{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T21:09:10Z","timestamp":1790629750065,"version":"4.1.0"},"reference-count":83,"publisher":"MDPI AG","issue":"1","license":[{"start":{"date-parts":[[2017,12,25]],"date-time":"2017-12-25T00:00:00Z","timestamp":1514160000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R03 1R03AI123885-01"],"award-info":[{"award-number":["R03 1R03AI123885-01"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["R01 GM 067260"],"award-info":[{"award-number":["R01 GM 067260"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"name":"SutroVax"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Vaccines"],"abstract":"<jats:p>Chlamydia is the most common cause of bacterial sexually transmitted infections worldwide. While infections resolve with antibiotic treatment, this is often neglected in women due to frequent asymptomatic infections, leading to disease progression and severe sequelae (pelvic inflammatory disease, ectopic pregnancy, infertility). Development of a vaccine against Chlamydia is crucial. Whole organism-based vaccines have short-lived activity, serovar\/subgroup-specific immunity and can cause adverse reactions in vaccinated subjects. The Chlamydia major outer membrane protein (MOMP) is a prime candidate for a subunit vaccine. MOMP contains four regions of sequence variability (variable domains, VDs) with B-cell and T-cell epitopes that elicit protective immunity. However, barriers for developing a MOMP-based vaccine include solubility, yield and refolding. We have engineered novel recombinant antigens in which the VDs are expressed into a carrier protein structurally similar to MOMP and suitable for recombinant expression at a high yield in a correctly folded and detergent-free form. Using a carrier such as the PorB porin from the human commensal organism N. lactamica, we show that PorB\/VD chimeric proteins are immunogenic, antigenic and cross-reactive with MOMP. VDs are unique for each serovar but if combined in a single vaccine, a broad coverage against the major Chlamydia serovars can be ensured.<\/jats:p>","DOI":"10.3390\/vaccines6010002","type":"journal-article","created":{"date-parts":[[2017,12,26]],"date-time":"2017-12-26T03:06:38Z","timestamp":1514257598000},"page":"2","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":9,"title":["Structural and Immunological Characterization of Novel Recombinant MOMP-Based Chlamydial Antigens"],"prefix":"10.3390","volume":"6","author":[{"given":"Guillermo","family":"Madico","sequence":"first","affiliation":[{"name":"National Emerging Infectious Diseases Laboratories, Boston University, Boston, MA 02118, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8598-4824","authenticated-orcid":false,"given":"Olga","family":"Gursky","sequence":"additional","affiliation":[{"name":"Department of Physiology &amp; Biophysics and the Amyloidosis Treatment and Research Center, Boston University School of Medicine, Boston, MA 02118, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jeff","family":"Fairman","sequence":"additional","affiliation":[{"name":"SutroVax, Inc., Foster City, CA 94404, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3106-0711","authenticated-orcid":false,"given":"Paola","family":"Massari","sequence":"additional","affiliation":[{"name":"Department of Immunology, Tufts University School of Medicine, Boston, MA 02111, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2017,12,25]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"428","DOI":"10.1056\/NEJM197802232980805","article-title":"Chlamydial infections (first of three parts)","volume":"298","author":"Schachter","year":"1978","journal-title":"N. Engl. J. Med."},{"key":"ref_2","unstructured":"Centers for Disease Control and Prevention (CDC) (2009). Chlamydia screening among sexually active young female enrollees of health plans\u2013United States, 2000\u20132007. MMWR Morb. Mortal. Wkly. Rep., 58, 362\u2013365."},{"key":"ref_3","first-page":"834","article-title":"Prevalence of Chlamydia trachomatis genital infection among persons aged 14\u201339 years\u2014United States, 2007\u20132012","volume":"63","author":"Torrone","year":"2014","journal-title":"MMWR Morb. Mortal. Wkly. Rep."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Newman, L., Rowley, J., Vander, H.S., Wijesooriya, N.S., Unemo, M., Low, N., Stevens, G., Gottlieb, S., Kiarie, J., and Temmerman, M. (2015). Global Estimates of the Prevalence and Incidence of Four Curable Sexually Transmitted Infections in 2012 Based on Systematic Review and Global Reporting. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0143304"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"849","DOI":"10.4049\/jimmunol.90.6.849","article-title":"Classification of trachoma virus strains by protection of mice from toxic death","volume":"90","author":"Wang","year":"1963","journal-title":"J. Immunol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"791","DOI":"10.1093\/infdis\/152.4.791","article-title":"Immunotyping of Chlamydia trachomatis with monoclonal antibodies","volume":"152","author":"Wang","year":"1985","journal-title":"J. Infect. Dis."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1836","DOI":"10.1086\/497341","article-title":"The unexpected impact of a Chlamydia trachomatis infection control program on susceptibility to reinfection","volume":"192","author":"Brunham","year":"2005","journal-title":"J. Infect. Dis."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1080\/00365540110077001","article-title":"Is the increase in notifications of Chlamydia trachomatis infections in Sweden the result of changes in prevalence, sampling frequency or diagnostic methods?","volume":"34","author":"Gotz","year":"2002","journal-title":"Scand. J. Infect. Dis."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"761","DOI":"10.1097\/OLQ.0000000000000028","article-title":"Pelvic inflammatory disease: Identifying research gaps--proceedings of a workshop sponsored by Department of Health and Human Services\/National Institutes of Health\/National Institute of Allergy and Infectious Diseases, November 3\u20134, 2011","volume":"40","author":"Darville","year":"2013","journal-title":"Sex Transm. Dis."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"765","DOI":"10.1056\/NEJMcp1412935","article-title":"Screening for Chlamydia trachomatis Infections in Women","volume":"376","author":"Wiesenfeld","year":"2017","journal-title":"N. Engl. J. Med."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"233","DOI":"10.1093\/infdis\/163.2.233","article-title":"Cofactors in male-female sexual transmission of human immunodeficiency virus type 1","volume":"163","author":"Plummer","year":"1991","journal-title":"J. Infect. Dis."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"230","DOI":"10.1159\/000156481","article-title":"Co-expression of HSV2 and Chlamydia trachomatis in HPV-positive cervical cancer and cervical intraepithelial neoplasia lesions is associated with aberrations in key intracellular pathways","volume":"51","author":"Paba","year":"2008","journal-title":"Intervirology"},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"De la Maza, L.M., Zhong, G., and Brunham, R.C. (2017). Update on Chlamydia trachomatis Vaccinology. Clin. Vaccine Immunol., 24.","DOI":"10.1128\/CVI.00543-16"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"960","DOI":"10.3201\/eid2106.141270","article-title":"Cost-effectiveness of Chlamydia vaccination programs for young women","volume":"21","author":"Chesson","year":"2015","journal-title":"Emerg. Infect. Dis."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"831","DOI":"10.1080\/21645515.2016.1252886","article-title":"Considerations for the rational design of a Chlamydia vaccine","volume":"13","author":"Liang","year":"2017","journal-title":"Hum. Vaccine Immunother."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1097\/QCO.0000000000000343","article-title":"Future prospects for new vaccines against sexually transmitted infections","volume":"30","author":"Gottlieb","year":"2017","journal-title":"Curr. Opin. Infect. Dis."},{"key":"ref_17","unstructured":"CDC (2014). Sexually transmitted diseases surveillance 2013, Prevention."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1055","DOI":"10.1128\/CVI.00131-10","article-title":"Correlates of protection induced by vaccination","volume":"17","author":"Plotkin","year":"2010","journal-title":"Clin. Vaccine Immunol."},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Li, L.X., and McSorley, S.J. (2013). B cells enhance antigen-specific CD4 T cell priming and prevent bacteria dissemination following Chlamydia muridarum genital tract infection. PLoS Pathog., 9.","DOI":"10.1371\/journal.ppat.1003707"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"4374","DOI":"10.1128\/IAI.00622-10","article-title":"CD4+ T cells and antibody are required for optimal major outer membrane protein vaccine-induced immunity to Chlamydia muridarum genital infection","volume":"78","author":"Farris","year":"2010","journal-title":"Infect. Immun."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"7536","DOI":"10.4049\/jimmunol.175.11.7536","article-title":"A predominant role for antibody in acquired immunity to chlamydial genital tract reinfection","volume":"175","author":"Morrison","year":"2005","journal-title":"J. Immunol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1133","DOI":"10.1016\/0002-9394(67)94095-0","article-title":"Pannus with experimental trachoma and inclusion conjunctivitis agent infection of Taiwan monkeys","volume":"63","author":"Wang","year":"1967","journal-title":"Am. J. Ophthalmol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"663","DOI":"10.1084\/jem.169.3.663","article-title":"Chlamydial disease pathogenesis. Ocular hypersensitivity elicited by a genus-specific 57-kD protein","volume":"169","author":"Morrison","year":"1989","journal-title":"J. Exp. Med."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1038\/nri1551","article-title":"Immunology of Chlamydia infection: Implications for a Chlamydia trachomatis vaccine","volume":"5","author":"Brunham","year":"2005","journal-title":"Nat. Rev. Immunol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1161","DOI":"10.1128\/iai.31.3.1161-1176.1981","article-title":"Purification and partial characterization of the major outer membrane protein of Chlamydia trachomatis","volume":"31","author":"Caldwell","year":"1981","journal-title":"Infect. Immun."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1854","DOI":"10.1110\/ps.3650102","article-title":"Prediction of the membrane-spanning beta-strands of the major outer membrane protein of Chlamydia","volume":"11","author":"Bush","year":"2002","journal-title":"Protein Sci."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"6222","DOI":"10.1128\/JB.00552-07","article-title":"Structural and functional analyses of the major outer membrane protein of Chlamydia trachomatis","volume":"189","author":"Sun","year":"2007","journal-title":"J. Bacteriol."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Feher, V.A., Randall, A., Baldi, P., Bush, R.M., de la Maza, L.M., and Amaro, R.E. (2013). A 3-dimensional trimeric beta-barrel model for Chlamydia MOMP contains conserved and novel elements of Gram-negative bacterial porins. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0068934"},{"key":"ref_29","unstructured":"Paul, W.E. (1993). T-cell-mediated immune regulation: Help and suppression. Fundamental Immunology, Raven Press, Ltd.. [3rd ed.]."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"4000","DOI":"10.1073\/pnas.85.11.4000","article-title":"Mapping antigenic domains expressed by Chlamydia trachomatis major outer membrane protein genes","volume":"85","author":"Baehr","year":"1988","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1719","DOI":"10.1128\/IAI.68.3.1719-1723.2000","article-title":"T-cell epitopes in variable segments of Chlamydia trachomatis major outer membrane protein elicit serovar-specific immune responses in infected humans","volume":"68","author":"Ortiz","year":"2000","journal-title":"Infect. Immun."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"4147","DOI":"10.1128\/iai.59.11.4147-4153.1991","article-title":"Functional and structural mapping of Chlamydia trachomatis species-specific major outer membrane protein epitopes by use of neutralizing monoclonal antibodies","volume":"59","author":"Peterson","year":"1991","journal-title":"Infect. Immun."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"575","DOI":"10.1016\/S0264-410X(97)00206-5","article-title":"Monoclonal immunoglobulin A antibody to the major outer membrane protein of the Chlamydia trachomatis mouse pneumonitis biovar protects mice against a chlamydial genital challenge","volume":"15","author":"Pal","year":"1997","journal-title":"Vaccine"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"817","DOI":"10.1084\/jem.167.3.817","article-title":"High-resolution mapping of serovar-specific and common antigenic determinants of the major outer membrane protein of Chlamydia trachomatis","volume":"167","author":"Stephens","year":"1988","journal-title":"J. Exp. Med."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"978","DOI":"10.1093\/infdis\/jiv137","article-title":"Protection Against Chlamydia trachomatis Infection and Upper Genital Tract Pathological Changes by Vaccine-Promoted Neutralizing Antibodies Directed to the VD4 of the Major Outer Membrane Protein","volume":"212","author":"Olsen","year":"2015","journal-title":"J. Infect. Dis."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1126\/science.95.2454.49.b","article-title":"An unidentified virus which produces pneumonia and systemic infection in mice","volume":"95","author":"Nigg","year":"1942","journal-title":"Science"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"2094","DOI":"10.1128\/iai.62.8.3354-3362.1994","article-title":"Intravaginal inoculation of mice with the Chlamydia trachomatis mouse pneumonitis biovar results in infertility","volume":"62","author":"Pal","year":"1994","journal-title":"Infect. Immun."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"5276","DOI":"10.1016\/j.vaccine.2011.05.013","article-title":"Induction of protection against vaginal shedding and infertility by a recombinant Chlamydia vaccine","volume":"29","author":"Carmichael","year":"2011","journal-title":"Vaccine"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"4210","DOI":"10.1128\/IAI.72.7.4210-4216.2004","article-title":"New murine model for the study of Chlamydia trachomatis genitourinary tract infections in males","volume":"72","author":"Pal","year":"2004","journal-title":"Infect. Immun."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"986","DOI":"10.1128\/IAI.00881-10","article-title":"Vaccination against Chlamydia genital infection utilizing the murine C. muridarum model","volume":"79","author":"Farris","year":"2011","journal-title":"Infect. Immun."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"244","DOI":"10.1016\/j.micinf.2013.11.009","article-title":"A vaccine formulated with a combination of TLR-2 and TLR-9 adjuvants and the recombinant major outer membrane protein elicits a robust immune response and significant protection against a Chlamydia muridarum challenge","volume":"16","author":"Cheng","year":"2014","journal-title":"Microbes Infect."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"920","DOI":"10.1016\/j.micinf.2013.08.005","article-title":"Vaccination with major outer membrane protein proteosomes elicits protection in mice against a Chlamydia respiratory challenge","volume":"15","author":"Tifrea","year":"2013","journal-title":"Microbes Infect."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"8063","DOI":"10.4049\/jimmunol.0804375","article-title":"Chlamydia trachomatis native major outer membrane protein induces partial protection in nonhuman primates: Implication for a trachoma transmission-blocking vaccine","volume":"182","author":"Kari","year":"2009","journal-title":"J. Immunol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"766","DOI":"10.1016\/j.vaccine.2005.08.074","article-title":"Immunization with the Chlamydia trachomatis major outer membrane protein, using adjuvants developed for human vaccines, can induce partial protection in a mouse model against a genital challenge","volume":"24","author":"Pal","year":"2006","journal-title":"Vaccine"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1093\/abbs\/gmu016","article-title":"A multi-epitope vaccine based on Chlamydia trachomatis major outer membrane protein induces specific immunity in mice","volume":"46","author":"Tu","year":"2014","journal-title":"Acta Biochim. Biophys. Sin."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"1213","DOI":"10.1016\/j.vaccine.2005.08.097","article-title":"Mucosal immunization with recombinant MOMP genetically linked with modified cholera toxin confers protection against Chlamydia trachomatis infection","volume":"24","author":"Singh","year":"2006","journal-title":"Vaccine"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1023","DOI":"10.1016\/0264-410X(95)00017-U","article-title":"Protective efficacy of a parenterally administered MOMP-derived synthetic oligopeptide vaccine in a murine model of Chlamydia trachomatis genital tract infection: Serum neutralizing IgG antibodies do not protect against chlamydial genital tract infection","volume":"13","author":"Su","year":"1995","journal-title":"Vaccine"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"459","DOI":"10.1016\/S0264-410X(98)00219-9","article-title":"Vaccination of mice with DNA plasmids coding for the Chlamydia trachomatis major outer membrane protein elicits an immune response but fails to protect against a genital challenge","volume":"17","author":"Pal","year":"1999","journal-title":"Vaccine"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"6798","DOI":"10.1128\/IAI.68.12.6798-6806.2000","article-title":"Induction of protective immunity against Chlamydia trachomatis genital infection by a vaccine based on major outer membrane protein-lipophilic immune response-stimulating complexes","volume":"68","author":"Igietseme","year":"2000","journal-title":"Infect. Immun."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"4107","DOI":"10.1007\/s00253-014-5517-x","article-title":"Hepatitis B virus surface antigen as delivery vector can enhance Chlamydia trachomatis MOMP multi-epitope immune response in mice","volume":"98","author":"Zhu","year":"2014","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_51","first-page":"2085","article-title":"Chlamydia trachomatis recombinant MOMP encapsulated in PLGA nanoparticles triggers primarily T helper 1 cellular and antibody immune responses in mice: A desirable candidate nanovaccine","volume":"8","author":"Fairley","year":"2013","journal-title":"Int. J. Nanomed."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1311","DOI":"10.1016\/j.nano.2014.02.009","article-title":"Poly(lactic acid)-poly(ethylene glycol) nanoparticles provide sustained delivery of a Chlamydia trachomatis recombinant MOMP peptide and potentiate systemic adaptive immune responses in mice","volume":"10","author":"Dixit","year":"2014","journal-title":"Nanomedicine"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"5942","DOI":"10.1016\/j.vaccine.2012.07.038","article-title":"In vitro assessment of halobacterial gas vesicles as a Chlamydia vaccine display and delivery system","volume":"30","author":"Childs","year":"2012","journal-title":"Vaccine"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"194","DOI":"10.1016\/j.pep.2011.08.010","article-title":"A novel chimeric MOMP antigen expressed in Escherichia coli, Arabidopsis thaliana, and Daucus carota as a potential Chlamydia trachomatis vaccine candidate","volume":"80","author":"Kalbina","year":"2011","journal-title":"Protein Expr. Purif."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"698","DOI":"10.1016\/j.vaccine.2012.11.039","article-title":"Protective efficacy against Chlamydophila psittaci by oral immunization based on transgenic rice expressing MOMP in mice","volume":"31","author":"Zhang","year":"2013","journal-title":"Vaccine"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"1097","DOI":"10.1128\/IAI.70.3.1097-1105.2002","article-title":"Dendritic cells pulsed with a recombinant chlamydial major outer membrane protein antigen elicit a CD4(+) type 2 rather than type 1 immune response that is not protective","volume":"70","author":"Shaw","year":"2002","journal-title":"Infect. Immun."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"1741","DOI":"10.1128\/IAI.00734-12","article-title":"Vaccination with the recombinant major outer membrane protein elicits antibodies to the constant domains and induces cross-serovar protection against intranasal challenge with Chlamydia trachomatis","volume":"81","author":"Tifrea","year":"2013","journal-title":"Infect. Immun."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"3417","DOI":"10.1128\/IAI.00683-12","article-title":"The amino acid sequence of Neisseria lactamica PorB surface-exposed loops influences Toll-like receptor 2-dependent cell activation","volume":"80","author":"Toussi","year":"2012","journal-title":"Infect. Immun."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"3497","DOI":"10.1093\/nar\/gkg500","article-title":"Multiple sequence alignment with the Clustal series of programs","volume":"31","author":"Chenna","year":"2003","journal-title":"Nucleic Acids Res."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"501","DOI":"10.4049\/jimmunol.177.1.501","article-title":"The meningococcal vaccine candidate GNA1870 binds the complement regulatory protein factor H and enhances serum resistance","volume":"177","author":"Madico","year":"2006","journal-title":"J. Immunol."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"4489","DOI":"10.4049\/jimmunol.178.7.4489","article-title":"Factor H binding and function in sialylated pathogenic neisseriae is influenced by gonococcal, but not meningococcal, porin","volume":"178","author":"Madico","year":"2007","journal-title":"J. Immunol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"2432","DOI":"10.1128\/iai.62.6.2432-2439.1994","article-title":"Expression of large amounts of neisserial porin proteins in Escherichia coli and refolding of the proteins into native trimers","volume":"62","author":"Qi","year":"1994","journal-title":"Infect. Immun."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"W252","DOI":"10.1093\/nar\/gku340","article-title":"SWISS-MODEL: Modelling protein tertiary and quaternary structure using evolutionary information","volume":"42","author":"Biasini","year":"2014","journal-title":"Nucleic Acids Res."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1093\/bioinformatics\/bti770","article-title":"The SWISS-MODEL workspace: A web-based environment for protein structure homology modelling","volume":"22","author":"Arnold","year":"2006","journal-title":"Bioinformatics"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"W576","DOI":"10.1093\/nar\/gkv402","article-title":"NGL Viewer: A web application for molecular visualization","volume":"43","author":"Rose","year":"2015","journal-title":"Nucleic Acids Res."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"440","DOI":"10.1016\/j.jsb.2013.12.006","article-title":"Crystallographic analysis of Neisseria meningitidis PorB extracellular loops potentially implicated in TLR2 recognition","volume":"185","author":"Kattner","year":"2014","journal-title":"J. Struct. Biol."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1016\/j.pep.2005.04.021","article-title":"Improved purification of native meningococcal porin PorB and studies on its structure\/function","volume":"44","author":"Massari","year":"2005","journal-title":"Protein Expr. Purif."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"8153","DOI":"10.1128\/IAI.73.12.8153-8160.2005","article-title":"Vaccination with the Chlamydia trachomatis major outer membrane protein can elicit an immune response as protective as that resulting from inoculation with live bacteria","volume":"73","author":"Pal","year":"2005","journal-title":"Infect. Immun."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"786","DOI":"10.1016\/j.vaccine.2007.11.080","article-title":"The PorB porin from commensal Neisseria lactamica induces Th1 and Th2 immune responses to ovalbumin in mice and is a potential immune adjuvant","volume":"26","author":"Liu","year":"2008","journal-title":"Vaccine"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"1525","DOI":"10.1099\/mic.0.2007\/015479-0","article-title":"Variation in the Neisseria lactamica porin, and its relationship to meningococcal PorB","volume":"154","author":"Bennett","year":"2008","journal-title":"Microbiology"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"2406","DOI":"10.1128\/IAI.67.5.2406-2413.1999","article-title":"Structural and evolutionary inference from molecular variation in neisseria porins","volume":"67","author":"Derrick","year":"1999","journal-title":"Infect. Immun."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"25329","DOI":"10.1074\/jbc.273.39.25329","article-title":"Characterization of the structure, function, and conformational stability of PorB class 3 protein from Neisseria meningitidis. A porin with unusual physicochemical properties","volume":"273","author":"Minetti","year":"1998","journal-title":"J. Biol. Chem."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"4812","DOI":"10.1128\/IAI.70.9.4812-4817.2002","article-title":"Immunization with the Chlamydia trachomatis mouse pneumonitis major outer membrane protein by use of CpG oligodeoxynucleotides as an adjuvant induces a protective immune response against an intranasal chlamydial challenge","volume":"70","author":"Pal","year":"2002","journal-title":"Infect. Immun."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"5314","DOI":"10.1128\/IAI.00681-10","article-title":"Human airway epithelial cell responses to Neisseria lactamica and purified porin via Toll-like receptor 2-dependent signaling","volume":"78","author":"Liu","year":"2010","journal-title":"Infect. Immun."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1128\/IAI.01062-12","article-title":"Toll-Like Receptor 2-Dependent Activity of Native Major Outer Membrane Protein Proteosomes of Chlamydia trachomatis","volume":"81","author":"Massari","year":"2013","journal-title":"Infect. Immun."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"10","DOI":"10.4049\/jimmunol.139.1.10","article-title":"B cell stimulatory factor-1 (interleukin 4) prepares resting murine B cells to secrete IgG1 upon subsequent stimulation with bacterial lipopolysaccharide","volume":"139","author":"Snapper","year":"1987","journal-title":"J. Immunol."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1016\/j.imlet.2004.09.011","article-title":"Do Th1 or Th2 sequence motifs exist in proteins? Identification of amphipatic immunomodulatory domains in Helicobacter pylori catalase","volume":"96","author":"Guy","year":"2005","journal-title":"Immunol. Lett."},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Desclozeaux, M., Robbins, A., Jelocnik, M., Khan, S.A., Hanger, J., Gerdts, V., Potter, A., Polkinghorne, A., and Timms, P. (2017). Immunization of a wild koala population with a recombinant Chlamydia pecorum Major Outer Membrane Protein (MOMP) or Polymorphic Membrane Protein (PMP) based vaccine: New insights into immune response, protection and clearance. PLoS ONE, 12.","DOI":"10.1371\/journal.pone.0178786"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"112","DOI":"10.1093\/infdis\/137.2.112","article-title":"Carriage of Neisseria meningitidis and Neisseria lactamica in infants and children","volume":"137","author":"Gold","year":"1978","journal-title":"J. Infect. Dis."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"591","DOI":"10.1017\/S0950268800066449","article-title":"The Stonehouse survey: Nasopharyngeal carriage of meningococci and Neisseria lactamica","volume":"99","author":"Cartwright","year":"1987","journal-title":"Epidemiol. Infect."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"1512","DOI":"10.1093\/cid\/civ098","article-title":"Nasal Inoculation of the Commensal Neisseria lactamica Inhibits Carriage of Neisseria meningitidis by Young Adults: A Controlled Human Infection Study","volume":"60","author":"Deasy","year":"2015","journal-title":"Clin. Infect. Dis."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"6348","DOI":"10.1128\/IAI.01062-06","article-title":"Immunization with live Neisseria lactamica protects mice against meningococcal challenge and can elicit serum bactericidal antibodies","volume":"74","author":"Li","year":"2006","journal-title":"Infect. Immun."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"3652","DOI":"10.4049\/jimmunol.0902468","article-title":"Neisseria lactamica selectively induces mitogenic proliferation of the naive B cell pool via cell surface Ig","volume":"185","author":"Vaughan","year":"2010","journal-title":"J. Immunol."}],"container-title":["Vaccines"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-393X\/6\/1\/2\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T18:55:29Z","timestamp":1760208929000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-393X\/6\/1\/2"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2017,12,25]]},"references-count":83,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2018,3]]}},"alternative-id":["vaccines6010002"],"URL":"https:\/\/doi.org\/10.3390\/vaccines6010002","relation":{},"ISSN":["2076-393X"],"issn-type":[{"value":"2076-393X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2017,12,25]]}}}