Research advances in the mechanisms of infection and antibiotic resistance mediated by outer membrane vesicle_West China Medical Journal

Authors：

ZHANG Lun ^1,2 , JING Xiaopeng ¹ , LIU Yi ¹ ,  ZENG Ji ¹

1. Department of Clinical Laboratory, Wuhan Fourth Hospital, Wuhan, Hubei 430000, P. R. China;
2. Hubei Key Laboratory of Sports Injury and Precision Therapy, Wuhan Fourth Hospital, Wuhan, Hubei 430000, P. R. China;

Corresponding?author：

ZENG Ji, Email: whzjmicro@163.com

Keywords：

Outer membrane vesicle; infection; drug resistance; review

DOI：

10.7507/1002-0179.202508018

Video：

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Abstract Full text Figures/Tables Video References Cited by

Outer membrane vesicle (OMV) serves as a crucial carrier in bacterial communication and drug resistance transmission. This review summarizes the formation mechanisms and compositional characteristics of OMV regulated by outer membrane cross-linking, lipids, and signaling molecules. It focuses on the diverse mechanisms by which OMV mediate drug resistance, including horizontal transfer and expression regulation of resistance genes, enhanced efflux capabilities, involvement in biofilm formation, and modulation of host immune responses. Finally, it explores the research direction of OMV as novel anti-infection target, aiming to provide a theoretical basis for overcoming bacterial resistance challenges and developing new therapeutic strategies.

Citation： ZHANG Lun, JING Xiaopeng, LIU Yi, ZENG Ji. Research advances in the mechanisms of infection and antibiotic resistance mediated by outer membrane vesicle. West China Medical Journal, 2026, 41(7): 1207-1212. doi: 10.7507/1002-0179.202508018 Copy

1.	Aggarwal R, Mahajan P, Pandiya S, et al. Antibiotic resistance: a global crisis, problems and solutions[J]. Crit Rev Microbiol, 2024, 50(5): 896-921.
2.	Cecil JD, Sirisaengtaksin N, O’Brien-Simpson NM, et al. Outer membrane vesicle-host cell interactions[J]. Microbiol Spectr, 2019, 7(1): 10.
3.	Sartorio MG, Pardue EJ, Feldman MF, et al. Bacterial outer membrane vesicles: from discovery to applications[J]. Annu Rev Microbiol, 2021, 75: 609-630.
4.	Schwechheimer C, Kuehn MJ. Outer-membrane vesicles from Gram-negative bacteria: biogenesis and functions[J]. Nat Rev Microbiol, 2015, 13(10): 605-619.
5.	Kim JY, Suh JW, Kang JS, et al. Gram-negative bacteria’s outer membrane vesicles[J]. Infect Chemother, 2023, 55(1): 1-9.
6.	Schwechheimer C, Kulp A, Kuehn MJ. Modulation of bacterial outer membrane vesicle production by envelope structure and content[J]. BMC Microbiol, 2014, 14: 324.
7.	Cassin EK, Tseng BS. Pushing beyond the envelope: the potential roles of OprF in Pseudomonas aeruginosa biofilm formation and pathogenicity[J]. J Bacteriol, 2019, 201(18): e00050-19.
8.	Schwechheimer C, Rodriguez DL, Kuehn MJ. NlpI-mediated modulation of outer membrane vesicle production through peptidoglycan dynamics in Escherichia coli[J]. Microbiologyopen, 2015, 4(3): 375-389.
9.	Lappann M, Otto A, Becher D, et al. Comparative proteome analysis of spontaneous outer membrane vesicles and purified outer membranes of Neisseria meningitidis[J]. J Bacteriol, 2013, 195(19): 4425-4435.
10.	McBroom AJ, Kuehn MJ. Release of outer membrane vesicles by Gram-negative bacteria is a novel envelope stress response[J]. Mol Microbiol, 2007, 63(2): 545-558.
11.	Macdonald IA, Kuehn MJ. Stress-induced outer membrane vesicle production by Pseudomonas aeruginosa[J]. J Bacteriol, 2013, 195(13): 2971-2981.
12.	Roden JA, Wells DH, Chomel BB, et al. Hemin binding protein C is found in outer membrane vesicles and protects Bartonella henselae against toxic concentrations of hemin[J]. Infect Immun, 2012, 80(3): 929-942.
13.	McMahon KJ, Castelli ME, García Vescovi E, et al. Biogenesis of outer membrane vesicles in Serratia marcescens is thermoregulated and can be induced by activation of the Rcs phosphorelay system[J]. J Bacteriol, 2012, 194(12): 3241-3249.
14.	Frias A, Manresa A, de Oliveira E, et al. Membrane vesicles: a common feature in the extracellular matter of cold-adapted antarctic bacteria[J]. Microb Ecol, 2010, 59(3): 476-486.
15.	Chowdhury C, Jagannadham MV. Virulence factors are released in association with outer membrane vesicles of Pseudomonas syringae pv. tomato T1 during normal growth[J]. Biochim Biophys Acta, 2013, 1834(1): 231-239.
16.	Kulkarni HM, Swamy ChV, Jagannadham MV. Molecular characterization and functional analysis of outer membrane vesicles from the antarctic bacterium Pseudomonas syringae suggest a possible response to environmental conditions[J]. J Proteome Res, 2014, 13(3): 1345-1358.
17.	Tashiro Y, Inagaki A, Shimizu M, et al. Characterization of phospholipids in membrane vesicles derived from Pseudomonas aeruginosa[J]. Biosci Biotechnol Biochem, 2011, 75(3): 605-607.
18.	Murphy K, Park AJ, Hao Y, et al. Influence of O polysaccharides on biofilm development and outer membrane vesicle biogenesis in Pseudomonas aeruginosa PAO1[J]. J Bacteriol, 2014, 196(7): 1306-1317.
19.	Yamaguchi M, Sato K, Yukitake H, et al. A Porphyromonas gingivalis mutant defective in a putative glycosyltransferase exhibits defective biosynthesis of the polysaccharide portions of lipopolysaccharide, decreased gingipain activities, strong autoaggregation, and increased biofilm formation[J]. Infect Immun, 2010, 78(9): 3801-3812.
20.	Huang C, Cao W, Zhou S, et al. Biogenesis mechanisms, regulatory strategies, and applications of bacterial extracellular vesicles[J]. Crit Rev Biotechnol, 2025, 45(8): 1700-1716.
21.	Zhao X, Wei Y, Bu Y, et al. Review on bacterial outer membrane vesicles: structure, vesicle formation, separation and biotechnological applications[J]. Microb Cell Fact, 2025, 24(1): 27.
22.	Dai K, Liao B, Huang X, et al. Consistency in bacterial extracellular vesicle production: key to their application in human health[J]. Extracell Vesicles Circ Nucl Acids, 2025, 6(1): 1-20.
23.	Gon?alves MO, Pincela Lins PM, Kassab G, et al. Outer membrane vesicles (OMVs) and their therapeutic potential as anti-infectious agents[J]. Nano Trends, 2025, 11: 100129.
24.	Meena H, Nandy S, Patil VS, et al. Lipidomics in understanding microbial biofilms[C]//Omics approaches in biofilm research: perspectives and applications. Cham: Springer Nature Switzerland, 2025: 287-310.
25.	Li L, Xu X, Cheng P, et al. Klebsiella pneumoniae derived outer membrane vesicles mediated bacterial virulence, antibiotic resistance, host immune responses and clinical applications[J]. Virulence, 2025, 16(1): 2449722.
26.	Li P, Luo W, Xiang TX, et al. Horizontal gene transfer via OMVs co-carrying virulence and antimicrobial-resistant genes is a novel way for the dissemination of carbapenem-resistant hypervirulent Klebsiella pneumoniae[J]. Front Microbiol, 2022, 13: 945972.
27.	Dhurve G, Madikonda AK, Jagannadham MV, et al. Outer membrane vesicles of Acinetobacter baumannii DS002 are selectively enriched with TonB-dependent transporters and play a key role in iron acquisition[J]. Microbiol Spectr, 2022, 10(2): e0029322.
28.	Wachino JI. Horizontal gene transfer systems for spread of antibiotic resistance in Gram-negative bacteria[J]. Microbiol Immunol, 2025, 69(7): 367-376.
29.	Chen LJ, Jing XP, Meng DL, et al. Newly detected transmission of blaKPC-2 by outer membrane vesicles in Klebsiella pneumoniae[J]. Curr Med Sci, 2023, 43(1): 80-85.
30.	Dell’Annunziata F, Dell’Aversana C, Doti N, et al. Outer membrane vesicles derived from Klebsiella pneumoniae are a driving force for horizontal gene transfer[J]. Int J Mol Sci, 2021, 22(16): 8732.
31.	Wang M, Yao Y, Yang Y, et al. The characterization of outer membrane vesicles (OMVs) and their role in mediating antibiotic-resistance gene transfer through natural transformation in Riemerella anatipestifer[J]. Poult Sci, 2025, 104(2): 104730.
32.	He M, Yin S, Huang X, et al. Insights into the regulatory role of bacterial sncRNA and its extracellular delivery via OMVs[J]. Appl Microbiol Biotechnol, 2024, 108(1): 29.
33.	Parker A, Gottesman S. Small RNA regulation of TolC, the outer membrane component of bacterial multidrug transporters[J]. J Bacteriol, 2016, 198(7): 1101-1113.
34.	Gutierrez A, Laureti L, Crussard S, et al. β-Lactam antibiotics promote bacterial mutagenesis via an RpoS-mediated reduction in replication fidelity[J]. Nat Commun, 2013, 4: 1610.
35.	Nishino K, Yamasaki S, Hayashi-Nishino M, et al. Effect of overexpression of small non-coding DsrA RNA on multidrug efflux in Escherichia coli[J]. J Antimicrob Chemother, 2011, 66(2): 291-296.
36.	Sarshar M, Scribano D, Palamara AT, et al. The Acinetobacter baumannii model can explain the role of small non-coding RNAs as potential mediators of host-pathogen interactions[J]. Front Mol Biosci, 2022, 9: 1088783.
37.	Eyraud A, Tattevin P, Chabelskaya S, et al. A small RNA controls a protein regulator involved in antibiotic resistance in Staphylococcus aureus[J]. Nucleic Acids Res, 2014, 42(8): 4892-4905.
38.	Lucena ACR, Ferrarini MG, de Oliveira WK, et al. Modulation of Klebsiella pneumoniae outer membrane vesicle protein cargo under antibiotic treatment[J]. Biomedicines, 2023, 11(6): 1515.
39.	Park AJ, Surette MD, Khursigara CM. Antimicrobial targets localize to the extracellular vesicle-associated proteome of Pseudomonas aeruginosa grown in a biofilm[J]. Front Microbiol, 2014, 5: 464.
40.	Farrugia C, Stafford GP, Murdoch C. Porphyromonas gingivalis outer membrane vesicles increase vascular permeability[J]. J Dent Res, 2020, 99(13): 1494-1501.
41.	Wang J, Terrasse R, Bafna JA, et al. Electrophysiological characterization of transport across outer-membrane channels from Gram-negative bacteria in presence of lipopolysaccharides[J]. Angew Chem Int Ed Engl, 2020, 59(22): 8517-8521.
42.	Jasim R, Han ML, Zhu Y, et al. Lipidomic analysis of the outer membrane vesicles from paired polymyxin-susceptible and -resistant Klebsiella pneumoniae clinical isolates[J]. Int J Mol Sci, 2018, 19(8): 2356.
43.	Sáenz JP, Grosser D, Bradley AS, et al. Hopanoids as functional analogues of cholesterol in bacterial membranes[J]. Proc Natl Acad Sci USA, 2015, 112(38): 11971-11976.
44.	Kim SW, Park SB, Im SP, et al. Outer membrane vesicles from β-lactam-resistant Escherichia coli enable the survival of β-lactam-susceptible E. coli in the presence of β-lactam antibiotics[J]. Sci Rep, 2018, 8(1): 5402.
45.	Miao B, Wang D, Yu L, et al. Mechanism and nanotechnological-based therapeutics for tolerance and resistance of bacterial biofilms[J]. Microbiol Res, 2025, 292: 127987.
46.	Wang W, Chanda W, Zhong M. The relationship between biofilm and outer membrane vesicles: a novel therapy overview[J]. FEMS Microbiol Lett, 2015, 362(15): fnv117.
47.	Juodeikis R, Carding SR. Outer membrane vesicles: biogenesis, functions, and issues[J]. Microbiol Mol Biol Rev, 2022, 86(4): e0003222.
48.	Zhao Z, Wang L, Miao J, et al. Regulation of the formation and structure of biofilms by quorum sensing signal molecules packaged in outer membrane vesicles[J]. Sci Total Environ, 2022, 806(4): 151403.
49.	Chen Z, Liu Y, Jiang L, et al. Bacterial outer membrane vesicles increase polymyxin resistance in Pseudomonas aeruginosa while inhibiting its quorum sensing[J]. J Hazard Mater, 2024, 478: 135588.
50.	Seike S, Kobayashi H, Ueda M, et al. Outer membrane vesicles released from aeromonas strains are involved in the biofilm formation[J]. Front Microbiol, 2021, 11: 613650.
51.	Grenier D. Porphyromonas gingivalis outer membrane vesicles mediate coaggregation and piggybacking of treponema denticola and Lachnoanaerobaculum saburreum[J]. Int J Dent, 2013, 2013: 305476.
52.	Schaar V, Nordstr?m T, M?rgelin M, et al. Moraxella catarrhalis outer membrane vesicles carry β-lactamase and promote survival of Streptococcus pneumoniae and Haemophilus influenzae by inactivating amoxicillin[J]. Antimicrob Agents Chemother, 2011, 55(8): 3845-3853.
53.	Schaar V, Uddb?ck I, Nordstr?m T, et al. Group A streptococci are protected from amoxicillin-mediated killing by vesicles containing β-lactamase derived from Haemophilus influenzae[J]. J Antimicrob Chemother, 2014, 69(1): 117-120.
54.	Capodimonte L, Meireles FTP, Bahr G, et al. OXA β-lactamases from Acinetobacter spp. are membrane bound and secreted into outer membrane vesicles[J]. mBio, 2025, 16(2): e0334324.
55.	Li L, Zhang Y, Weng L, et al. NDM-5-carried outer membrane vesicles impair the efficacy of antibiotics against bacterial infections[J]. Antimicrob Agents Chemother, 2025, 69(5): e0180524.
56.	Zhou D, Yang X, Gao Y, et al. The mechanism of outer membrane vesicle-mediated resistance to carbapenem antibiotics[J]. Microb Pathog, 2025, 205: 107654.
57.	He X, Yuan F, Lu F, et al. Vancomycin-induced biofilm formation by methicillin-resistant Staphylococcus aureus is associated with the secretion of membrane vesicles[J]. Microb Pathog, 2017, 110: 225-231.
58.	Kumaraswamy M, Wiull K, Joshi B, et al. Bacterial membrane-derived vesicles attenuate vancomycin activity against methicillin-resistant Staphylococcus aureus[J]. Microorganisms, 2021, 9(10): 2055.
59.	Kaparakis-Liaskos M, Ferrero RL. Immune modulation by bacterial outer membrane vesicles[J]. Nat Rev Immunol, 2015, 15(6): 375-387.
60.	Waller T, Kesper L, Hirschfeld J, et al. Porphyromonas gingivalis outer membrane vesicles induce selective tumor necrosis factor tolerance in a toll-like receptor 4- and mTOR-dependent manner[J]. Infect Immun, 2016, 84(4): 1194-1204.
61.	Tan Y, Zanoni I, Cullen TW, et al. Mechanisms of toll-like receptor 4 endocytosis reveal a common immune-evasion strategy used by pathogenic and commensal bacteria[J]. Immunity, 2015, 43(5): 909-922.
62.	Winter J, Letley D, Rhead J, et al. Helicobacter pylori membrane vesicles stimulate innate pro- and anti-inflammatory responses and induce apoptosis in Jurkat T cells[J]. Infect Immun, 2014, 82(4): 1372-1381.
63.	Hock BD, McKenzie JL, Keenan JI. Helicobacter pylori outer membrane vesicles inhibit human T cell responses via induction of monocyte COX-2 expression[J]. Pathog Dis, 2017, 75(4): 10.
64.	Medzhitov R. The spectrum of inflammatory responses[J]. Science, 2021, 374(6571): 1070-1075.
65.	Cecil JD, O’Brien-Simpson NM, Lenzo JC, et al. Outer membrane vesicles prime and activate macrophage inflammasomes and cytokine secretion in vitro and in vivo[J]. Front Immunol, 2017, 8: 1017.
66.	Turner KL, Cahill BK, Dilello SK, et al. Porin loss impacts the host inflammatory response to outer membrane vesicles of Klebsiella pneumoniae[J]. Antimicrob Agents Chemother, 2015, 60(3): 1360-1369.
67.	Ye C, Li W, Yang Y, et al. Inappropriate use of antibiotics exacerbates inflammation through OMV-induced pyroptosis in MDR Klebsiella pneumoniae infection[J]. Cell Rep, 2021, 36(12): 109750.

1. Aggarwal R, Mahajan P, Pandiya S, et al. Antibiotic resistance: a global crisis, problems and solutions[J]. Crit Rev Microbiol, 2024, 50(5): 896-921.
2. Cecil JD, Sirisaengtaksin N, O’Brien-Simpson NM, et al. Outer membrane vesicle-host cell interactions[J]. Microbiol Spectr, 2019, 7(1): 10.
3. Sartorio MG, Pardue EJ, Feldman MF, et al. Bacterial outer membrane vesicles: from discovery to applications[J]. Annu Rev Microbiol, 2021, 75: 609-630.
4. Schwechheimer C, Kuehn MJ. Outer-membrane vesicles from Gram-negative bacteria: biogenesis and functions[J]. Nat Rev Microbiol, 2015, 13(10): 605-619.
5. Kim JY, Suh JW, Kang JS, et al. Gram-negative bacteria’s outer membrane vesicles[J]. Infect Chemother, 2023, 55(1): 1-9.
6. Schwechheimer C, Kulp A, Kuehn MJ. Modulation of bacterial outer membrane vesicle production by envelope structure and content[J]. BMC Microbiol, 2014, 14: 324.
7. Cassin EK, Tseng BS. Pushing beyond the envelope: the potential roles of OprF in Pseudomonas aeruginosa biofilm formation and pathogenicity[J]. J Bacteriol, 2019, 201(18): e00050-19.
8. Schwechheimer C, Rodriguez DL, Kuehn MJ. NlpI-mediated modulation of outer membrane vesicle production through peptidoglycan dynamics in Escherichia coli[J]. Microbiologyopen, 2015, 4(3): 375-389.
9. Lappann M, Otto A, Becher D, et al. Comparative proteome analysis of spontaneous outer membrane vesicles and purified outer membranes of Neisseria meningitidis[J]. J Bacteriol, 2013, 195(19): 4425-4435.
10. McBroom AJ, Kuehn MJ. Release of outer membrane vesicles by Gram-negative bacteria is a novel envelope stress response[J]. Mol Microbiol, 2007, 63(2): 545-558.
11. Macdonald IA, Kuehn MJ. Stress-induced outer membrane vesicle production by Pseudomonas aeruginosa[J]. J Bacteriol, 2013, 195(13): 2971-2981.
12. Roden JA, Wells DH, Chomel BB, et al. Hemin binding protein C is found in outer membrane vesicles and protects Bartonella henselae against toxic concentrations of hemin[J]. Infect Immun, 2012, 80(3): 929-942.
13. McMahon KJ, Castelli ME, García Vescovi E, et al. Biogenesis of outer membrane vesicles in Serratia marcescens is thermoregulated and can be induced by activation of the Rcs phosphorelay system[J]. J Bacteriol, 2012, 194(12): 3241-3249.
14. Frias A, Manresa A, de Oliveira E, et al. Membrane vesicles: a common feature in the extracellular matter of cold-adapted antarctic bacteria[J]. Microb Ecol, 2010, 59(3): 476-486.
15. Chowdhury C, Jagannadham MV. Virulence factors are released in association with outer membrane vesicles of Pseudomonas syringae pv. tomato T1 during normal growth[J]. Biochim Biophys Acta, 2013, 1834(1): 231-239.
16. Kulkarni HM, Swamy ChV, Jagannadham MV. Molecular characterization and functional analysis of outer membrane vesicles from the antarctic bacterium Pseudomonas syringae suggest a possible response to environmental conditions[J]. J Proteome Res, 2014, 13(3): 1345-1358.
17. Tashiro Y, Inagaki A, Shimizu M, et al. Characterization of phospholipids in membrane vesicles derived from Pseudomonas aeruginosa[J]. Biosci Biotechnol Biochem, 2011, 75(3): 605-607.
18. Murphy K, Park AJ, Hao Y, et al. Influence of O polysaccharides on biofilm development and outer membrane vesicle biogenesis in Pseudomonas aeruginosa PAO1[J]. J Bacteriol, 2014, 196(7): 1306-1317.
19. Yamaguchi M, Sato K, Yukitake H, et al. A Porphyromonas gingivalis mutant defective in a putative glycosyltransferase exhibits defective biosynthesis of the polysaccharide portions of lipopolysaccharide, decreased gingipain activities, strong autoaggregation, and increased biofilm formation[J]. Infect Immun, 2010, 78(9): 3801-3812.
20. Huang C, Cao W, Zhou S, et al. Biogenesis mechanisms, regulatory strategies, and applications of bacterial extracellular vesicles[J]. Crit Rev Biotechnol, 2025, 45(8): 1700-1716.
21. Zhao X, Wei Y, Bu Y, et al. Review on bacterial outer membrane vesicles: structure, vesicle formation, separation and biotechnological applications[J]. Microb Cell Fact, 2025, 24(1): 27.
22. Dai K, Liao B, Huang X, et al. Consistency in bacterial extracellular vesicle production: key to their application in human health[J]. Extracell Vesicles Circ Nucl Acids, 2025, 6(1): 1-20.
23. Gon?alves MO, Pincela Lins PM, Kassab G, et al. Outer membrane vesicles (OMVs) and their therapeutic potential as anti-infectious agents[J]. Nano Trends, 2025, 11: 100129.
24. Meena H, Nandy S, Patil VS, et al. Lipidomics in understanding microbial biofilms[C]//Omics approaches in biofilm research: perspectives and applications. Cham: Springer Nature Switzerland, 2025: 287-310.
25. Li L, Xu X, Cheng P, et al. Klebsiella pneumoniae derived outer membrane vesicles mediated bacterial virulence, antibiotic resistance, host immune responses and clinical applications[J]. Virulence, 2025, 16(1): 2449722.
26. Li P, Luo W, Xiang TX, et al. Horizontal gene transfer via OMVs co-carrying virulence and antimicrobial-resistant genes is a novel way for the dissemination of carbapenem-resistant hypervirulent Klebsiella pneumoniae[J]. Front Microbiol, 2022, 13: 945972.
27. Dhurve G, Madikonda AK, Jagannadham MV, et al. Outer membrane vesicles of Acinetobacter baumannii DS002 are selectively enriched with TonB-dependent transporters and play a key role in iron acquisition[J]. Microbiol Spectr, 2022, 10(2): e0029322.
28. Wachino JI. Horizontal gene transfer systems for spread of antibiotic resistance in Gram-negative bacteria[J]. Microbiol Immunol, 2025, 69(7): 367-376.
29. Chen LJ, Jing XP, Meng DL, et al. Newly detected transmission of blaKPC-2 by outer membrane vesicles in Klebsiella pneumoniae[J]. Curr Med Sci, 2023, 43(1): 80-85.
30. Dell’Annunziata F, Dell’Aversana C, Doti N, et al. Outer membrane vesicles derived from Klebsiella pneumoniae are a driving force for horizontal gene transfer[J]. Int J Mol Sci, 2021, 22(16): 8732.
31. Wang M, Yao Y, Yang Y, et al. The characterization of outer membrane vesicles (OMVs) and their role in mediating antibiotic-resistance gene transfer through natural transformation in Riemerella anatipestifer[J]. Poult Sci, 2025, 104(2): 104730.
32. He M, Yin S, Huang X, et al. Insights into the regulatory role of bacterial sncRNA and its extracellular delivery via OMVs[J]. Appl Microbiol Biotechnol, 2024, 108(1): 29.
33. Parker A, Gottesman S. Small RNA regulation of TolC, the outer membrane component of bacterial multidrug transporters[J]. J Bacteriol, 2016, 198(7): 1101-1113.
34. Gutierrez A, Laureti L, Crussard S, et al. β-Lactam antibiotics promote bacterial mutagenesis via an RpoS-mediated reduction in replication fidelity[J]. Nat Commun, 2013, 4: 1610.
35. Nishino K, Yamasaki S, Hayashi-Nishino M, et al. Effect of overexpression of small non-coding DsrA RNA on multidrug efflux in Escherichia coli[J]. J Antimicrob Chemother, 2011, 66(2): 291-296.
36. Sarshar M, Scribano D, Palamara AT, et al. The Acinetobacter baumannii model can explain the role of small non-coding RNAs as potential mediators of host-pathogen interactions[J]. Front Mol Biosci, 2022, 9: 1088783.
37. Eyraud A, Tattevin P, Chabelskaya S, et al. A small RNA controls a protein regulator involved in antibiotic resistance in Staphylococcus aureus[J]. Nucleic Acids Res, 2014, 42(8): 4892-4905.
38. Lucena ACR, Ferrarini MG, de Oliveira WK, et al. Modulation of Klebsiella pneumoniae outer membrane vesicle protein cargo under antibiotic treatment[J]. Biomedicines, 2023, 11(6): 1515.
39. Park AJ, Surette MD, Khursigara CM. Antimicrobial targets localize to the extracellular vesicle-associated proteome of Pseudomonas aeruginosa grown in a biofilm[J]. Front Microbiol, 2014, 5: 464.
40. Farrugia C, Stafford GP, Murdoch C. Porphyromonas gingivalis outer membrane vesicles increase vascular permeability[J]. J Dent Res, 2020, 99(13): 1494-1501.
41. Wang J, Terrasse R, Bafna JA, et al. Electrophysiological characterization of transport across outer-membrane channels from Gram-negative bacteria in presence of lipopolysaccharides[J]. Angew Chem Int Ed Engl, 2020, 59(22): 8517-8521.
42. Jasim R, Han ML, Zhu Y, et al. Lipidomic analysis of the outer membrane vesicles from paired polymyxin-susceptible and -resistant Klebsiella pneumoniae clinical isolates[J]. Int J Mol Sci, 2018, 19(8): 2356.
43. Sáenz JP, Grosser D, Bradley AS, et al. Hopanoids as functional analogues of cholesterol in bacterial membranes[J]. Proc Natl Acad Sci USA, 2015, 112(38): 11971-11976.
44. Kim SW, Park SB, Im SP, et al. Outer membrane vesicles from β-lactam-resistant Escherichia coli enable the survival of β-lactam-susceptible E. coli in the presence of β-lactam antibiotics[J]. Sci Rep, 2018, 8(1): 5402.
45. Miao B, Wang D, Yu L, et al. Mechanism and nanotechnological-based therapeutics for tolerance and resistance of bacterial biofilms[J]. Microbiol Res, 2025, 292: 127987.
46. Wang W, Chanda W, Zhong M. The relationship between biofilm and outer membrane vesicles: a novel therapy overview[J]. FEMS Microbiol Lett, 2015, 362(15): fnv117.
47. Juodeikis R, Carding SR. Outer membrane vesicles: biogenesis, functions, and issues[J]. Microbiol Mol Biol Rev, 2022, 86(4): e0003222.
48. Zhao Z, Wang L, Miao J, et al. Regulation of the formation and structure of biofilms by quorum sensing signal molecules packaged in outer membrane vesicles[J]. Sci Total Environ, 2022, 806(4): 151403.
49. Chen Z, Liu Y, Jiang L, et al. Bacterial outer membrane vesicles increase polymyxin resistance in Pseudomonas aeruginosa while inhibiting its quorum sensing[J]. J Hazard Mater, 2024, 478: 135588.
50. Seike S, Kobayashi H, Ueda M, et al. Outer membrane vesicles released from aeromonas strains are involved in the biofilm formation[J]. Front Microbiol, 2021, 11: 613650.
51. Grenier D. Porphyromonas gingivalis outer membrane vesicles mediate coaggregation and piggybacking of treponema denticola and Lachnoanaerobaculum saburreum[J]. Int J Dent, 2013, 2013: 305476.
52. Schaar V, Nordstr?m T, M?rgelin M, et al. Moraxella catarrhalis outer membrane vesicles carry β-lactamase and promote survival of Streptococcus pneumoniae and Haemophilus influenzae by inactivating amoxicillin[J]. Antimicrob Agents Chemother, 2011, 55(8): 3845-3853.
53. Schaar V, Uddb?ck I, Nordstr?m T, et al. Group A streptococci are protected from amoxicillin-mediated killing by vesicles containing β-lactamase derived from Haemophilus influenzae[J]. J Antimicrob Chemother, 2014, 69(1): 117-120.
54. Capodimonte L, Meireles FTP, Bahr G, et al. OXA β-lactamases from Acinetobacter spp. are membrane bound and secreted into outer membrane vesicles[J]. mBio, 2025, 16(2): e0334324.
55. Li L, Zhang Y, Weng L, et al. NDM-5-carried outer membrane vesicles impair the efficacy of antibiotics against bacterial infections[J]. Antimicrob Agents Chemother, 2025, 69(5): e0180524.
56. Zhou D, Yang X, Gao Y, et al. The mechanism of outer membrane vesicle-mediated resistance to carbapenem antibiotics[J]. Microb Pathog, 2025, 205: 107654.
57. He X, Yuan F, Lu F, et al. Vancomycin-induced biofilm formation by methicillin-resistant Staphylococcus aureus is associated with the secretion of membrane vesicles[J]. Microb Pathog, 2017, 110: 225-231.
58. Kumaraswamy M, Wiull K, Joshi B, et al. Bacterial membrane-derived vesicles attenuate vancomycin activity against methicillin-resistant Staphylococcus aureus[J]. Microorganisms, 2021, 9(10): 2055.
59. Kaparakis-Liaskos M, Ferrero RL. Immune modulation by bacterial outer membrane vesicles[J]. Nat Rev Immunol, 2015, 15(6): 375-387.
60. Waller T, Kesper L, Hirschfeld J, et al. Porphyromonas gingivalis outer membrane vesicles induce selective tumor necrosis factor tolerance in a toll-like receptor 4- and mTOR-dependent manner[J]. Infect Immun, 2016, 84(4): 1194-1204.
61. Tan Y, Zanoni I, Cullen TW, et al. Mechanisms of toll-like receptor 4 endocytosis reveal a common immune-evasion strategy used by pathogenic and commensal bacteria[J]. Immunity, 2015, 43(5): 909-922.
62. Winter J, Letley D, Rhead J, et al. Helicobacter pylori membrane vesicles stimulate innate pro- and anti-inflammatory responses and induce apoptosis in Jurkat T cells[J]. Infect Immun, 2014, 82(4): 1372-1381.
63. Hock BD, McKenzie JL, Keenan JI. Helicobacter pylori outer membrane vesicles inhibit human T cell responses via induction of monocyte COX-2 expression[J]. Pathog Dis, 2017, 75(4): 10.
64. Medzhitov R. The spectrum of inflammatory responses[J]. Science, 2021, 374(6571): 1070-1075.
65. Cecil JD, O’Brien-Simpson NM, Lenzo JC, et al. Outer membrane vesicles prime and activate macrophage inflammasomes and cytokine secretion in vitro and in vivo[J]. Front Immunol, 2017, 8: 1017.
66. Turner KL, Cahill BK, Dilello SK, et al. Porin loss impacts the host inflammatory response to outer membrane vesicles of Klebsiella pneumoniae[J]. Antimicrob Agents Chemother, 2015, 60(3): 1360-1369.
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Research advances in the mechanisms of infection and antibiotic resistance mediated by outer membrane vesicle

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