derived from engineered yersinia pestis. Infect Immun. 2020;88:e00081-20.
78. Takaki K, Tahara YO, Nakamichi N, et al. Multilamellar and multivesicular outer membrane vesicles produced by a Buttiauxella agrestis tolB mutant. Appl Environ Microbiol. 2020;86:e01131-20.
79. Nevermann J, Silva A, Otero C, et al. Identification of genes involved in biogenesis of outer membrane vesicles (OMVs) in salmonella enterica serovar typhi. Front. Microbiol. 2019;10:104.
80. Irene C, Fantappiè L, Caproni E, et al. Bacterial outer membrane vesicles engineered with lipidated antigens as a platform for Staphylococcus aureus vaccine. Proc. Natl. Acad. Sci. U.S.A. 2019;116:21780-8.
81. Needham BD, Carroll SM, Giles DK, Georgiou G, Whiteley M, Trent MS. Modulating the innate immune response by combinatorial engineering of endotoxin. Proc. Natl. Acad. Sci. U.S.A. 2013;110:1464-9.
82. Van Der Ley P, Steeghs L, Hamstra HJ, Ten Hove J, Zomer B, Van Alphen L. Modification of lipid a biosynthesis in Neisseria meningitidis lpxL mutants: influence on lipopolysaccharide structure, toxicity, and adjuvant activity. Infect Immun. 2001;69:5981-90.
83. Gujrati V, Prakash J, Malekzadeh-Najafabadi J, et al. Bioengineered bacterial vesicles as biological nano-heaters for optoacoustic imaging. Nat Commun. 2019;10:1114.
84. Jumper J, Evans R, Pritzel A, et al. Highly accurate protein structure prediction with AlphaFold. Nature. 2021;596:583-9.
85. Lin Z, Akin H, Rao R, et al. Evolutionary-scale prediction of atomic-level protein structure with a language model. Science. 2023;379:1123-30.
86. Wang J, Lisanza S, Juergens D, et al. Scaffolding protein functional sites using deep learning. Science. 2022;377:387-94.
87. Gujrati V, Kim S, Kim S, et al. Bioengineered bacterial outer membrane vesicles as cell-specific drug-delivery vehicles for cancer therapy. ACS Nano. 2014;8:1525-37.
88. Jiang L, Driedonks TA, Jong WS, et al. A bacterial extracellular vesicle‐based intranasal vaccine against SARS‐CoV‐2 protects against disease and elicits neutralizing antibodies to wild‐type and Delta variants. J Extracell Vesicle. 2022;11:e12192.
89. Ren C, Li Y, Cong Z, Li Z, Xie L, Wu S. Bioengineered bacterial outer membrane vesicles encapsulated Polybia-mastoparan I fusion peptide as a promising nanoplatform for bladder cancer immune-modulatory chemotherapy. Front. Immunol. 2023;14:1129771.
90. Sung BH, Von Lersner A, Guerrero J, et al. A live cell reporter of exosome secretion and uptake reveals pathfinding behavior of migrating cells. Nat Commun. 2020;11:2092.
91. Liu H, Song P, Zhang H, et al. Synthetic biology‐based bacterial extracellular vesicles displaying BMP‐2 and CXCR4 to ameliorate osteoporosis. J Extracell Vesicle. 2024;13:e12429.
92. Ivanova A, Badertscher L, O’Driscoll G, et al. Creating designer engineered extracellular vesicles for diverse ligand display, target recognition, and controlled protein loading and delivery. Adv Sci. 2023;10:2304389.
93. Sun J, Tan L, Ye B, Bi X. Engineered outer membrane vesicles as nanosized immune cell engagers for enhanced solid tumor immunotherapy. ACS Nano. 2024;18:30332-44.
94. Han D, Wang F, Ma Y, et al. Redirecting antigens by engineered photosynthetic bacteria and derived outer membrane vesicles for enhanced cancer immunotherapy. ACS Nano. 2023;17:18716-31.
95. Cui C, Guo T, Zhang S, et al. Bacteria-derived outer membrane vesicles engineered with over-expressed pre-miRNA as delivery nanocarriers for cancer therapy. Nanomed-Nanotechnol. 2022;45:102585.
96. Santos NL, Bustos SO, Reis PP, Chammas R, Andrade LNS. Extracellular vesicle-packaged miR-195-5p sensitizes melanoma to targeted therapy with kinase inhibitors. Cells. 2023;12:1317.
97. Sork H, Corso G, Krjutskov K, et al. Heterogeneity and interplay of the extracellular vesicle small RNA transcriptome and proteome. Sci Rep. 2018;8:10813.
98. You Y, Tian Y, Yang Z, et al. Intradermally delivered mRNA-encapsulating extracellular vesicles for collagen-replacement therapy. Nat. Biomed. Eng. 2023;7:887-900.
99. Cheng K, Zhao R, Li Y, et al. Bioengineered bacteria-derived outer membrane vesicles as a versatile antigen display platform for tumor vaccination via Plug-and-Display technology. Nat Commun. 2021;12:2041.
100. Yue Y, Xu J, Li Y, et al. Antigen-bearing outer membrane vesicles as tumour vaccines produced in situ by ingested genetically engineered bacteria. Nat. Biomed. Eng. 2022;6:898-909.
101. Zhuang W, Wang Y, Lei Y, et al. Phytochemical engineered bacterial outer membrane vesicles for photodynamic effects promoted immunotherapy. Nano Lett. 2022;22:4491-500.
102. Shen H, Chen S, Zhang C, et al. Engineered delivery platform based on bacterial outer membrane vesicles for efficient labeling of 131I and improved radio-immunotherapy. Chem Eng J. 2024;490:151837.
103. Wang D, Liu C, You S, et al. Bacterial vesicle-cancer cell hybrid membrane-coated nanoparticles for tumor specific immune activation and photothermal therapy. ACS Appl Mater Interfaces. 2020;12:41138-47.
104. Rezaei Adriani R, Mousavi Gargari SL, Bakherad H, Amani J. Anti-EGFR bioengineered bacterial outer membrane vesicles as targeted immunotherapy candidate in triple-negative breast tumor murine model. Sci Rep. 2023;13:16403.
105. Sepahdar Z, Miroliaei M, Bouzari S, Khalaj V, Salimi M. Surface engineering of Escherichia coli-derived OMVs as promising nano-carriers to target EGFR-overexpressing breast cancer cells. Front. Pharmacol. 2021;12:719289.
106. Thomas SC, Madaan T, Kamble NS, Siddiqui NA, Pauletti GM, Kotagiri N. Engineered bacteria enhance immunotherapy and targeted therapy through stromal remodeling of tumors. Adv Healthc Mater. 2021;11:2101487.
107. Murase K. Cytolysin A (ClyA): a bacterial virulence factor with potential applications in nanopore technology, vaccine development, and tumor therapy. Toxins. 2022;14:78.
108. Zaruba M, Roschitz L, Sami H, Ogris M, Gerner W, Metzner C. Surface modification of E. coli outer membrane vesicles with glycosylphosphatidylinositol-anchored proteins: generating pro/eukaryote chimera constructs. Membranes. 2021;11:428.
109. Chen T, Wang K, Chi X, et al. Construction of a bacterial surface display system based on outer membrane protein F. Microb Cell Fact. 2019;18:70.
110. Tian T, Zhang H, He C, et al. Surface functionalized exosomes as targeted drug delivery vehicles for cerebral ischemia therapy. Biomaterials. 2018;150:137-49.
111. Qi H, Liu C, Long L, et al. Blood exosomes endowed with magnetic and targeting properties for cancer therapy. ACS Nano. 2016;10:3323-33.
112. Kim HK, Choi Y, Kim KH, et al. Scalable production of siRNA‐encapsulated extracellular vesicles for the inhibition of KRAS‐mutant cancer using acoustic shock waves. J Extracell Vesicle. 2024;13:e12508.
113. Yang X, Shi G, Lin Z, et al. Pathogen-targeting biomineralized bacterial outer membrane vesicles for eradicating both intracellular and extracellular Staphylococcus aureus. J Controlled Release. 2025;382:113702.
114. Gerritzen MJH, Maas RHW, Van Den Ijssel J, et al. High dissolved oxygen tension triggers outer membrane vesicle formation by Neisseria meningitidis. Microb Cell Fact. 2018;17:157.
115. Wang Y, Fan Y, Zhang X, et al. In situ production and precise release of bioactive GM-CSF and siRNA by engineered bacteria for macrophage reprogramming in cancer immunotherapy. Biomaterials. 2025;317:123037.
116. Li Q, Zhang Z, Wang F, et al. Reversible zwitterionic coordination enables rapid, high-yield, and high-purity isolation of extracellular vesicles from biofluids. Sci. Adv. 2023;9:eadf4568.
117. Ye Z, Ji M, Wu K, et al. In‐sequence high-specificity dual-reporter unlocking of fluorescent probe enables the precise identification of atherosclerotic plaques. Angew Chem Int Ed. 2022;61:e202204518.
118. Han C, Zhang X, Pang G, et al. Hydrogel microcapsules containing engineered bacteria for sustained production and release of protein drugs. Biomaterials. 2022;287:121619.
119. Wang P, Lan G, Xu B, et al. α-Synuclein-carrying astrocytic extracellular vesicles in Parkinson pathogenesis and diagnosis. Transl Neurodegener. 2023;12:40.
120. Chen L, Valentine JL, Huang C, et al. Outer membrane vesicles displaying engineered glycotopes elicit protective antibodies. Proc. Natl. Acad. Sci. U.S.A. 2016;113.
121. Tian H, Li B, Xu T, et al. Outer Membrane vesicles derived from Salmonella enterica serotype typhimurium can deliver Shigella flexneri 2a O-polysaccharide antigen to prevent Shigella flexneri 2a infection in mice. Appl Environ Microbiol. 2021;87:e00968-21.
122. Van Den Berg Van Saparoea HB, Houben D, Kuijl C, Luirink J, Jong WSP. Combining protein ligation systems to expand the functionality of semi-synthetic outer membrane vesicle nanoparticles. Front. Microbiol. 2020;11:890.
123. Gao J, Su Y, Wang Z. Engineering bacterial membrane nanovesicles for improved therapies in infectious diseases and cancer. Adv Drug Delivery Rev. 2022;186:114340.
124. Brezgin S, Danilik O, Yudaeva A, et al. Basic guide for approaching drug delivery with extracellular vesicles. Int J Mol Sci. 2024;25:10401.
125. Feng L, Dong Z, Tao D, Zhang Y, Liu Z. The acidic tumor microenvironment: a target for smart cancer nano-theranostics. Natl Sci Rev. 2018;5:269-86.
126. Jan K, Hassan N, James A, Hussain I, Rashid SM. Exploring molecular targets in cancer: unveiling the anticancer potential of Paeoniflorin through a comprehensive analysis of diverse signaling pathways and recent advances. JBM. 2024;11:e99010014.
127. Kim OY, Park HT, Dinh NTH, et al. Bacterial outer membrane vesicles suppress tumor by interferon-γ-mediated antitumor response. Nat Commun. 2017;8:626.
128. Chen Q, Huang G, Wu W, et al. A hybrid eukaryotic-prokaryotic nanoplatform with photothermal modality for enhanced antitumor vaccination. Adv Mater. 2020;32:1908185.
129. Sagnella SM, Trieu J, Brahmbhatt H, et al. Targeted doxorubicin-loaded bacterially derived nano-cells for the treatment of neuroblastoma. Mol Cancer Ther. 2018;17:1012-23.
130. Chen Q, Bai H, Wu W, et al. Bioengineering bacterial vesicle-coated polymeric nanomedicine for enhanced cancer immunotherapy and metastasis prevention. Nano Lett. 2019;20:11-21.
131. Gu T, Wang M, Niu J, Chu Y, Guo K, Peng L. Outer membrane vesicles derived from E. coli as novel vehicles for transdermal and tumor targeting delivery. Nanoscale. 2020;12:18965-77.
132. Maerz JK, Steimle A, Lange A, Bender A, Fehrenbacher B, Frick J. Outer membrane vesicles blebbing contributes to B. Vulgatus mpk-mediated immune response silencing. Gut Microbes. 2017;9:1-12.
133. Seo M, Park E, Ko S, Choi E, Kim S. Therapeutic effects of kefir grain Lactobacillus-derived extracellular vesicles in mice with 2,4,6-trinitrobenzene sulfonic acid-induced inflammatory bowel disease. J Dairy Sci. 2018;101:8662-71.
134. Kim M, Choi SJ, Choi H, et al. Lactobacillus plantarum-derived extracellular vesicles protect atopic dermatitis induced by Staphylococcus aureus-derived extracellular vesicles. Allergy Asthma Immunol Res. 2018;10:516.
135. Dean SN, Rimmer MA, Turner KB, et al. Lactobacillus acidophilus membrane vesicles as a vehicle of bacteriocin delivery. Front. Microbiol. 2020;11:710.
136. Wang X, Eagen WJ, Lee JC. Orchestration of human macrophage NLRP3 inflammasome activation by Staphylococcus aureus extracellular vesicles. Proc. Natl. Acad. Sci. U.S.A. 2020;117:3174-84.
137. Van Der Ley P, Schijns VE. Outer membrane vesicle-based intranasal vaccines. Curr Opin Immunol. 2023;84:102376.
138. Holst J, Martin D, Arnold R, et al. Properties and clinical performance of vaccines containing outer membrane vesicles from Neisseria meningitidis. Vaccine. 2009;27:B3-B12.
139. Li Y, Ma X, Yue Y, et al. Rapid surface display of mRNA antigens by bacteria-derived outer membrane vesicles for a personalized tumor vaccine. Adv Mater. 2022;34:2109984.
140. Schaack B, Hindré T, Quansah N, Hannani D, Mercier C, Laurin D. Microbiota-derived extracellular vesicles detected in human blood from healthy donors. Int J Mol Sci. 2022;23:13787.
141. Yoo JY, Rho M, You Y, et al. 16S rRNA gene-based metagenomic analysis reveals differences in bacteria-derived extracellular vesicles in the urine of pregnant and non-pregnant women. Exp Mol Med. 2016;48:e208.
142. Han P, Bartold PM, Salomon C, Ivanovski S. Salivary outer membrane vesicles and DNA methylation of small extracellular vesicles as biomarkers for periodontal status: a pilot study. Int J Mol Sci. 2021;22:2423.
143. Yoon H, Kim N, Park J, et al. Analysis of the gut microbiome using extracellular vesicles in the urine of patients with colorectal cancer. Korean J Intern Med. 2023;38:27-38.
144. Liu C, Yazdani N, Moran CS, et al. Unveiling clinical applications of bacterial extracellular vesicles as natural nanomaterials in disease diagnosis and therapeutics. Acta Biomater. 2024;180:18-45.
145. Liang L, Yang C, Liu L, et al. Commensal bacteria-derived extracellular vesicles suppress ulcerative colitis through regulating the macrophages polarization and remodeling the gut microbiota. Microb Cell Fact. 2022;21:88.
146. Meng F, Li L, Zhang Z, et al. Biosynthetic neoantigen displayed on bacteria derived vesicles elicit systemic antitumour immunity. J Extracell Vesicle. 2022;11:12289.
147. Liu H, Li M, Zhang T, et al. Engineered bacterial extracellular vesicles for osteoporosis therapy. Chem Eng J. 2022;450:138309.
148. Liu H, Wu Y, Wang F, et al. Bone-targeted engineered bacterial extracellular vesicles delivering miRNA to treat osteoporosis. Compos Part B-Eng. 2023;267:111047.
149. J. Jansen G, P. Schouten G, Wiersma M. Advancements in analytical methods for studying the human gut microbiome. JBM. 2024;12:e99010038.
150. Liu JH, Chen CY, Liu ZZ, et al. Extracellular vesicles from child gut microbiota enter into bone to preserve bone mass and strength. Adv Sci. 2021;8:2004831.
151. Wang T, Mo L, Ou J, et al. Proteus mirabilis vesicles induce mitochondrial apoptosis by regulating miR96-5p/Abca1 to inhibit Osteoclastogenesis and bone loss. Front. Immunol. 2022;13:833040.
152. Han F, Wang K, Shen K, et al. Extracellular vesicles from Lactobacillus druckerii inhibit hypertrophic scar fibrosis. J Nanobiotechnol. 2023;21:113.
153. Zhou H, Tan X, Chen G, et al. Extracellular vesicles of commensal skin microbiota alleviate cutaneous inflammation in atopic dermatitis mouse model by re-establishing skin homeostasis. J Investig Dermatol. 2025;145:312-322.e9.
154. Chen Y, Huang X, Liu A, et al. Lactobacillus Reuteri vesicles regulate mitochondrial function of macrophages to promote mucosal and cutaneous wound healing. Adv Sci. 2024;11:2309725.
155. Chen Y, Ou Z, Pang M, et al. Extracellular vesicles derived from Akkermansia muciniphila promote placentation and mitigate preeclampsia in a mouse model. J Extracell Vesicle. 2023;12:12328.
156. Choi J, Kwon H, Kim Y, Han P. Extracellular vesicles from gram-positive and gram-negative probiotics remediate stress-induced depressive behavior in mice. Mol Neurobiol. 2022;59:2715-28.
157. Lee DH, Park HK, Lee HR, et al. Immunoregulatory effects of Lactococcus lactis‐derived extracellular vesicles in allergic asthma. Clin Transl Allergy. 2022;12:e12138.
158. Wu F, Ren Y, Lv W, et al. Generating dual structurally and functionally skin-mimicking hydrogels by crosslinking cell-membrane compartments. Nat Commun. 2024;15:802.
159. Wei D, Dao J, Liu H, Chen G. Suspended polyhydroxyalkanoate microspheres as 3D carriers for mammalian cell growth. Artif Cell Nanomed B. 2018;46:473-83.
160. Wei DX, Dao JW, Chen GQ. A micro-ark for cells: highly open porous polyhydroxyalkanoate microspheres as injectable scaffolds for tissue regeneration. Adv Mater. 2018;30:1802273.
161. Yang Y, Huang Y, Yang J, et al. Umbilical cord mesenchymal stem cell-derived exosomes promote wound healing and skin regeneration via the regulation of inflammation and angiogenesis. Front. Bioeng. Biotechnol. 2025;13:1641709.
162. Kang S, Jeon S, Baek H, et al. Lactobacillus-derived artificial extracellular vesicles for skin rejuvenation and prevention of photo-aging. Biomater. Sci. 2025;13:2026-35.
163. Tao Y, Ren H, Chen W, et al. Extracellular vesicles from Lactiplantibacillus plantarum: a novel approach to lowering 5-ASA dosage in ulcerative colitis. Food Bioscience. 2025;69:106917.
164. Koeppen K, Hampton TH, Jarek M, et al. A novel mechanism of host-pathogen interaction through sRNA in bacterial outer membrane vesicles. PLoS Pathog. 2016;12:e1005672.
165. Toyofuku M, Cárcamo-Oyarce G, Yamamoto T, et al. Prophage-triggered membrane vesicle formation through peptidoglycan damage in Bacillus subtilis. Nat Commun. 2017;8:481.







