{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,6]],"date-time":"2026-11-06T22:05:46Z","timestamp":1794002746162,"version":"4.8.0"},"reference-count":105,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,2,5]],"date-time":"2026-02-05T00:00:00Z","timestamp":1770249600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Immunol."],"abstract":"<jats:p>Avian influenza virus (AIV) poses a persistent threat to global poultry production and public health. Long-term immunization programs have established a foundational immune barrier, significantly mitigating the risk of highly pathogenic avian influenza outbreaks. However, the high mutability of AIV, complex biosafety requirements, and the accelerating scale of poultry production underscore the need for enhanced mucosal protection. Mucosal immunity represents a critical defense against respiratory virus invasion in poultry, rapidly mobilizing local antibodies, cellular immune responses, and innate defense mechanisms. Recent advances in mucosal vaccine platforms\u2014including viral vectors, nucleic acid vaccines, nanoparticle-based delivery systems, and water- or spray-administered formulations\u2014have demonstrated potential in poultry models to enhance local immune responses, reduce viral shedding, and improve herd-level immune uniformity. This review provides a systematic overview of the unique structural and immunological features of avian mucosal tissues, summarizes the latest developments in mucosal vaccine technologies, and discusses their potential applications and challenges within integrated avian influenza control strategies.<\/jats:p>","DOI":"10.3389\/fimmu.2026.1766957","type":"journal-article","created":{"date-parts":[[2026,2,5]],"date-time":"2026-02-05T06:45:09Z","timestamp":1770273909000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":4,"title":["Advances and prospects of mucosal vaccination in the prevention and control of avian influenza"],"prefix":"10.3389","volume":"17","author":[{"given":"Xianfeng","family":"Hui","sequence":"first","affiliation":[{"name":"Department of Immunology, School of Basic Medical Sciences, Henan Medical University","place":["Xinxiang, China"]},{"name":"Henan Collaborative Innovation Center of Molecular Diagnosis and Laboratory Medicine, School of Medical Technology, Henan Medical University","place":["Xinxiang, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaowei","family":"Tian","sequence":"additional","affiliation":[{"name":"Department of Pathogenic Biology, School of Basic Medical Sciences, Henan Medical University","place":["Xinxiang, China"]},{"name":"Xinxiang Engineering Technology Research Center of Immune Checkpoint Drug for Liver-Intestinal Tumors, Henan Medical University","place":["Xinxiang, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shihuan","family":"Ding","sequence":"additional","affiliation":[{"name":"Department of Immunology, School of Basic Medical Sciences, Henan Medical University","place":["Xinxiang, China"]},{"name":"Henan Collaborative Innovation Center of Molecular Diagnosis and Laboratory Medicine, School of Medical Technology, Henan Medical University","place":["Xinxiang, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ge","family":"Gao","sequence":"additional","affiliation":[{"name":"Henan Collaborative Innovation Center of Molecular Diagnosis and Laboratory Medicine, School of Medical Technology, Henan Medical University","place":["Xinxiang, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shuoxiang","family":"Gao","sequence":"additional","affiliation":[{"name":"Department of Pathogenic Biology, School of Basic Medical Sciences, Henan Medical University","place":["Xinxiang, China"]},{"name":"Xinxiang Engineering Technology Research Center of Immune Checkpoint Drug for Liver-Intestinal Tumors, Henan Medical University","place":["Xinxiang, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aiping","family":"Sun","sequence":"additional","affiliation":[{"name":"Department of Immunology, School of Basic Medical Sciences, Henan Medical University","place":["Xinxiang, China"]},{"name":"Xinxiang Engineering Technology Research Center of Immune Checkpoint Drug for Liver-Intestinal Tumors, Henan Medical University","place":["Xinxiang, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tiesuo","family":"Zhao","sequence":"additional","affiliation":[{"name":"Department of Immunology, School of Basic Medical Sciences, Henan Medical University","place":["Xinxiang, China"]},{"name":"Xinxiang Engineering Technology Research Center of Immune Checkpoint Drug for Liver-Intestinal Tumors, Henan Medical University","place":["Xinxiang, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hui","family":"Wang","sequence":"additional","affiliation":[{"name":"Henan Collaborative Innovation Center of Molecular Diagnosis and Laboratory Medicine, School of Medical Technology, Henan Medical University","place":["Xinxiang, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,2,5]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.03146-14","article-title":"Influenza virus reservoirs and intermediate hosts: dogs, horses, and new possibilities for influenza virus exposure of humans","volume":"89","author":"Parrish","year":"2015","journal-title":"J Virol"},{"key":"B2","doi-asserted-by":"publisher","DOI":"10.1126\/science.aaf8852","article-title":"Role for migratory wild birds in the global spread of avian influenza H5N8","volume":"354","author":"Global Consortium for","year":"2016","journal-title":"Science."},{"key":"B3","doi-asserted-by":"publisher","DOI":"10.46234\/ccdcw2025.235","article-title":"Epidemiological and genetic characterization of three H9N2 viruses causing human infections - changsha city, hunan province, China, april 2025","volume":"7","author":"Huang","year":"2025","journal-title":"China CDC Wkly."},{"key":"B4","doi-asserted-by":"publisher","DOI":"10.3201\/eid3108.250373","article-title":"Transmission dynamics of highly pathogenic avian influenza A(H5N1) and A(H5N6) viruses in wild birds, South Korea, 2023-2024","volume":"31","author":"Seo","year":"2025","journal-title":"Emerg Infect Dis"},{"key":"B5","doi-asserted-by":"publisher","first-page":"2155072","DOI":"10.1080\/22221751.2022.2155072","article-title":"Alarming situation of emerging H5 and H7 avian influenza and effective control strategies","volume":"12","author":"Shi","year":"2023","journal-title":"Emerg Microbes Infect"},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.3201\/eid2501.181259","article-title":"Influenza H5\/H7 virus vaccination in poultry and reduction of zoonotic infections, guangdong province, China, 2017-18","volume":"25","author":"Wu","year":"2019","journal-title":"Emerg Infect Dis"},{"key":"B7","doi-asserted-by":"publisher","first-page":"ofz197","DOI":"10.1093\/ofid\/ofz197","article-title":"Comparison of avian influenza virus contamination in the environment before and after massive poultry H5\/H7 vaccination in zhejiang province, China","volume":"6","author":"Cheng","year":"2019","journal-title":"Open Forum Infect Dis"},{"key":"B8","doi-asserted-by":"publisher","first-page":"1249","DOI":"10.1186\/s12864-024-11152-x","article-title":"Evidence of an emerging triple-reassortant H3N3 avian influenza virus in China","volume":"25","author":"He","year":"2024","journal-title":"BMC Genomics"},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2024.1402235","article-title":"Diversity of genotypes and pathogenicity of H9N2 avian influenza virus derived from wild bird and domestic poultry","volume":"15","author":"Yang","year":"2024","journal-title":"Front Microbiol"},{"key":"B10","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0332213","article-title":"Characterisation of a novel chicken-derived H3N3 avian influenza virus detected in China in 2023: Pathogenicity and immunogenicity","volume":"20","author":"Wang","year":"2025","journal-title":"PloS One"},{"key":"B11","doi-asserted-by":"publisher","DOI":"10.3389\/fphys.2018.00492","article-title":"What makes the harderian gland transcriptome different from other chicken immune tissues? A gene expression comparative analysis","volume":"9","author":"Deist","year":"2018","journal-title":"Front Physiol"},{"key":"B12","doi-asserted-by":"publisher","first-page":"4054","DOI":"10.1038\/s41467-022-31840-0","article-title":"Human-type sialic acid receptors contribute to avian influenza A virus binding and entry by hetero-multivalent interactions","volume":"13","author":"Liu","year":"2022","journal-title":"Nat Commun"},{"key":"B13","doi-asserted-by":"publisher","first-page":"230","DOI":"10.1038\/s41541-025-01280-0","article-title":"Advances and prospects of respiratory mucosal vaccines: mechanisms, technologies, and clinical applications","volume":"10","author":"Li","year":"2025","journal-title":"NPJ Vaccines"},{"key":"B14","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2025.1573037","article-title":"Nanotechnology-driven advances in intranasal vaccine delivery systems against infectious diseases","volume":"16","author":"Zhang","year":"2025","journal-title":"Front Immunol"},{"key":"B15","doi-asserted-by":"publisher","first-page":"480","DOI":"10.1186\/s12951-024-02758-0","article-title":"Tailoring biomaterials for vaccine delivery","volume":"22","author":"Zhuo","year":"2024","journal-title":"J Nanobiotechnology."},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.1021\/acsnano.3c10797","article-title":"Nanoplatform based intranasal vaccines: current progress and clinical challenges","volume":"18","author":"Bai","year":"2024","journal-title":"ACS Nano."},{"key":"B17","doi-asserted-by":"publisher","first-page":"e180784","DOI":"10.1172\/jci.insight.180784","article-title":"An intranasally administered adenovirus-vectored SARS-CoV-2 vaccine induces robust mucosal secretory IgA","volume":"9","author":"Sun","year":"2024","journal-title":"JCI Insight"},{"key":"B18","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1080\/01652176.2023.2228360","article-title":"The immune system of chicken and its response to H9N2 avian influenza virus","volume":"43","author":"Yang","year":"2023","journal-title":"Vet Q"},{"key":"B19","doi-asserted-by":"publisher","first-page":"1185","DOI":"10.1038\/s42003-024-06815-0","article-title":"The gut microbiota and its metabolite butyrate shape metabolism and antiviral immunity along the gut-lung axis in the chicken","volume":"7","author":"Saint-Martin","year":"2024","journal-title":"Commun Biol"},{"key":"B20","doi-asserted-by":"publisher","first-page":"134","DOI":"10.3390\/vaccines12020134","article-title":"Advances in poultry vaccines: leveraging biotechnology for improving vaccine development, stability, and delivery","volume":"12","author":"Abdelaziz","year":"2024","journal-title":"Vaccines (Basel)."},{"key":"B21","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2025.06.044","article-title":"Interferons in health and disease","volume":"188","author":"Boehmer","year":"2025","journal-title":"Cell."},{"key":"B22","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1038\/s41541-022-00485-x","article-title":"Towards the future exploration of mucosal mRNA vaccines against emerging viral diseases; lessons from existing next-generation mucosal vaccine strategies","volume":"7","author":"Hameed","year":"2022","journal-title":"NPJ Vaccines"},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.1093\/infdis\/jiae379","article-title":"An update on highly pathogenic avian influenza A(H5N1) virus, clade 2","volume":"230","author":"Webby","year":"2024","journal-title":"3.4.4b. J Infect Dis"},{"key":"B24","doi-asserted-by":"publisher","first-page":"101070","DOI":"10.1016\/j.lanmic.2024.101070","article-title":"Ocular infectivity and replication of a clade 2.3.4.4b A(H5N1) influenza virus associated with human conjunctivitis in a dairy farm worker in the USA: an in-vitro and ferret study","volume":"6","author":"Belser","year":"2025","journal-title":"Lancet Microbe"},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.02144-08","article-title":"Early control of H5N1 influenza virus replication by the type I interferon response in mice","volume":"83","author":"Szretter","year":"2009","journal-title":"J Virol"},{"key":"B26","doi-asserted-by":"publisher","first-page":"919","DOI":"10.1016\/j.cell.2025.01.019","article-title":"Receptor binding, structure, and tissue tropism of cattle-infecting H5N1 avian influenza virus hemagglutinin","volume":"188","author":"Song","year":"2025","journal-title":"Cell."},{"key":"B27","doi-asserted-by":"publisher","first-page":"1541","DOI":"10.3390\/ijms18071541","article-title":"The interplay between the host receptor and influenza virus hemagglutinin and neuraminidase","volume":"18","author":"Byrd-Leotis","year":"2017","journal-title":"Int J Mol Sci"},{"key":"B28","doi-asserted-by":"publisher","first-page":"e00649-19","DOI":"10.1128\/JVI.00649-19","article-title":"TMPRSS2 is the major activating protease of influenza A virus in primary human airway cells and influenza B virus in human type II pneumocytes","volume":"93","author":"Limburg","year":"2019","journal-title":"J Virol"},{"key":"B29","doi-asserted-by":"publisher","first-page":"e00080-19","DOI":"10.1128\/JVI.00080-19","article-title":"Tropism and infectivity of a seasonal A(H1N1) and a highly pathogenic avian A(H5N1) influenza virus in primary differentiated ferret nasal epithelial cell cultures","volume":"93","author":"Zeng","year":"2019","journal-title":"J Virol"},{"key":"B30","doi-asserted-by":"publisher","DOI":"10.1016\/j.chom.2016.01.001","article-title":"The interaction between respiratory pathogens and mucus","volume":"19","author":"Zanin","year":"2016","journal-title":"Cell Host Microbe"},{"key":"B31","doi-asserted-by":"publisher","DOI":"10.5713\/ab.24.0117","article-title":"In vitro analysis of antiviral immune response against avian influenza virus in chicken tracheal epithelial cells","volume":"37","author":"Heo","year":"2024","journal-title":"Anim Biosci"},{"key":"B32","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.00976-15","article-title":"Repeated low-dose influenza virus infection causes severe disease in mice: a model for vaccine evaluation","volume":"89","author":"Song","year":"2015","journal-title":"J Virol"},{"key":"B33","doi-asserted-by":"publisher","first-page":"182","DOI":"10.1038\/s41426-018-0181-3","article-title":"Pathogenesis and genetic characteristics of novel reassortant low-pathogenic avian influenza H7 viruses isolated from migratory birds in the Republic of Korea in the winter of 2016-2017","volume":"7","author":"Lee","year":"2018","journal-title":"Emerg Microbes Infect"},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.2141\/jpsa.0190091","article-title":"Three-dimensional analysis of the nasolacrimal duct and nasal cavity and arrangement of mucosal tissue in chickens","volume":"57","author":"Watanabe","year":"2020","journal-title":"J Poult Sci"},{"key":"B35","doi-asserted-by":"publisher","first-page":"1418","DOI":"10.3390\/vaccines10091418","article-title":"Immune evaluation of avian influenza virus HAr protein expressed in dunaliella salina in the mucosa of chicken","volume":"10","author":"Castellanos-Huerta","year":"2022","journal-title":"Vaccines (Basel)"},{"key":"B36","doi-asserted-by":"publisher","DOI":"10.1016\/j.vaccine.2018.05.093","article-title":"Examination of the effects of virus inactivation methods on the induction of antibody- and cell-mediated immune responses against whole inactivated H9N2 avian influenza virus vaccines in chickens","volume":"36","author":"Astill","year":"2018","journal-title":"Vaccine."},{"key":"B37","doi-asserted-by":"publisher","first-page":"862","DOI":"10.3390\/v12080862","article-title":"Emerging role of mucosal vaccine in preventing infection with avian influenza A viruses","volume":"12","author":"Wang","year":"2020","journal-title":"Viruses."},{"key":"B38","doi-asserted-by":"publisher","DOI":"10.2478\/v10181-011-0047-2","article-title":"Local immunity of the respiratory mucosal system in chickens and Turkeys","volume":"14","author":"Smialek","year":"2011","journal-title":"Pol J Vet Sci"},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.1016\/j.dci.2017.03.028","article-title":"gammadelta T cells represent a major spontaneously cytotoxic cell population in the chicken","volume":"73","author":"Fenzl","year":"2017","journal-title":"Dev Comp Immunol"},{"key":"B40","doi-asserted-by":"publisher","DOI":"10.1093\/jimmun\/vkaf083","article-title":"Chicken TCRgammadelta+CD8alpha+T cells are antigen-specific and protective in H9N2 AIV infection","volume":"214","author":"Dai","year":"2025","journal-title":"J Immunol"},{"key":"B41","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2013.00185","article-title":"Multi-faceted functions of secretory IgA at mucosal surfaces","volume":"4","author":"Corthesy","year":"2013","journal-title":"Front Immunol"},{"key":"B42","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2025.1576766","article-title":"Compensatory mechanisms in gammadelta T cell-deficient chickens following Salmonella infection","volume":"16","author":"Tetzlaff","year":"2025","journal-title":"Front Immunol"},{"key":"B43","doi-asserted-by":"publisher","first-page":"e2350503","DOI":"10.1002\/eji.202350503","article-title":"Loss of alphabeta but not gammadelta T cells in chickens causes a severe phenotype","volume":"53","author":"von Heyl","year":"2023","journal-title":"Eur J Immunol"},{"key":"B44","doi-asserted-by":"publisher","first-page":"28","DOI":"10.1016\/j.dci.2008.07.018","article-title":"Induction of mucosal immunity in the avian Harderian gland with a replication-deficient Ad5 vector expressing avian influenza H5 hemagglutinin","volume":"33","author":"van Ginkel","year":"2009","journal-title":"Dev Comp Immunol"},{"key":"B45","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.01434-08","article-title":"Association of increased pathogenicity of Asian H5N1 highly pathogenic avian influenza viruses in chickens with highly efficient viral replication accompanied by early destruction of innate immune responses","volume":"83","author":"Suzuki","year":"2009","journal-title":"J Virol"},{"key":"B46","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.01265-06","article-title":"Inhibition of retinoic acid-inducible gene I-mediated induction of beta interferon by the NS1 protein of influenza A virus","volume":"81","author":"Mibayashi","year":"2007","journal-title":"J Virol"},{"key":"B47","doi-asserted-by":"publisher","DOI":"10.1016\/j.chom.2009.04.006","article-title":"Influenza A virus NS1 targets the ubiquitin ligase TRIM25 to evade recognition by the host viral RNA sensor RIG-I","volume":"5","author":"Gack","year":"2009","journal-title":"Cell Host Microbe"},{"key":"B48","doi-asserted-by":"publisher","first-page":"394","DOI":"10.1038\/s41594-018-0058-0","article-title":"Defective germline reprogramming rewires the spermatogonial transcriptome","volume":"25","author":"Vasiliauskaite","year":"2018","journal-title":"Nat Struct Mol Biol"},{"key":"B49","doi-asserted-by":"publisher","first-page":"14751","DOI":"10.1038\/ncomms14751","article-title":"An NS-segment exonic splicing enhancer regulates influenza A virus replication in mammalian cells","volume":"8","author":"Huang","year":"2017","journal-title":"Nat Commun"},{"key":"B50","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.01175-13","article-title":"Increased acid stability of the hemagglutinin protein enhances H5N1 influenza virus growth in the upper respiratory tract but is insufficient for transmission in ferrets","volume":"87","author":"Zaraket","year":"2013","journal-title":"J Virol"},{"key":"B51","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1003831","article-title":"A single amino acid in the stalk region of the H1N1pdm influenza virus HA protein affects viral fusion, stability and infectivity","volume":"10","author":"Cotter","year":"2014","journal-title":"PloS Pathog"},{"key":"B52","doi-asserted-by":"publisher","first-page":"7621","DOI":"10.3390\/ijms22147621","article-title":"LC-ESI-QTOF-MS\/MS Analysis, Cytotoxic, Antiviral, Antioxidant, and Enzyme Inhibitory Properties of Four Extracts of Geranium pyrenaicum Burm. f.: A Good Gift from the Natural Treasure","volume":"22","author":"Swiatek","year":"2021","journal-title":"Int J Mol Sci"},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.1016\/j.it.2009.09.004","article-title":"Innate immune responses to influenza A H5N1: friend or foe","volume":"30","author":"Peiris","year":"2009","journal-title":"Trends Immunol"},{"key":"B54","doi-asserted-by":"publisher","DOI":"10.1038\/nature15514","article-title":"Cleavage of GSDMD by inflammatory caspases determines pyroptotic cell death","volume":"526","author":"Shi","year":"2015","journal-title":"Nature."},{"key":"B55","doi-asserted-by":"publisher","DOI":"10.2353\/ajpath.2007.070248","article-title":"Human and avian influenza viruses target different cells in the lower respiratory tract of humans and other mammals","volume":"171","author":"van Riel","year":"2007","journal-title":"Am J Pathol"},{"key":"B56","doi-asserted-by":"publisher","DOI":"10.1017\/S0950268810003122","article-title":"An overview of the epidemic of highly pathogenic H5N1 avian influenza virus in Egypt: epidemiology and control challenges","volume":"139","author":"Abdelwhab","year":"2011","journal-title":"Epidemiol Infect"},{"key":"B57","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-540-92165-3_6","article-title":"Influenza vaccines for avian species","volume":"333","author":"Kapczynski","year":"2009","journal-title":"Curr Top Microbiol Immunol"},{"key":"B58","doi-asserted-by":"publisher","first-page":"699","DOI":"10.1038\/nri2836","article-title":"Antiviral immune responses in the genital tract: clues for vaccines","volume":"10","author":"Iwasaki","year":"2010","journal-title":"Nat Rev Immunol"},{"key":"B59","doi-asserted-by":"publisher","first-page":"110744","DOI":"10.1016\/j.vetmic.2025.110744","article-title":"Enhanced protection through genotype-matched bivalent H9N2-Newcastle disease virus vaccination: Comparative efficacy against contemporary field strains in specific-pathogen-free chickens","volume":"310","author":"Abdelhalim","year":"2025","journal-title":"Vet Microbiol"},{"key":"B60","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2013.00222","article-title":"Secretory igA: designed for anti-microbial defense","volume":"4","author":"Brandtzaeg","year":"2013","journal-title":"Front Immunol"},{"key":"B61","doi-asserted-by":"publisher","first-page":"e00408-20","DOI":"10.1128\/JVI.00408-20","article-title":"Potential role of nonneutralizing igA antibodies in cross-protective immunity against influenza A viruses of multiple hemagglutinin subtypes","volume":"94","author":"Okuya","year":"2020","journal-title":"J Virol"},{"key":"B62","doi-asserted-by":"publisher","first-page":"a037978","DOI":"10.1101\/cshperspect.a037978","article-title":"Influenza-specific lung-resident memory CD8(+) T cells","volume":"13","author":"van de Wall","year":"2021","journal-title":"Cold Spring Harb Perspect Biol"},{"key":"B63","doi-asserted-by":"publisher","first-page":"e0053021","DOI":"10.1128\/JVI.00530-21","article-title":"CCR2 regulates vaccine-induced mucosal T-cell memory to influenza A virus","volume":"95","author":"Lee","year":"2021","journal-title":"J Virol"},{"key":"B64","doi-asserted-by":"publisher","DOI":"10.1016\/j.vaccine.2016.02.011","article-title":"Vaccination with virus-like particles containing H5 antigens from three H5N1 clades protects chickens from H5N1 and H5N8 influenza viruses","volume":"34","author":"Kapczynski","year":"2016","journal-title":"Vaccine."},{"key":"B65","doi-asserted-by":"publisher","first-page":"2178","DOI":"10.3390\/vaccines10122178","article-title":"Protective efficacy of inactivated H9N2 vaccine in Turkey poults under both experimental and field conditions","volume":"10","author":"Elfeil","year":"2022","journal-title":"Vaccines (Basel)"},{"key":"B66","doi-asserted-by":"publisher","DOI":"10.1056\/NEJMoa066092","article-title":"Duration of humoral immunity to common viral and vaccine antigens","volume":"357","author":"Amanna","year":"2007","journal-title":"N Engl J Med"},{"key":"B67","doi-asserted-by":"publisher","first-page":"36","DOI":"10.1099\/vir.0.024992-0","article-title":"Rapid evolution of low-pathogenic H9N2 avian influenza viruses following poultry vaccination programmes","volume":"92","author":"Park","year":"2011","journal-title":"J Gen Virol"},{"key":"B68","doi-asserted-by":"publisher","DOI":"10.1126\/sciimmunol.abj5129","article-title":"Intranasal priming induces local lung-resident B cell populations that secrete protective mucosal antiviral IgA","volume":"6","author":"Oh","year":"2021","journal-title":"Sci Immunol"},{"key":"B69","doi-asserted-by":"publisher","first-page":"101397","DOI":"10.1016\/j.coviro.2024.101397","article-title":"Tissue-resident memory T cells in protective immunity to influenza virus","volume":"65","author":"Lee","year":"2024","journal-title":"Curr Opin Virol"},{"key":"B70","doi-asserted-by":"publisher","DOI":"10.1038\/s41385-021-00478-4","article-title":"Spatial, temporal and molecular dynamics of swine influenza virus-specific CD8 tissue resident memory T cells","volume":"15","author":"Martini","year":"2022","journal-title":"Mucosal Immunol"},{"key":"B71","doi-asserted-by":"publisher","first-page":"e28","DOI":"10.4110\/in.2021.21.e28","article-title":"A \u201cPrime and deploy\u201d Strategy for universal influenza vaccine targeting nucleoprotein induces lung-resident memory CD8 T cells","volume":"21","author":"Chung","year":"2021","journal-title":"Immune Netw"},{"key":"B72","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1038\/nri.2016.117","article-title":"Early local immune defences in the respiratory tract","volume":"17","author":"Iwasaki","year":"2017","journal-title":"Nat Rev Immunol"},{"key":"B73","doi-asserted-by":"publisher","first-page":"198","DOI":"10.4172\/1747-0862.1000035","article-title":"Avian influenza vaccination in Egypt: Limitations of the current strategy","volume":"3","author":"Peyre","year":"2009","journal-title":"J Mol Genet Med"},{"key":"B74","doi-asserted-by":"publisher","first-page":"2498574","DOI":"10.1080\/22221751.2025.2498574","article-title":"Epidemiology and evolutionary dynamics of H9N2 avian influenza virus in Bangladesh","volume":"14","author":"Islam","year":"2025","journal-title":"Emerg Microbes Infect"},{"key":"B75","doi-asserted-by":"publisher","DOI":"10.3389\/fphys.2022.934413","article-title":"Association between Serum Cystatin C levels and long-term cardiovascular outcomes and all-cause mortality in older patients with obstructive sleep apnea","volume":"13","author":"Li","year":"2022","journal-title":"Front Physiol"},{"key":"B76","doi-asserted-by":"publisher","first-page":"100230","DOI":"10.1016\/j.xcrm.2021.100230","article-title":"A single intranasal dose of chimpanzee adenovirus-vectored vaccine protects against SARS-CoV-2 infection in rhesus macaques","volume":"2","author":"Hassan","year":"2021","journal-title":"Cell Rep Med"},{"key":"B77","doi-asserted-by":"publisher","first-page":"169","DOI":"10.1016\/j.cell.2020.08.026","article-title":"A single-dose intranasal chAd vaccine protects upper and lower respiratory tracts against SARS-coV-2","volume":"183","author":"Hassan","year":"2020","journal-title":"Cell."},{"key":"B78","doi-asserted-by":"publisher","first-page":"592","DOI":"10.1038\/nri3251","article-title":"Recent progress in mucosal vaccine development: potential and limitations","volume":"12","author":"Lycke","year":"2012","journal-title":"Nat Rev Immunol"},{"key":"B79","doi-asserted-by":"publisher","DOI":"10.1016\/j.jconrel.2024.10.031","article-title":"Microparticle and nanoparticle-based influenza vaccines","volume":"376","author":"Ontiveros-Padilla","year":"2024","journal-title":"J Control Release."},{"key":"B80","doi-asserted-by":"publisher","DOI":"10.1021\/acsnano.4c04631","article-title":"An inhalable nanoshield for effective prevention of influenza virus infections","volume":"18","author":"Li","year":"2024","journal-title":"ACS Nano."},{"key":"B81","doi-asserted-by":"publisher","DOI":"10.2147\/IJN.S241492","article-title":"Induction of robust immune responses by cpG-ODN-loaded hollow polymeric nanoparticles for antiviral and vaccine applications in chickens","volume":"15","author":"Lin","year":"2020","journal-title":"Int J Nanomedicine."},{"key":"B82","doi-asserted-by":"publisher","DOI":"10.1016\/j.smim.2017.08.014","article-title":"Major findings and recent advances in virus-like particle (VLP)-based vaccines","volume":"34","author":"Mohsen","year":"2017","journal-title":"Semin Immunol"},{"key":"B83","doi-asserted-by":"publisher","DOI":"10.1038\/nm.3927","article-title":"Hemagglutinin-stem nanoparticles generate heterosubtypic influenza protection","volume":"21","author":"Yassine","year":"2015","journal-title":"Nat Med"},{"key":"B84","doi-asserted-by":"publisher","DOI":"10.1021\/acs.nanolett.5b03528","article-title":"An orthogonal array optimization of lipid-like nanoparticles for mRNA delivery in vivo","volume":"15","author":"Li","year":"2015","journal-title":"Nano Lett"},{"key":"B85","doi-asserted-by":"publisher","DOI":"10.1016\/j.ymthe.2020.04.018","article-title":"A multi-targeting, nucleoside-modified mRNA influenza virus vaccine provides broad protection in mice","volume":"28","author":"Freyn","year":"2020","journal-title":"Mol Ther"},{"key":"B86","doi-asserted-by":"publisher","DOI":"10.1016\/j.ymthe.2018.12.011","article-title":"Improving mRNA-based therapeutic gene delivery by expression-augmenting 3\u2019 UTRs identified by cellular library screening","volume":"27","author":"Orlandini von Niessen","year":"2019","journal-title":"Mol Ther"},{"key":"B87","doi-asserted-by":"publisher","DOI":"10.1038\/nrd4278","article-title":"mRNA-based therapeutics\u2013developing a new class of drugs","volume":"13","author":"Sahin","year":"2014","journal-title":"Nat Rev Drug Discov"},{"key":"B88","doi-asserted-by":"publisher","first-page":"eabh0755","DOI":"10.1126\/scitranslmed.abh0755","article-title":"Intranasal ChAdOx1 nCoV-19\/AZD1222 vaccination reduces viral shedding after SARS-CoV-2 D614G challenge in preclinical models","volume":"13","author":"van Doremalen","year":"2021","journal-title":". Sci Transl Med"},{"key":"B89","doi-asserted-by":"publisher","first-page":"683","DOI":"10.3390\/microorganisms9040683","article-title":"Lactiplantibacillus plantarum as a potential adjuvant and delivery system for the development of SARS-coV-2 oral vaccines","volume":"9","author":"Villena","year":"2021","journal-title":"Microorganisms."},{"key":"B90","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2017.04.008","article-title":"Homeostatic immunity and the microbiota","volume":"46","author":"Belkaid","year":"2017","journal-title":"Immunity."},{"key":"B91","doi-asserted-by":"publisher","first-page":"185","DOI":"10.1186\/s12934-022-01911-4","article-title":"Oral or intranasal immunization with recombinant Lactobacillus plantarum displaying head domain of Swine Influenza A virus hemagglutinin protects mice from H1N1 virus","volume":"21","author":"Zhang","year":"2022","journal-title":"Microb Cell Fact."},{"key":"B92","doi-asserted-by":"publisher","DOI":"10.1128\/JVI.79.5.2910-2919.2005","article-title":"Synthetic double-stranded RNA poly(I:C) combined with mucosal vaccine protects against influenza virus infection","volume":"79","author":"Ichinohe","year":"2005","journal-title":"J Virol"},{"key":"B93","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2017.04.008","article-title":"STING is a direct innate immune sensor of cyclic di-GMP","volume":"478","author":"Burdette","year":"2011","journal-title":"Nature."},{"key":"B94","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2017.04.008","article-title":"Delivery of foreign antigens by engineered outer membrane vesicle vaccines","volume":"107","author":"Chen","year":"2010","journal-title":"Proc Natl Acad Sci U S A."},{"key":"B95","doi-asserted-by":"publisher","first-page":"9647","DOI":"10.1038\/s41467-025-64651-0","article-title":"Live attenuated influenza vaccine with low proportions of defective interfering particles elicits robust immunogenicity and cross-protection","volume":"16","author":"Wu","year":"2025","journal-title":"Nat Commun"},{"key":"B96","doi-asserted-by":"publisher","DOI":"10.1016\/j.vaccine.2020.07.019","article-title":"Efficacy and safety of a live attenuated influenza vaccine in Chinese healthy children aged 3\u201317 years in one study center of a randomized, double-blind, placebo-controlled phase 3 clinical trial, 2016\/17 season","volume":"38","author":"Wang","year":"2020","journal-title":"Vaccine."},{"key":"B97","doi-asserted-by":"publisher","DOI":"10.1016\/S2213-2600(22)00131-X","article-title":"Safety and immunogenicity of a live-attenuated influenza virus vector-based intranasal SARS-CoV-2 vaccine in adults: randomised, double-blind, placebo-controlled, phase 1 and 2 trials","volume":"10","author":"Zhu","year":"2022","journal-title":"Lancet Respir Med"},{"key":"B98","doi-asserted-by":"publisher","first-page":"e0078622","DOI":"10.1128\/jvi.00786-22","article-title":"PSMD12-mediated M1 ubiquitination of influenza A virus at K102 regulates viral replication","volume":"96","author":"Hui","year":"2022","journal-title":"J Virol"},{"key":"B99","doi-asserted-by":"publisher","DOI":"10.1016\/j.ymthe.2017.11.017","article-title":"Self-Amplifying RNA Vaccines Give Equivalent Protection against Influenza to mRNA Vaccines but at Much Lower Doses","volume":"26","author":"Vogel","year":"2018","journal-title":"Mol Ther"},{"key":"B100","doi-asserted-by":"publisher","first-page":"3133","DOI":"10.1038\/s41467-025-58504-z","article-title":"Intranasal influenza virus-vectored vaccine offers protection against clade 2.3.4.4b H5N1 infection in small animal models","volume":"16","author":"Liu","year":"2025","journal-title":"Nat Commun"},{"key":"B101","doi-asserted-by":"publisher","first-page":"2511132","DOI":"10.1080\/22221751.2025.2511132","article-title":"Computationally designed haemagglutinin with nanocage plug-and-display elicits pan-H5 influenza vaccine responses","volume":"14","author":"Huang","year":"2025","journal-title":"Emerg Microbes Infect"},{"key":"B102","doi-asserted-by":"publisher","first-page":"101","DOI":"10.1186\/s13567-025-01506-y","article-title":"Review of respiratory syndromes in poultry: pathogens, prevention, and control measures","volume":"56","author":"Liu","year":"2025","journal-title":"Vet Res"},{"key":"B103","doi-asserted-by":"publisher","first-page":"192","DOI":"10.1637\/aviandiseases-D-24-00007","article-title":"Immune responses in the harderian gland after newcastle disease vaccination in chickens with maternal antibodies","volume":"68","author":"Espejo","year":"2024","journal-title":"Avian Dis"},{"key":"B104","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2019.03064","article-title":"Oral treatments with probiotics and live salmonella vaccine induce unique changes in gut neurochemicals and microbiome in chickens","volume":"10","author":"Redweik","year":"2019","journal-title":"Front Microbiol"},{"key":"B105","doi-asserted-by":"publisher","first-page":"101324","DOI":"10.1016\/j.psj.2021.101324","article-title":"Research Note: Effects of Escherichia coli co-infection on the protective efficacy assessment of two common infectious bronchitis vaccines","volume":"100","author":"Shen","year":"2021","journal-title":"Poult Sci"}],"container-title":["Frontiers in Immunology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2026.1766957\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,2,5]],"date-time":"2026-02-05T06:45:10Z","timestamp":1770273910000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2026.1766957\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,5]]},"references-count":105,"alternative-id":["10.3389\/fimmu.2026.1766957"],"URL":"https:\/\/doi.org\/10.3389\/fimmu.2026.1766957","relation":{},"ISSN":["1664-3224"],"issn-type":[{"value":"1664-3224","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,2,5]]},"article-number":"1766957"}}