VACUNAS CON VESÍCULAS DE MEMBRANA EXTERNA: ¿EL FUTURO DE LAS VACUNAS?
DOI:
https://doi.org/10.32399/icuap.rdic.2448-5829.2023.27.1193Palabras clave:
Vacunas; Bacterias multirresistentes; Secreción de microvesículas; Respuesta inmune; Membrana extracelularResumen
Prevenir la propagación de una enfermedad, sea por patógenos virales o bacterianos, implica
brindar protección a miles de millones de personas y los factores subsecuentes durante
el proceso. La vacunación, es sin duda, la manera más efectiva de lograr esto. Muchos países
priorizan la realización de campañas de vacunación procurando el bienestar en salud pública
de sus habitantes. El desarrollo de vacunas se ha situado en 6 tipos principales, cuya respuesta
es efectiva para muchas enfermedades, sin embargo, algunas demostraron tener limitantes
para determinadas condiciones clínicas. Por ello, la identificación de nuevos candidatos para
desarrollar vacunas ha captado la atención de muchos grupos de investigación. De acuerdo con
la respuesta inmune humoral y celular inducida por una vacuna, es que se mide su efectividad
contra una enfermedad. Las Microvesículas (MVs) son vesículas de tamaño nanométrico que
se secretan al espacio extracelular, cuyo contenido se conforma de carbohidratos, proteínas,
lípidos, ácidos nucleicos (ADN y ARN) y otros metabolitos. Dicha conformación puede utilizarse
como un vehículo que transporte para antígenos y moléculas inmunogénicas directamente a las
células diana. Ya se ha desarrollado una vacuna para la bacteria multirresistente Neisseria gonorrhoeae
utilizando vesículas de membrana externa (OMVs), cuya respuesta inmune fue prometedora.
En esta revisión, se estudiará la participación de las microvesículas secretadas por
bacterias Gram-positivas y Gram-negativas de distintos aislados, procesos de patogenicidad en
los que participan, utilidad en el transporte de moléculas inmunogénicas y su efectividad como
componentes vacunales contra bacterias multirresistentes.
Citas
Azze, R. F. O. (2019). A meningococcal B vaccine induces cross-protection against gonorrhea.
Clinical and Experimental Vaccine Research, 8(2), 110–115. https://doi.org/10.7774/
cevr.2019.8.2.110
Bomberger, J. M., Maceachran, D. P., Coutermarsh, B. A., Ye, S., O’Toole, G. A., & Stanton, B.
A. (2009). Long-distance delivery of bacterial virulence factors by Pseudomonas aeruginosa
outer membrane vesicles. PLoS pathogens, 5(4), e1000382. https://doi.org/10.1371/journal.
ppat.1000382
Chen, H., Zhou, M., Zeng, Y., Lv, Z., Wang, P., & Han, L. (2022, June 16). Recent advances in biomedical
applications of bacterial outer membrane vesicles. Journal of Materials Chemistry B. Royal
Society of Chemistry. https://doi.org/10.1039/d2tb00683a
Durham A. E. (2018). An evaluation of serum gentamicin concentrations and bacterial susceptibility
to gentamicin in equine practice. Journal of veterinary internal medicine, 32(3), 1194–1201.
https://doi.org/10.1111/jvim.15078
Elsevier. (s. f.). Células eucariotas y procariotas: ¿Sabrías distinguirlas? Te damos las claves.
Elsevier Connect. https://www.elsevier.com/es-es/connect/medicina/celulas-eucariotas-y-procariotas-
sabrias-distinguirlas-te-damos-las-claves#:~:text=Las%20c%C3%A9lulas%20de%20
los%20animales,griego%20%C2%ABn%C3%BAcleo%20primitivo%C2%BB
European Medicines Agency. (2018). Bexsero (vacuna antimeningocócica del grupo B [ADNr,
componente, adsorbida]). European Medicines Agency. https://www.ema.europa.eu/en/documents/
overview/bexsero-epar-summary-public_es.pdf
Furuyama, N., & Sircili, M. P. (2021). Outer membrane vesicles (OMVs) produced by gram-negative
bacteria: Structure, functions, biogenesis, and vaccine application. BioMed Research International.
Hindawi Limited. https://doi.org/10.1155/2021/1490732
Gao, J., Wang, S., & Wang, Z. (2017). High yield, scalable and remotely drug-loaded neutrophil-
derived extracellular vesicles (EVs) for anti-inflammation therapy. Biomaterials, 135, 62–73.
https://doi.org/10.1016/j.biomaterials.2017.05.003
Gao, W., Fang, R. H., Thamphiwatana, S., Luk, B. T., Li, J., Angsantikul, P., … Zhang, L. (2015). Modulating
antibacterial immunity via bacterial membrane-coated nanoparticles. Nano Letters, 15(2),
1403–1409. https://doi.org/10.1021/nl504798g
González-Melado, F. J., & Di Pietro, M. L. (2020). The vaccine against COVID-19 and institutional
trust. La vacuna frente a la COVID-19 y la confianza institucional. Enfermedades infecciosas
y microbiologia clinica (English ed.), 39(10), 510–515. Advance online publication. https://doi.
org/10.1016/j.eimc.2020.08.001
González-Vázquez, María Cristina, Guerra-Martínez, Álvaro Asaf, Escobedo-Herrera, Betsabé,
& Carabarin Lima, Alejandro. (2022). Microvesículas bacterianas secretadas y su potencial uso
en el desarrollo de vacunas. Alianzas y tendencias BUAP, 7(28), 47–74. https://doi.org/10.5281/
zenodo.7425479
Huang, W., Zhang, Q., Li, W., Chen, Y., Shu, C., Li, Q., … Ma, Y. (2019). Anti-outer membrane vesicle
antibodies increase antibiotic sensitivity of pan-drug-resistant Acinetobacter baumannii.
Frontiers in Microbiology, 10(JUN). https://doi.org/10.3389/fmicb.2019.01379
135
Año 9, No. 27, 2023, pp. 125-137
Jahromi, L. P., & Fuhrmann, G. (2021, June 1). Bacterial extracellular vesicles: Understanding biology
promotes applications as nanopharmaceuticals. Advanced Drug Delivery Reviews. Elsevier
B.V. https://doi.org/10.1016/j.addr.2021.03.012
Jan, A. T. (2017). Outer Membrane Vesicles (OMVs) of gram-negative bacteria: A perspective
update. Frontiers in Microbiology, 8(JUN). https://doi.org/10.3389/fmicb.2017.01053
Kadurugamuwa, J. L., & Beveridge, T. J. (1995). Virulence factors are released from Pseudomonas
aeruginosa in association with membrane vesicles during normal growth and exposure to gentamicin:
A novel mechanism of enzyme secretion. Journal of Bacteriology, 177(14), 3998–4008.
https://doi.org/10.1128/jb.177.14.3998-4008.1995
Kim, J. H., Lee, J., Park, J., & Gho, Y. S. (2015, April 1). Gram-negative and Gram-positive bacterial
extracellular vesicles. Seminars in Cell and Developmental Biology. Academic Press. https://doi.
org/10.1016/j.semcdb.2015.02.006
Kim, S. Y., Kim, S. I., Yun, S. H., Shin, M., Lee, Y. C., & Lee, J. C. (2020). Proteins in outer membrane
vesicles produced by burkholderia cepacia are responsible for pro-inflammatory responses in
epithelial cells. Journal of Bacteriology and Virology, 50(4), 227–234. https://doi.org/10.4167/
jbv.2020.50.4.227
Klimentová, J., & Stulík, J. (2015). Methods of isolation and purification of outer membrane vesicles
from gram-negative bacteria. Microbiological Research, 170, 1-9. https://doi.org/10.1016/j.
micres.2014.09.006
Krammer, F. (2020, October 22). SARS-CoV-2 vaccines in development. Nature. Nature Research.
https://doi.org/10.1038/s41586-020-2798-3
Leduc, I., Connolly, K. L., Begum, A., Underwood, K., Darnell, S., Shafer, W. M., Balthazar, J. T., Macintyre,
A. N., Sempowski, G. D., Duncan, J. A., Little, M. B., Rahman, N., Garges, E. C., & Jerse, A.
E. (2020). The serogroup B meningococcal outer membrane vesicle-based vaccine 4CMenB induces
cross-species protection against Neisseria gonorrhoeae. PLoS pathogens, 16(12), e1008602.
https://doi.org/10.1371/journal.ppat.1008602
Longtin, J., Dion, R., Simard, M., Betala Belinga, J. F., Longtin, Y., Lefebvre, B., Labbé, A. C., Deceuninck,
G., & De Wals, P. (2017). Possible Impact of Wide-scale Vaccination Against Serogroup B
Neisseria Meningitidis on Gonorrhea Incidence Rates in One Region of Quebec, Canada. Open
Forum Infectious Diseases, 4(Suppl 1), S734–S735. https://doi.org/10.1093/ofid/ofx180.002
Marchetti, M., Errecalde, J., & Mestorino, N. (2011). Dirección para correspondencia: Resistencia
bacteriana a Los antimicrobianos ocasionada por bombas de Eflujo. Impacto en la Multirresistencia.
AnAlectA Vet, 40(312), 40–53.
Matthias, K. A., Connolly, K. L., Begum, A. A., Jerse, A. E., MacIntyre, A. N., Sempowski, G. D., &
Bash, M. C. (2022). Meningococcal Detoxified Outer Membrane Vesicle Vaccines Enhance Gonococcal
Clearance in a Murine Infection Model. In Journal of Infectious Diseases (Vol. 225, pp.
650–660). Oxford University Press. https://doi.org/10.1093/infdis/jiab450
Meldolesi J. (2018). Exosomes and Ectosomes in Intercellular Communication. Current biology:
CB, 28(8), R435–R444. https://doi.org/10.1016/j.cub.2018.01.059
Milani, G., Lana, T., Bresolin, S., Aveic, S., Pastò, A., Frasson, C., & Te Kronnie, G. (2017). Expression
profiling of circulating microvesicles reveals intercellular transmission of oncogenic
pathways. Molecular Cancer Research, 15(6), 683–695. https://doi.org/10.1158/1541-7786.MCR-
16-0307
136
Año 9, No. 27, 2023, pp. 125-137
Nikaido H. (2009). Multidrug resistance in bacteria. Annual review of biochemistry, 78, 119–146.
https://doi.org/10.1146/annurev.biochem.78.082907.145923
Paniagua R., Nistal M., Sesma P., Álvarez-Uría M., Fraile B., Anadón R. & Sáez F. J. (2007). Biología
Celular 3ra edición. McGRAW-HILL - Interamericana de España, S. A. U. http://librodigital.sangregorio.
edu.ec/librosusgp/18073.pdf
Pérez-Cruz, C., Delgado, L., López-Iglesias, C., & Mercade, E. (2015). Outer-inner membrane vesicles
naturally secreted by gram-negative pathogenic bacteria. PLoS ONE, 10(1). https://doi.
org/10.1371/journal.pone.0116896
Petousis-Harris, H., Paynter, J., Morgan, J., Saxton, P., McArdle, B., Goodyear-Smith, F., & Black,
S. (2017). Effectiveness of a group B outer membrane vesicle meningococcal vaccine against
gonorrhoea in New Zealand: a retrospective case-control study. Lancet (London, England),
390(10102), 1603–1610. https://doi.org/10.1016/S0140-6736(17)31449-6
Pfizer, (s. f.). Understanding Six Types of Vaccine Technologies. https://www.pfizer.com/news/
articles/understanding_six_types_of_vaccine_technologies
Preado J., V. (2001). Conceptos microbiológicos de Streptococcus pneumoniae: Basic microbiological
aspects. Revista Chilena de Infectología, 18. https://doi.org/10.4067/s0716-
10182001000000002
Rodríguez, H. (2022, 3 abril). En 2050 la resistencia a los antibióticos será responsable de 10 millones
de muertes anuales. www.nationalgeographic.com.es. https://www.nationalgeographic.
com.es/ciencia/2050-resistencia-a-antibioticos-sera-responsable-10-millones-muertes-anuales_
18090
Semchenko, E. A., & Seib, K. L. (2022, January 1). Outer membrane vesicle vaccines for Neisseria
gonorrhoeae. Nature Reviews Urology. Nature Research. https://doi.org/10.1038/s41585-021-
00534-5
Shkair, L., Garanina, E. E., Stott, R. J., Foster, T. L., Rizvanov, A. A., & Khaiboullina, S. F. (2021,
February 1). Membrane microvesicles as potential vaccine candidates. International Journal of
Molecular Sciences. MDPI AG. https://doi.org/10.3390/ijms22031142
St. Cyr, S., Barbee, L., Workowski, K. A., Bachmann, L. H., Pham, C., Schlanger, K., … Thorpe, P.
(2020). Update to CDC’s Treatment Guidelines for Gonococcal Infection, 2020. MMWR. Morbidity
and Mortality Weekly Report, 69(50), 1911–1916. https://doi.org/10.15585/mmwr.mm6950a6
Takada, K., Ohno, N., & Yadomae, T. (1994). Binding of lysozyme to lipopolysaccharide suppresses
tumor necrosis factor production in vivo. Infection and immunity, 62(4), 1171–1175. https://
doi.org/10.1128/iai.62.4.1171-1175.1994
Toyofuku, M., Nomura, N., & Eberl, L. (2019, January 1). Types and origins of bacterial membrane
vesicles. Nature Reviews Microbiology. Nature Publishing Group. https://doi.org/10.1038/
s41579-018-0112-2
United Nations. (s. f.). Las muertes por COVID-19 sumarían 15 millones entre 2020 y 2021 | Naciones
Unidas. https://www.un.org/es/desa/las-muertes-por-covid-19-sumar%C3%ADan-15-millones-
entre-2020-y-2021
Wei, S., Jiao, D., & Xing, W. (2022). A rapid method for isolation of bacterial extracellular vesicles
from culture media using epsilon-poly-L–lysine that enables immunological function research.
Frontiers in Immunology, 13. https://doi.org/10.3389/fimmu.2022.930510
Wei, S., Li, X., Wang, J., Wang, Y., Zhang, C., Dai, S., … Shan, B. (2022). Outer Membrane Vesicles
Secreted by Helicobacter pylori Transmitting Gastric Pathogenic Virulence Factors. ACS Omega,
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Media S.A. https://doi.org/10.3389/fimmu.2022.840550
Clinical and Experimental Vaccine Research, 8(2), 110–115. https://doi.org/10.7774/
cevr.2019.8.2.110
Bomberger, J. M., Maceachran, D. P., Coutermarsh, B. A., Ye, S., O’Toole, G. A., & Stanton, B.
A. (2009). Long-distance delivery of bacterial virulence factors by Pseudomonas aeruginosa
outer membrane vesicles. PLoS pathogens, 5(4), e1000382. https://doi.org/10.1371/journal.
ppat.1000382
Chen, H., Zhou, M., Zeng, Y., Lv, Z., Wang, P., & Han, L. (2022, June 16). Recent advances in biomedical
applications of bacterial outer membrane vesicles. Journal of Materials Chemistry B. Royal
Society of Chemistry. https://doi.org/10.1039/d2tb00683a
Durham A. E. (2018). An evaluation of serum gentamicin concentrations and bacterial susceptibility
to gentamicin in equine practice. Journal of veterinary internal medicine, 32(3), 1194–1201.
https://doi.org/10.1111/jvim.15078
Elsevier. (s. f.). Células eucariotas y procariotas: ¿Sabrías distinguirlas? Te damos las claves.
Elsevier Connect. https://www.elsevier.com/es-es/connect/medicina/celulas-eucariotas-y-procariotas-
sabrias-distinguirlas-te-damos-las-claves#:~:text=Las%20c%C3%A9lulas%20de%20
los%20animales,griego%20%C2%ABn%C3%BAcleo%20primitivo%C2%BB
European Medicines Agency. (2018). Bexsero (vacuna antimeningocócica del grupo B [ADNr,
componente, adsorbida]). European Medicines Agency. https://www.ema.europa.eu/en/documents/
overview/bexsero-epar-summary-public_es.pdf
Furuyama, N., & Sircili, M. P. (2021). Outer membrane vesicles (OMVs) produced by gram-negative
bacteria: Structure, functions, biogenesis, and vaccine application. BioMed Research International.
Hindawi Limited. https://doi.org/10.1155/2021/1490732
Gao, J., Wang, S., & Wang, Z. (2017). High yield, scalable and remotely drug-loaded neutrophil-
derived extracellular vesicles (EVs) for anti-inflammation therapy. Biomaterials, 135, 62–73.
https://doi.org/10.1016/j.biomaterials.2017.05.003
Gao, W., Fang, R. H., Thamphiwatana, S., Luk, B. T., Li, J., Angsantikul, P., … Zhang, L. (2015). Modulating
antibacterial immunity via bacterial membrane-coated nanoparticles. Nano Letters, 15(2),
1403–1409. https://doi.org/10.1021/nl504798g
González-Melado, F. J., & Di Pietro, M. L. (2020). The vaccine against COVID-19 and institutional
trust. La vacuna frente a la COVID-19 y la confianza institucional. Enfermedades infecciosas
y microbiologia clinica (English ed.), 39(10), 510–515. Advance online publication. https://doi.
org/10.1016/j.eimc.2020.08.001
González-Vázquez, María Cristina, Guerra-Martínez, Álvaro Asaf, Escobedo-Herrera, Betsabé,
& Carabarin Lima, Alejandro. (2022). Microvesículas bacterianas secretadas y su potencial uso
en el desarrollo de vacunas. Alianzas y tendencias BUAP, 7(28), 47–74. https://doi.org/10.5281/
zenodo.7425479
Huang, W., Zhang, Q., Li, W., Chen, Y., Shu, C., Li, Q., … Ma, Y. (2019). Anti-outer membrane vesicle
antibodies increase antibiotic sensitivity of pan-drug-resistant Acinetobacter baumannii.
Frontiers in Microbiology, 10(JUN). https://doi.org/10.3389/fmicb.2019.01379
135
Año 9, No. 27, 2023, pp. 125-137
Jahromi, L. P., & Fuhrmann, G. (2021, June 1). Bacterial extracellular vesicles: Understanding biology
promotes applications as nanopharmaceuticals. Advanced Drug Delivery Reviews. Elsevier
B.V. https://doi.org/10.1016/j.addr.2021.03.012
Jan, A. T. (2017). Outer Membrane Vesicles (OMVs) of gram-negative bacteria: A perspective
update. Frontiers in Microbiology, 8(JUN). https://doi.org/10.3389/fmicb.2017.01053
Kadurugamuwa, J. L., & Beveridge, T. J. (1995). Virulence factors are released from Pseudomonas
aeruginosa in association with membrane vesicles during normal growth and exposure to gentamicin:
A novel mechanism of enzyme secretion. Journal of Bacteriology, 177(14), 3998–4008.
https://doi.org/10.1128/jb.177.14.3998-4008.1995
Kim, J. H., Lee, J., Park, J., & Gho, Y. S. (2015, April 1). Gram-negative and Gram-positive bacterial
extracellular vesicles. Seminars in Cell and Developmental Biology. Academic Press. https://doi.
org/10.1016/j.semcdb.2015.02.006
Kim, S. Y., Kim, S. I., Yun, S. H., Shin, M., Lee, Y. C., & Lee, J. C. (2020). Proteins in outer membrane
vesicles produced by burkholderia cepacia are responsible for pro-inflammatory responses in
epithelial cells. Journal of Bacteriology and Virology, 50(4), 227–234. https://doi.org/10.4167/
jbv.2020.50.4.227
Klimentová, J., & Stulík, J. (2015). Methods of isolation and purification of outer membrane vesicles
from gram-negative bacteria. Microbiological Research, 170, 1-9. https://doi.org/10.1016/j.
micres.2014.09.006
Krammer, F. (2020, October 22). SARS-CoV-2 vaccines in development. Nature. Nature Research.
https://doi.org/10.1038/s41586-020-2798-3
Leduc, I., Connolly, K. L., Begum, A., Underwood, K., Darnell, S., Shafer, W. M., Balthazar, J. T., Macintyre,
A. N., Sempowski, G. D., Duncan, J. A., Little, M. B., Rahman, N., Garges, E. C., & Jerse, A.
E. (2020). The serogroup B meningococcal outer membrane vesicle-based vaccine 4CMenB induces
cross-species protection against Neisseria gonorrhoeae. PLoS pathogens, 16(12), e1008602.
https://doi.org/10.1371/journal.ppat.1008602
Longtin, J., Dion, R., Simard, M., Betala Belinga, J. F., Longtin, Y., Lefebvre, B., Labbé, A. C., Deceuninck,
G., & De Wals, P. (2017). Possible Impact of Wide-scale Vaccination Against Serogroup B
Neisseria Meningitidis on Gonorrhea Incidence Rates in One Region of Quebec, Canada. Open
Forum Infectious Diseases, 4(Suppl 1), S734–S735. https://doi.org/10.1093/ofid/ofx180.002
Marchetti, M., Errecalde, J., & Mestorino, N. (2011). Dirección para correspondencia: Resistencia
bacteriana a Los antimicrobianos ocasionada por bombas de Eflujo. Impacto en la Multirresistencia.
AnAlectA Vet, 40(312), 40–53.
Matthias, K. A., Connolly, K. L., Begum, A. A., Jerse, A. E., MacIntyre, A. N., Sempowski, G. D., &
Bash, M. C. (2022). Meningococcal Detoxified Outer Membrane Vesicle Vaccines Enhance Gonococcal
Clearance in a Murine Infection Model. In Journal of Infectious Diseases (Vol. 225, pp.
650–660). Oxford University Press. https://doi.org/10.1093/infdis/jiab450
Meldolesi J. (2018). Exosomes and Ectosomes in Intercellular Communication. Current biology:
CB, 28(8), R435–R444. https://doi.org/10.1016/j.cub.2018.01.059
Milani, G., Lana, T., Bresolin, S., Aveic, S., Pastò, A., Frasson, C., & Te Kronnie, G. (2017). Expression
profiling of circulating microvesicles reveals intercellular transmission of oncogenic
pathways. Molecular Cancer Research, 15(6), 683–695. https://doi.org/10.1158/1541-7786.MCR-
16-0307
136
Año 9, No. 27, 2023, pp. 125-137
Nikaido H. (2009). Multidrug resistance in bacteria. Annual review of biochemistry, 78, 119–146.
https://doi.org/10.1146/annurev.biochem.78.082907.145923
Paniagua R., Nistal M., Sesma P., Álvarez-Uría M., Fraile B., Anadón R. & Sáez F. J. (2007). Biología
Celular 3ra edición. McGRAW-HILL - Interamericana de España, S. A. U. http://librodigital.sangregorio.
edu.ec/librosusgp/18073.pdf
Pérez-Cruz, C., Delgado, L., López-Iglesias, C., & Mercade, E. (2015). Outer-inner membrane vesicles
naturally secreted by gram-negative pathogenic bacteria. PLoS ONE, 10(1). https://doi.
org/10.1371/journal.pone.0116896
Petousis-Harris, H., Paynter, J., Morgan, J., Saxton, P., McArdle, B., Goodyear-Smith, F., & Black,
S. (2017). Effectiveness of a group B outer membrane vesicle meningococcal vaccine against
gonorrhoea in New Zealand: a retrospective case-control study. Lancet (London, England),
390(10102), 1603–1610. https://doi.org/10.1016/S0140-6736(17)31449-6
Pfizer, (s. f.). Understanding Six Types of Vaccine Technologies. https://www.pfizer.com/news/
articles/understanding_six_types_of_vaccine_technologies
Preado J., V. (2001). Conceptos microbiológicos de Streptococcus pneumoniae: Basic microbiological
aspects. Revista Chilena de Infectología, 18. https://doi.org/10.4067/s0716-
10182001000000002
Rodríguez, H. (2022, 3 abril). En 2050 la resistencia a los antibióticos será responsable de 10 millones
de muertes anuales. www.nationalgeographic.com.es. https://www.nationalgeographic.
com.es/ciencia/2050-resistencia-a-antibioticos-sera-responsable-10-millones-muertes-anuales_
18090
Semchenko, E. A., & Seib, K. L. (2022, January 1). Outer membrane vesicle vaccines for Neisseria
gonorrhoeae. Nature Reviews Urology. Nature Research. https://doi.org/10.1038/s41585-021-
00534-5
Shkair, L., Garanina, E. E., Stott, R. J., Foster, T. L., Rizvanov, A. A., & Khaiboullina, S. F. (2021,
February 1). Membrane microvesicles as potential vaccine candidates. International Journal of
Molecular Sciences. MDPI AG. https://doi.org/10.3390/ijms22031142
St. Cyr, S., Barbee, L., Workowski, K. A., Bachmann, L. H., Pham, C., Schlanger, K., … Thorpe, P.
(2020). Update to CDC’s Treatment Guidelines for Gonococcal Infection, 2020. MMWR. Morbidity
and Mortality Weekly Report, 69(50), 1911–1916. https://doi.org/10.15585/mmwr.mm6950a6
Takada, K., Ohno, N., & Yadomae, T. (1994). Binding of lysozyme to lipopolysaccharide suppresses
tumor necrosis factor production in vivo. Infection and immunity, 62(4), 1171–1175. https://
doi.org/10.1128/iai.62.4.1171-1175.1994
Toyofuku, M., Nomura, N., & Eberl, L. (2019, January 1). Types and origins of bacterial membrane
vesicles. Nature Reviews Microbiology. Nature Publishing Group. https://doi.org/10.1038/
s41579-018-0112-2
United Nations. (s. f.). Las muertes por COVID-19 sumarían 15 millones entre 2020 y 2021 | Naciones
Unidas. https://www.un.org/es/desa/las-muertes-por-covid-19-sumar%C3%ADan-15-millones-
entre-2020-y-2021
Wei, S., Jiao, D., & Xing, W. (2022). A rapid method for isolation of bacterial extracellular vesicles
from culture media using epsilon-poly-L–lysine that enables immunological function research.
Frontiers in Immunology, 13. https://doi.org/10.3389/fimmu.2022.930510
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2023-09-01
Cómo citar
Díaz-Hernández, P. . (2023). VACUNAS CON VESÍCULAS DE MEMBRANA EXTERNA: ¿EL FUTURO DE LAS VACUNAS?. RD-ICUAP, 9(27), 125–137. https://doi.org/10.32399/icuap.rdic.2448-5829.2023.27.1193
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