{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,10]],"date-time":"2026-09-10T05:08:37Z","timestamp":1789016917068,"version":"build-2803163510"},"reference-count":117,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2026,9,8]],"date-time":"2026-09-08T00:00:00Z","timestamp":1788825600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100006978","name":"University of California, Berkeley","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100006978","id-type":"DOI","asserted-by":"crossref"}]},{"id":[{"id":"https:\/\/ror.org\/01an7q238","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Pathogens"],"abstract":"<jats:p>Attenuated Salmonella strains are a promising gene delivery vector in the development of vaccines targeting various human diseases. For example, the Ty21a vaccine is a Salmonella-based vaccine that elicits a robust local, cellular, and systemic immune response against Salmonella typhi infection. Human immunodeficiency virus (HIV) is responsible for the AIDS pandemic and requires lifelong treatment to manage. Human papillomavirus (HPV) is the cause of the vast majority of cervical cancer cases in the global population. Influenza is a seasonal, pandemic virus that can lead to high death tolls in humans and livestock and considerable financial losses. In this review, we outline current methods of protection and treatment for HIV, HPV, and influenza, as well as how a versatile Salmonella-based gene delivery vector\u2019s mechanism of action can lead to immunity. We also describe several studies that utilize many of these Salmonella-based vector vaccines to provide protection against infection with these viruses, as many of these Salmonella-based vectors exhibit different factors that may enhance their gene delivery, immune responses, and post-delivery clearance out of host cells. Finally, we discuss the benefits and drawbacks of using Salmonella-based vaccines and future directions for these vaccine candidates, as well as the implications of introducing these vaccines into the current landscape.<\/jats:p>","DOI":"10.3390\/pathogens15090955","type":"journal-article","created":{"date-parts":[[2026,9,8]],"date-time":"2026-09-08T09:02:24Z","timestamp":1788858144000},"page":"955","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Developing Antiviral Vaccines Based on Attenuated Salmonella"],"prefix":"10.3390","volume":"15","author":[{"given":"Ethan","family":"Ou","sequence":"first","affiliation":[{"name":"School of Public Health, University of California, Berkeley, CA 94720, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fenyong","family":"Liu","sequence":"additional","affiliation":[{"name":"School of Public Health, University of California, Berkeley, CA 94720, USA"},{"name":"Program in Comparative Biochemistry, University of California, Berkeley, CA 94720, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,8]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"112","DOI":"10.1038\/sj.icb.7100007","article-title":"Salmonella, the host and disease: A brief review","volume":"85","author":"Coburn","year":"2007","journal-title":"Immunol. Cell Biol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"1305","DOI":"10.1016\/S1286-4579(01)01491-5","article-title":"intracellular proliferation: Where, when and how?","volume":"3","year":"2001","journal-title":"Microbes Infect."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1146\/annurev.med.52.1.259","article-title":"Salmonella: A Model for Bacterial Pathogenesis","volume":"52","author":"Ohl","year":"2001","journal-title":"Annu. Rev. Med."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1586\/14760584.6.2.147","article-title":"Salmonella-based vaccines for infectious diseases","volume":"6","author":"Kwon","year":"2007","journal-title":"Expert Rev. Vaccines"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1128\/mr.57.1.183-289.1993","article-title":"Pathogenesis of human immunodeficiency virus infection","volume":"57","author":"Levy","year":"1993","journal-title":"Microbiol. Rev."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"343","DOI":"10.3934\/biophy.2015.3.343","article-title":"Morphology and ultrastructure of retrovirus particles","volume":"2","author":"Zhang","year":"2015","journal-title":"AIMS Biophys."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Gomez-Lucia, E. (2021). Animal Lentiviruses (Retroviridae), Elsevier.","DOI":"10.1016\/B978-0-12-814515-9.00005-9"},{"key":"ref_8","unstructured":"World Health Organization (2026, July 12). HIV Data and Statistics. Available online: https:\/\/www.who.int\/teams\/global-hiv-hepatitis-and-stis-programmes\/hiv\/strategic-information\/hiv-data-and-statistics."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"a006841","DOI":"10.1101\/cshperspect.a006841","article-title":"Origins of HIV and the AIDS Pandemic","volume":"1","author":"Sharp","year":"2011","journal-title":"Cold Spring Harb. Perspect. Med."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"e6","DOI":"10.4081\/idr.2013.s1.e6","article-title":"Pathogenesis of HIV infection","volume":"5","author":"Naif","year":"2013","journal-title":"Infect. Dis. Rep."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1016\/j.coviro.2016.03.004","article-title":"Current status and prospects of HIV treatment","volume":"18","author":"Cihlar","year":"2016","journal-title":"Curr. Opin. Virol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"765","DOI":"10.1016\/j.idc.2013.09.001","article-title":"HPV and HPV-Associated Diseases","volume":"27","author":"Dunne","year":"2013","journal-title":"Infect. Dis. Clin. N. Am."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Cubie, H.A., Cuschieri, K.S., and Tong, C.Y.W. (2012). Papillomaviruses and Polyomaviruses, Elsevier.","DOI":"10.1016\/B978-0-7020-4089-4.00060-3"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"F55","DOI":"10.1016\/j.vaccine.2012.06.083","article-title":"The Biology and Life-Cycle of Human Papillomaviruses","volume":"30","author":"Doorbar","year":"2012","journal-title":"Vaccine"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1128\/CMR.16.1.1-17.2003","article-title":"Human Papillomavirus and Cervical Cancer","volume":"16","author":"Burd","year":"2003","journal-title":"Clin. Microbiol. Rev."},{"key":"ref_16","unstructured":"World Health Organization (2026, July 12). Human Papillomavirus and Cancer. Available online: https:\/\/www.who.int\/news-room\/fact-sheets\/detail\/human-papilloma-virus-and-cancer."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"D49","DOI":"10.1016\/j.vaccine.2008.07.039","article-title":"The biology of influenza viruses","volume":"26","author":"Bouvier","year":"2008","journal-title":"Vaccine"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"e117","DOI":"10.1016\/S2352-3018(23)00264-3","article-title":"Prophylactic HIV-1 vaccine trials: Past, present, and future","volume":"11","author":"Nkolola","year":"2024","journal-title":"Lancet HIV"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"e95","DOI":"10.1016\/S2666-5247(21)00042-2","article-title":"Preliminary phase 1 results from an HIV vaccine candidate trial","volume":"2","author":"Venkatesan","year":"2021","journal-title":"Lancet Microbe"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"8954","DOI":"10.1128\/JVI.00754-11","article-title":"Mechanism of Neutralization by the Broadly Neutralizing HIV-1 Monoclonal Antibody VRC01","volume":"85","author":"Li","year":"2011","journal-title":"J. Virol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"eadn0223","DOI":"10.1126\/scitranslmed.adn0223","article-title":"Heterologous prime-boost vaccination drives early maturation of HIV broadly neutralizing antibody precursors in humanized mice","volume":"16","author":"Cottrell","year":"2024","journal-title":"Sci. Transl. Med."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"e186927","DOI":"10.1172\/JCI186927","article-title":"Safety and implementation of phase I randomized GLA-SE\u2013adjuvanted CH505TF gp120 HIV vaccine trial in newborns","volume":"135","author":"Violari","year":"2025","journal-title":"J. Clin. Investig."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1016\/S0140-6736(10)60676-9","article-title":"Antiretroviral therapy and management of HIV infection","volume":"376","author":"Volberding","year":"2010","journal-title":"Lancet"},{"key":"ref_24","first-page":"e515","article-title":"Current Scenario of HIV\/AIDS, Treatment Options, and Major Challenges with Compliance to Antiretroviral Therapy","volume":"8","author":"Bhatti","year":"2016","journal-title":"Cureus"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1016\/j.jiac.2016.10.002","article-title":"Current status of HIV\/AIDS in the ART era","volume":"23","author":"Yoshimura","year":"2017","journal-title":"J. Infect. Chemother."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Cheng, L., Wang, Y., and Du, J. (2020). Human Papillomavirus Vaccines: An Updated Review. Vaccines, 8.","DOI":"10.3390\/vaccines8030391"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1263","DOI":"10.2165\/00003495-200666090-00008","article-title":"Human Papillomavirus Quadrivalent (types 6, 11, 16, 18) Recombinant Vaccine (Gardasil??)","volume":"66","author":"Siddiqui","year":"2006","journal-title":"Drugs"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/j.1365-2710.2009.01150.x","article-title":"Quadrivalent human papillomavirus (HPV) vaccine: A review of safety, efficacy, and pharmacoeconomics","volume":"36","author":"Pomfret","year":"2011","journal-title":"J. Clin. Pharm. Ther."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"382","DOI":"10.1093\/infdis\/jiy516","article-title":"Bivalent Human Papillomavirus Vaccine Effectiveness in a Japanese Population: High Vaccine-Type\u2013Specific Effectiveness and Evidence of Cross-Protection","volume":"219","author":"Kudo","year":"2019","journal-title":"J. Infect. Dis."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"1643","DOI":"10.1016\/S1470-2045(20)30524-6","article-title":"Efficacy of the bivalent HPV vaccine against HPV 16\/18-associated precancer: Long-term follow-up results from the Costa Rica Vaccine Trial","volume":"21","author":"Porras","year":"2020","journal-title":"Lancet Oncol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"2143","DOI":"10.1016\/S0140-6736(17)31821-4","article-title":"Final efficacy, immunogenicity, and safety analyses of a nine-valent human papillomavirus vaccine in women aged 16\u201326 years: A randomised, double-blind trial","volume":"390","author":"Huh","year":"2017","journal-title":"Lancet"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"212","DOI":"10.1038\/s41591-018-0340-z","article-title":"Current and future influenza vaccines","volume":"25","author":"Yamayoshi","year":"2019","journal-title":"Nat. Med."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/j.0300-9475.2004.01382.x","article-title":"Influenza Virus: Immunity and Vaccination Strategies. Comparison of the Immune Response to Inactivated and Live, Attenuated Influenza Vaccines","volume":"59","author":"Cox","year":"2004","journal-title":"Scand. J. Immunol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"120","DOI":"10.54254\/2753-8818\/6\/20230193","article-title":"The advantages and disadvantages of different types of vaccines: DNA vaccine, mRNA vaccine, and inactivated vaccine","volume":"6","author":"Chen","year":"2023","journal-title":"Theor. Nat. Sci."},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Mokalla, V.R., Gundarapu, S., Kaushik, R.S., Rajput, M., and Tummala, H. (2025). Influenza Vaccines: Current Status, Adjuvant Strategies, and Efficacy. Vaccines, 13.","DOI":"10.3390\/vaccines13090962"},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Hong, Q., Liu, J., Wei, Y., and Wei, X. (2023). Application of Baculovirus Expression Vector System (BEVS) in Vaccine Development. Vaccines, 11.","DOI":"10.3390\/vaccines11071218"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1007\/BF02686090","article-title":"Intestinal innate immunity and the pathogenesis of Salmonella enteritis","volume":"37","author":"Srikanth","year":"2007","journal-title":"Immunol. Res."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"975","DOI":"10.1080\/21505594.2016.1270493","article-title":"The infectious intracellular lifestyle of Salmonella enterica relies on the adaptation to nutritional conditions within the Salmonella-containing vacuole","volume":"8","author":"Diacovich","year":"2017","journal-title":"Virulence"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"4699","DOI":"10.1128\/JB.184.17.4699-4708.2002","article-title":"Salmonella Type III Secretion-Associated Protein InvE Controls Translocation of Effector Proteins into Host Cells","volume":"184","author":"Kubori","year":"2002","journal-title":"J. Bacteriol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"5826","DOI":"10.1128\/IAI.00375-06","article-title":"Optimization of the Delivery of Heterologous Proteins by the Salmonella enterica Serovar Typhimurium Type III Secretion System for Vaccine Development","volume":"74","author":"Chen","year":"2006","journal-title":"Infect. Immun."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Lou, L., Zhang, P., Piao, R., and Wang, Y. (2019). Salmonella Pathogenicity Island 1 (SPI-1) and Its Complex Regulatory Network. Front. Cell. Infect. Microbiol., 9.","DOI":"10.3389\/fcimb.2019.00270"},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Worley, M.J. (2025). Salmonella Type III Secretion System Effectors. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms26062611"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1016\/j.mib.2008.01.002","article-title":"The Salmonella-containing vacuole\u2014Moving with the times","volume":"11","year":"2008","journal-title":"Curr. Opin. Microbiol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"19414","DOI":"10.1073\/pnas.1217554109","article-title":"Turning self-destructing Salmonella into a universal DNA vaccine delivery platform","volume":"109","author":"Kong","year":"2012","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"238","DOI":"10.1038\/291238a0","article-title":"Aromatic-dependent Salmonella typhimurium are non-virulent and effective as live vaccines","volume":"291","author":"Hoiseth","year":"1981","journal-title":"Nature"},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Mao, M., He, L., and Yan, Q. (2025). An updated overview on the bacterial PhoP\/PhoQ two-component signal transduction system. Front. Cell. Infect. Microbiol., 15.","DOI":"10.3389\/fcimb.2025.1509037"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"C36","DOI":"10.1016\/j.vaccine.2015.04.029","article-title":"Live attenuated vaccines for invasive Salmonella infections","volume":"33","author":"Tennant","year":"2015","journal-title":"Vaccine"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1186\/s12985-025-02645-6","article-title":"Revolutionizing immunization: A comprehensive review of mRNA vaccine technology and applications","volume":"22","author":"Leong","year":"2025","journal-title":"Virol. J."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1719","DOI":"10.1128\/JVI.02639-14","article-title":"Nuclear Import of Adenovirus DNA Involves Direct Interaction of Hexon with an N-Terminal Domain of the Nucleoporin Nup214","volume":"89","author":"Cassany","year":"2015","journal-title":"J. Virol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"270","DOI":"10.1038\/s41591-020-01194-5","article-title":"T cell and antibody responses induced by a single dose of ChAdOx1 nCoV-19 (AZD1222) vaccine in a phase 1\/2 clinical trial","volume":"27","author":"Ewer","year":"2021","journal-title":"Nat. Med."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"594","DOI":"10.1038\/s41586-020-2814-7","article-title":"COVID-19 vaccine BNT162b1 elicits human antibody and TH1 T cell responses","volume":"586","author":"Sahin","year":"2020","journal-title":"Nature"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"3680","DOI":"10.1128\/IAI.67.7.3680-3685.1999","article-title":"Mucosal and Systemic Immune Responses in Humans after Primary and Booster Immunizations with Orally Administered Invasive and Noninvasive Live Attenuated Bacteria","volume":"67","author":"Viret","year":"1999","journal-title":"Infect. Immun."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"233","DOI":"10.1016\/j.ijmm.2008.09.003","article-title":"Genetic stability of vaccine strain Salmonella Typhi Ty21a over 25 years","volume":"299","author":"Kopecko","year":"2009","journal-title":"Int. J. Med. Microbiol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"2640","DOI":"10.1111\/jam.15055","article-title":"Advances in the development of Salmonella-based vaccine strategies for protection against Salmonellosis in humans","volume":"131","author":"Sears","year":"2021","journal-title":"J. Appl. Microbiol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1016\/j.jcmgh.2017.08.002","article-title":"Systemic and Terminal Ileum Mucosal Immunity Elicited by Oral Immunization With the Ty21a Typhoid Vaccine in Humans","volume":"4","author":"Booth","year":"2017","journal-title":"Cell. Mol. Gastroenterol. Hepatol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"e00007-17","DOI":"10.1128\/MMBR.00007-17","article-title":"Vaccination against Salmonella Infection: The Mucosal Way","volume":"81","author":"Gayet","year":"2017","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"3577","DOI":"10.1016\/j.vaccine.2014.04.055","article-title":"Effectiveness of typhoid vaccination in US travelers","volume":"32","author":"Mahon","year":"2014","journal-title":"Vaccine"},{"key":"ref_58","unstructured":"U.S. Food and Drug Administration (2026, August 18). Vivotif\u00ae Typhoid Vaccine Live Oral Ty21a, Available online: https:\/\/www.fda.gov\/media\/75988\/download."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"717","DOI":"10.1093\/infdis\/136.6.717","article-title":"Evaluation of a UDP-Glucose-4-Epimeraseless Mutant of Salmonella typhi as a Live Oral Vaccine","volume":"136","author":"Gilman","year":"1977","journal-title":"J. Infect. Dis."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"292","DOI":"10.1093\/infdis\/145.3.292","article-title":"A Controlled Field Trial of Live Salmonella typhi Strain Ty 21a Oral Vaccine Against Typhoid: Three-Year Results","volume":"145","author":"Wahdan","year":"1982","journal-title":"J. Infect. Dis."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"1049","DOI":"10.1016\/S0140-6736(87)90480-6","article-title":"Large-Scale Field Trial of Ty21a Live Oral Typhoid Vaccine in Enteric-Coated Capsule Formulation","volume":"329","author":"Levine","year":"1987","journal-title":"Lancet"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"845","DOI":"10.1016\/S0264-410X(01)00391-7","article-title":"Safety and immunogenicity of live recombinant Salmonella enterica serovar Typhi Ty21a expressing urease A and B from Helicobacter pylori in human volunteers","volume":"20","author":"Bumann","year":"2001","journal-title":"Vaccine"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"1065","DOI":"10.1136\/gut.2007.145839","article-title":"Correlation of T cell response and bacterial clearance in human volunteers challenged with Helicobacter pylori revealed by randomised controlled vaccination with Ty21a-based Salmonella vaccines","volume":"57","author":"Aebischer","year":"2008","journal-title":"Gut"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"2273","DOI":"10.1016\/j.vaccine.2003.11.020","article-title":"Impact of vector-priming on the immunogenicity of a live recombinant Salmonella enterica serovar typhi Ty21a vaccine expressing urease A and B from Helicobacter pylori in human volunteers","volume":"22","author":"Metzger","year":"2004","journal-title":"Vaccine"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"766","DOI":"10.1093\/infdis\/159.4.766","article-title":"Comparative Efficacy of Two, Three, or Four Doses of TY21a Live Oral Typhoid Vaccine in Enteric-Coated Capsules: A Field Trial in an Endemic Area","volume":"159","author":"Ferreccio","year":"1989","journal-title":"J. Infect. Dis."},{"key":"ref_66","first-page":"209","article-title":"Recombinant Salmonella Vaccination Technology and Its Application to Human Bacterial Pathogens","volume":"14","author":"Zhang","year":"2013","journal-title":"Curr. Pharm. Biotechnol."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"6260","DOI":"10.1158\/0008-5472.CAN-14-1169","article-title":"Effective Cancer Vaccine Platform Based on Attenuated Salmonella and a Type III Secretion System","volume":"74","author":"Xu","year":"2014","journal-title":"Cancer Res."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"2622","DOI":"10.1002\/ijc.24957","article-title":"Novel cancer vaccine based on genes of Salmonella pathogenicity island 2","volume":"126","author":"Xiong","year":"2010","journal-title":"Int. J. Cancer"},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Vendrell, A., Mongini, C., Gravisaco, M.J., Canellada, A., Tesone, A.I., Goin, J.C., and Waldner, C.I. (2016). An Oral Salmonella-Based Vaccine Inhibits Liver Metastases by Promoting Tumor-Specific T-Cell-Mediated Immunity in Celiac and Portal Lymph Nodes: A Preclinical Study. Front. Immunol., 7.","DOI":"10.3389\/fimmu.2016.00072"},{"key":"ref_70","doi-asserted-by":"crossref","unstructured":"Liu, Y., Gong, H., Zhu, J., and Liu, F. (2026). An Oral Vaccine Derived from Attenuated Salmonella Producing Murine Cytomegalovirus M24 Protein Induces Successful Antiviral Immune Responses in Mice. Vaccines, 14.","DOI":"10.3390\/vaccines14030279"},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Liu, Y., Gong, H., Zhu, J., and Liu, F. (2025). Oral Vaccination with Attenuated Salmonella Expressing Viral M25 Protein Effectively Protects Mice Against Murine Cytomegalovirus Infection. Pathogens, 14.","DOI":"10.3390\/pathogens14040314"},{"key":"ref_72","doi-asserted-by":"crossref","unstructured":"Gong, H., Liu, Y., Yan, B., and Liu, F. (2025). Antiviral Immune Responses Against Murine Cytomegalovirus Induced by an Oral Salmonella-Based Vaccine Expressing Viral M33 Protein. Microorganisms, 13.","DOI":"10.3390\/microorganisms13071510"},{"key":"ref_73","doi-asserted-by":"crossref","unstructured":"Liu, Y., Gong, H., Zhu, J., and Liu, F. (2025). An Oral Salmonella-Based Vaccine Expressing Viral M43 Protein Elicits Effective Immunity Against Murine Cytomegalovirus in Mice. Pathogens, 14.","DOI":"10.3390\/pathogens14090902"},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Liu, Y., Gong, H., Zhu, J., and Liu, F. (2025). Effective Immune Protection of Mice from Murine Cytomegalovirus Infection by Oral Salmonella-Based Vaccine Expressing Viral M78 Antigen. Vaccines, 13.","DOI":"10.3390\/vaccines13020137"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"561","DOI":"10.1016\/0264-410X(94)00016-G","article-title":"Construction and characterization of a Salmonella typhi-based human immunodeficiency virus type 1 vector vaccine","volume":"13","author":"Fouts","year":"1995","journal-title":"Vaccine"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"810","DOI":"10.1016\/0264-410X(91)90218-U","article-title":"Construction of genetically defined double aro mutants of Salmonella typhi","volume":"9","author":"Hone","year":"1991","journal-title":"Vaccine"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1186\/1743-422X-6-87","article-title":"Oral vaccination with a recombinant Salmonella vaccine vector provokes systemic HIV-1 subtype C Gag-specific CD4+ Th1 and Th2 cell immune responses in mice","volume":"6","author":"Bourn","year":"2009","journal-title":"Virol. J."},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Klingler, J., Anton, H., R\u00e9al, E., Zeiger, M., Moog, C., M\u00e9ly, Y., and Boutant, E. (2020). How HIV-1 Gag Manipulates Its Host Cell Proteins: A Focus on Interactors of the Nucleocapsid Domain. Viruses, 12.","DOI":"10.3390\/v12080888"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"5733","DOI":"10.1016\/j.vaccine.2012.07.008","article-title":"Salmonella typhi Ty21a bacterial ghost vector augments HIV-1 gp140 DNA vaccine-induced peripheral and mucosal antibody responses via TLR4 pathway","volume":"30","author":"Wen","year":"2012","journal-title":"Vaccine"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"11440","DOI":"10.1073\/pnas.1101414108","article-title":"Trimeric HIV-1 glycoprotein gp140 immunogens and native HIV-1 envelope glycoproteins display the same closed and open quaternary molecular architectures","volume":"108","author":"Harris","year":"2011","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"18542","DOI":"10.1073\/pnas.1422269112","article-title":"Stable, uncleaved HIV-1 envelope glycoprotein gp140 forms a tightly folded trimer with a native-like structure","volume":"111","author":"Kovacs","year":"2014","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"29556","DOI":"10.1038\/srep29556","article-title":"Live attenuated Salmonella displaying HIV-1 10E8 epitope on fimbriae: Systemic and mucosal immune responses in BALB\/c mice by mucosal administration","volume":"6","author":"Li","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"e0331923","DOI":"10.1128\/mbio.03319-23","article-title":"Genome-wide fitness analysis of Salmonella enterica reveals aroA mutants are attenuated due to iron restriction in vitro","volume":"15","author":"Rooke","year":"2024","journal-title":"mBio"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1007\/s002620050302","article-title":"Induction of an antibody response in mice against human papillomavirus (HPV) type 16 after immunization with HPV recombinant Salmonella strains","volume":"43","author":"Krul","year":"1996","journal-title":"Cancer Immunol. Immunother."},{"key":"ref_85","doi-asserted-by":"crossref","unstructured":"Tomai\u0107, V. (2016). Functional Roles of E6 and E7 Oncoproteins in HPV-Induced Malignancies at Diverse Anatomical Sites. Cancers, 8.","DOI":"10.3390\/cancers8100095"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"750","DOI":"10.1128\/IAI.72.2.750-756.2004","article-title":"Immunogenicity against Human Papillomavirus Type 16 Virus-Like Particles Is Strongly Enhanced by the PhoPc Phenotype in Salmonella enterica Serovar Typhimurium","volume":"72","author":"Baud","year":"2004","journal-title":"Infect. Immun."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1016\/j.virol.2013.05.038","article-title":"The papillomavirus major capsid protein L1","volume":"445","author":"Buck","year":"2013","journal-title":"Virology"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"9361","DOI":"10.1073\/pnas.0803801105","article-title":"Regulated programmed lysis of recombinant Salmonella in host tissues to release protective antigens and confer biological containment","volume":"105","author":"Kong","year":"2008","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"20083","DOI":"10.1038\/s41598-021-99425-3","article-title":"Attenuated Salmonella carrying plasmid co-expressing HPV16 L1 and siRNA-E6 for cervical cancer therapy","volume":"11","author":"Chen","year":"2021","journal-title":"Sci. Rep."},{"key":"ref_90","doi-asserted-by":"crossref","unstructured":"Pei, Z., Jiang, X., Yang, Z., Ren, X., Gong, H., Reeves, M., Sheng, J., Wang, Y., Pan, Z., and Liu, F. (2015). Oral Delivery of a Novel Attenuated Salmonella Vaccine Expressing Influenza A Virus Proteins Protects Mice against H5N1 and H1N1 Viral Infection. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0129276"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"102224","DOI":"10.1016\/j.mtbio.2025.102224","article-title":"Harnessing Salmonella as a potent vaccine delivery platform: Targeted HA\/NA epitope presentation via alphavirus RdRp-driven expression to boost immune efficacy","volume":"34","author":"Kwon","year":"2025","journal-title":"Mater. Today Bio"},{"key":"ref_92","doi-asserted-by":"crossref","unstructured":"Hajam, I.A., and Lee, J.H. (2017). An Influenza HA and M2e Based Vaccine Delivered by a Novel Attenuated Salmonella Mutant Protects Mice against Homologous H1N1 Infection. Front. Microbiol., 8.","DOI":"10.3389\/fmicb.2017.00872"},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"38780","DOI":"10.18632\/oncotarget.16353","article-title":"A consensus-hemagglutinin-based vaccine delivered by an attenuated Salmonella mutant protects chickens against heterologous H7N1 influenza virus","volume":"8","author":"Hyoung","year":"2017","journal-title":"Oncotarget"},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"103844","DOI":"10.1016\/j.psj.2024.103844","article-title":"A low-endotoxic Salmonella enterica Gallinarum serovar delivers infectious bronchitis virus immunogens via a dual-promoter vector system that drives protective immune responses through MHC class-I and -II activation in chickens","volume":"103","author":"Hewawaduge","year":"2024","journal-title":"Poult. Sci."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"110264","DOI":"10.1016\/j.vetmic.2024.110264","article-title":"Enhancement of protective efficacy of recombinant attenuated Salmonella typhimurium delivering H9N2 avian influenza virus hemagglutinins(HA) antigen vaccine candidate strains by C-C motif chemokine ligand 5 in chickens(chCCL5)","volume":"298","author":"Ma","year":"2024","journal-title":"Vet. Microbiol."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"110546","DOI":"10.1016\/j.vetmic.2025.110546","article-title":"Recombinant Salmonella vector delivery 3M2e-ferritin fusion nanoparticles provide cross protection against H9N2 and H7N9 avian influenza viruses","volume":"305","author":"Sun","year":"2025","journal-title":"Vet. Microbiol."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"104616","DOI":"10.1016\/j.psj.2024.104616","article-title":"Delivery of dendritic cells targeting 3M2e-HA2 nanoparticles with a CpG adjuvant via lysosomal escape of Salmonella enhances protection against H9N2 avian influenza virus","volume":"104","author":"Jia","year":"2025","journal-title":"Poult. Sci."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"e0142125","DOI":"10.1128\/spectrum.01421-25","article-title":"Protection against H9N2 challenge in chickens via Salmonella with a focA mutation delivered by a dendritic cell-targeted avian influenza virus NA protein tetrame","volume":"13","author":"Ge","year":"2025","journal-title":"Microbiol. Spectr."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1111\/j.1365-2958.2005.04668.x","article-title":"The PhoP\/PhoQ system controls the intramacrophage type three secretion system of Salmonella enterica","volume":"57","author":"Bijlsma","year":"2005","journal-title":"Mol. Microbiol."},{"key":"ref_100","doi-asserted-by":"crossref","unstructured":"Ou, E., and Liu, F. (2026). Anti-Influenza Oral Vaccines Derived from Attenuated Salmonella, IntechOpen.","DOI":"10.5772\/intechopen.1015331"},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1016\/j.jim.2019.06.008","article-title":"Self-destructing Salmonella via temperature induced gene E of phage PhiX174 improves influenza HA DNA vaccine immune protection against H1N1 infection in mice model","volume":"472","author":"Kamble","year":"2019","journal-title":"J. Immunol. Methods"},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"429","DOI":"10.4236\/ojas.2015.54045","article-title":"Molecular Significance of lon and cpxR Genes in the Pathogenicity of Salmonella","volume":"5","author":"Nandre","year":"2015","journal-title":"Open J. Anim. Sci."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"993","DOI":"10.1128\/jvi.44.3.993-1002.1982","article-title":"Lytic action of cloned phi X174 gene E","volume":"44","author":"Young","year":"1982","journal-title":"J. Virol."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"1166","DOI":"10.1038\/s42003-025-08595-7","article-title":"Structural basis for Salmonella infection by two Microviridae phages","volume":"8","author":"Hu","year":"2025","journal-title":"Commun. Biol."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"2","DOI":"10.1186\/s13567-024-01413-8","article-title":"Deletion of pagL and arnT genes involved in LPS structure and charge modulation in the Salmonella genome confer reduced endotoxicity and retained efficient protection against wild-type Salmonella Gallinarum challenge in chicken","volume":"56","author":"Aganja","year":"2025","journal-title":"Vet. Res."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"76","DOI":"10.1186\/s12929-019-0572-3","article-title":"M2e-based universal influenza vaccines: A historical overview and new approaches to development","volume":"26","author":"Mezhenskaya","year":"2019","journal-title":"J. Biomed. Sci."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"104023","DOI":"10.1016\/j.dci.2021.104023","article-title":"Characterization of immunological properties of chicken chemokine CC motif ligand 5 using new monoclonal antibodies","volume":"119","author":"Lu","year":"2021","journal-title":"Dev. Comp. Immunol."},{"key":"ref_108","doi-asserted-by":"crossref","unstructured":"Rodrigues, M.Q., Alves, P.M., and Rold\u00e3o, A. (2021). Functionalizing Ferritin Nanoparticles for Vaccine Development. Pharmaceutics, 13.","DOI":"10.3390\/pharmaceutics13101621"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1084\/jem.20092203","article-title":"CD83 increases MHC II and CD86 on dendritic cells by opposing IL-10\u2013driven MARCH1-mediated ubiquitination and degradation","volume":"208","author":"Tze","year":"2011","journal-title":"J. Exp. Med."},{"key":"ref_110","doi-asserted-by":"crossref","unstructured":"Brandum, E.P., J\u00f8rgensen, A.S., Rosenkilde, M.M., and Hjort\u00f8, G.M. (2021). Dendritic Cells and CCR7 Expression: An Important Factor for Autoimmune Diseases, Chronic Inflammation, and Cancer. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22158340"},{"key":"ref_111","doi-asserted-by":"crossref","unstructured":"Gao, R., Zhang, J., Geng, H., Wang, Y., Kang, X., Geng, S., Jiao, X.A., and Barrow, P. (2022). The Loss of focA Gene Increases the Ability of Salmonella Enteritidis to Exit from Macrophages and Boosts Early Extraintestinal Spread for Systemic Infection in a Mouse Model. Microorganisms, 10.","DOI":"10.3390\/microorganisms10081557"},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"e00440-18","DOI":"10.1128\/IAI.00440-18","article-title":"Improved Tolerability of a Salmonella enterica Serovar Typhimurium Live-Attenuated Vaccine Strain Achieved by Balancing Inflammatory Potential with Immunogenicity","volume":"86","author":"Higginson","year":"2018","journal-title":"Infect. Immun."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"1835","DOI":"10.1128\/IAI.01655-06","article-title":"Candidate Live, Attenuated Salmonella enterica Serotype Typhimurium Vaccines with Reduced Fecal Shedding Are Immunogenic and Effective Oral Vaccines","volume":"75","author":"Jansen","year":"2007","journal-title":"Infect. Immun."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"1269","DOI":"10.1073\/pnas.1113246109","article-title":"Gut inflammation can boost horizontal gene transfer between pathogenic and commensal Enterobacteriaceae","volume":"109","author":"Stecher","year":"2012","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1093\/abbs\/gmt121","article-title":"De-oncogenic HPV E6\/E7 vaccine gets enhanced antigenicity and promotes tumoricidal synergy with cisplatin","volume":"46","author":"Chen","year":"2014","journal-title":"Acta Biochim. Biophys. Sin."},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"2761","DOI":"10.1016\/j.vaccine.2011.01.093","article-title":"Room temperature stabilization of oral, live attenuated Salmonella enterica serovar Typhi-vectored vaccines","volume":"29","author":"Ohtake","year":"2011","journal-title":"Vaccine"},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"5675","DOI":"10.1016\/j.vaccine.2008.08.024","article-title":"Formulation of a live bacterial vaccine for stable room temperature storage results in loss of acid, bile and bile salt resistance","volume":"26","author":"Edwards","year":"2008","journal-title":"Vaccine"}],"container-title":["Pathogens"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-0817\/15\/9\/955\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,10]],"date-time":"2026-09-10T04:18:15Z","timestamp":1789013895000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-0817\/15\/9\/955"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,8]]},"references-count":117,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["pathogens15090955"],"URL":"https:\/\/doi.org\/10.3390\/pathogens15090955","relation":{},"ISSN":["2076-0817"],"issn-type":[{"value":"2076-0817","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,8]]}}}