{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T14:29:27Z","timestamp":1787063367423,"version":"build-2736575974"},"reference-count":114,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,6,18]],"date-time":"2026-06-18T00:00:00Z","timestamp":1781740800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100010661","name":"Horizon 2020 Framework Programme","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100010661","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Immunol."],"abstract":"<jats:p>\n                    Domestic pigs constitute a significant global livestock resource and are increasingly recognized as valuable biomedical models, enabling fundamental research. In contrast, immunological studies on wild boar remain limited. To address this knowledge gap, we conducted a comparative analysis of baseline immune phenotypes in age-matched cohorts of domestic pigs and wild boar. Additionally, we adapted a staining and flow cytometry gating strategy based on human immunophenotyping protocols to define conventional memory T cell subsets in both species, validating this approach through proliferation assays and cytokine expression profiling. Our results demonstrated that wild boar exhibit a decreased frequency of polymorphonuclear neutrophils and natural killer cells in peripheral blood. Swine Leukocyte Antigen class II (SLA-DR) expression was significantly reduced across all professional antigen-presenting cell populations. Furthermore, the proportion of T lymphocytes was elevated in wild boar relative to domestic pigs. The T cell compartment in wild boar was characterized by a pronounced bias toward \u03b3\u03b4 T cells, accompanied by reduced frequencies of differentiated CD4\n                    <jats:sup>+<\/jats:sup>\n                    CD8\u03b1\n                    <jats:sup>+<\/jats:sup>\n                    \u03b1\u03b2 T cells and late-stage memory T cells. Notably, these cells still expressed markers indicative of high activation. Our findings highlight intrinsic immunological differences between domestic pigs and wild boar and provide a foundational framework for future functional immunological studies, particularly in the context of infectious disease research and immunization strategies.\n                  <\/jats:p>","DOI":"10.3389\/fimmu.2026.1844742","type":"journal-article","created":{"date-parts":[[2026,6,18]],"date-time":"2026-06-18T04:37:17Z","timestamp":1781757437000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":1,"title":["How the wild things are: comparative T cell phenotyping and memory T cell identification in wild boar and domestic pigs"],"prefix":"10.3389","volume":"17","author":[{"given":"Alexander","family":"Sch\u00e4fer","sequence":"first","affiliation":[{"name":"Institute of Diagnostic Virology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health","place":["Greifswald-Insel Riems, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Virginia","family":"Friedrichs","sequence":"additional","affiliation":[{"name":"Institute of Diagnostic Virology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health","place":["Greifswald-Insel Riems, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Paul","family":"Deutschmann","sequence":"additional","affiliation":[{"name":"Institute of Diagnostic Virology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health","place":["Greifswald-Insel Riems, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Claudia","family":"Klein","sequence":"additional","affiliation":[{"name":"Friedrich-Loeffler-Institut, Institute of Farm Animal Genetics","place":["Neustadt, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sandra","family":"Blome","sequence":"additional","affiliation":[{"name":"Institute of Diagnostic Virology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health","place":["Greifswald-Insel Riems, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ulrike","family":"Blohm","sequence":"additional","affiliation":[{"name":"Institute of Immunology, Friedrich-Loeffler-Institut, Federal Research Institute for Animal Health","place":["Greifswald-Insel Riems, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,6,18]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.1016\/j.molimm.2021.03.014","article-title":"The pig as a medical model for acquired respiratory diseases and dysfunctions: an immunological perspective","volume":"135","author":"Bertho","year":"2021","journal-title":"Mol Immunol"},{"key":"B2","doi-asserted-by":"publisher","DOI":"10.1016\/j.molimm.2021.04.004","article-title":"Swine as biomedical animal model for T-cell research-success and potential for transmittable and non-transmittable human diseases","volume":"135","author":"K\u00e4ser","year":"2021","journal-title":"Mol Immunol"},{"key":"B3","doi-asserted-by":"publisher","DOI":"10.1016\/j.tim.2011.11.002","article-title":"The pig: a model for human infectious diseases","volume":"20","author":"Meurens","year":"2012","journal-title":"Trends Microbiol"},{"key":"B4","doi-asserted-by":"publisher","first-page":"287","DOI":"10.1007\/s00441-020-03206-9","article-title":"The pig as a model for immunology research","volume":"380","author":"Pabst","year":"2020","journal-title":"Cell Tissue Res"},{"key":"B5","doi-asserted-by":"publisher","first-page":"1380","DOI":"10.3389\/fimmu.2019.01380","article-title":"Porcine invariant natural killer T cells: functional profiling and dynamics in steady state and viral infections","volume":"10","author":"Sch\u00e4fer","year":"2019","journal-title":"Front Immunol"},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.1038\/s41590-018-0068-4","article-title":"Recognition of microbial viability via TLR8 drives TFH cell differentiation and vaccine responses","volume":"19","author":"Ugolini","year":"2018","journal-title":"Nat Immunol"},{"key":"B7","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2009.0086","article-title":"Wild boars as sources for infectious diseases in livestock and humans","volume":"364","author":"Meng","year":"2009","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"key":"B8","doi-asserted-by":"publisher","DOI":"10.3389\/fvets.2022.844209","article-title":"The role of the wild boar spreading African swine fever virus in Asia: another underestimated problem","volume":"9","author":"Cadenas-Fern\u00e1ndez","year":"2022","journal-title":"Front Vet Sci"},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.3390\/vetsci7010015","article-title":"The African swine fever epidemic in wild boar (Sus scrofa) in Lithuania (2014-2018)","volume":"7","author":"Ma\u010diulskis","year":"2020","journal-title":"Vet Sci"},{"key":"B10","doi-asserted-by":"publisher","first-page":"4189","DOI":"10.1038\/s41598-019-40962-3","article-title":"Epidemiological evaluation of Latvian control measures for African swine fever in wild boar on the basis of surveillance data","volume":"9","author":"Schulz","year":"2019","journal-title":"Sci Rep"},{"key":"B11","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms14811","article-title":"The comparative immunology of wild and laboratory mice mus musculus domesticus","author":"Abolins","year":"2017","journal-title":"Nat Commun"},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.1038\/nature17655","article-title":"Normalizing the environment recapitulates adult human immune traits in laboratory mice","author":"Beura","year":"2016","journal-title":"Nature"},{"key":"B13","doi-asserted-by":"publisher","DOI":"10.1111\/j.1440-1711.2006.01440.x","article-title":"Increased IL-4 production and attenuated proliferative and pro-inflammatory responses of splenocytes from wild caught rats rattus norvegicus","author":"Lesher","year":"2006","journal-title":"Immunol Cell Biol"},{"key":"B14","doi-asserted-by":"publisher","DOI":"10.1038\/cmi.2011.61","article-title":"Lymphocyte phenotypes in wild caught rats suggest potential mechanisms underlying increased immune sensitivity in post industrial environments","volume":"9","author":"Trama","year":"2012","journal-title":"Cell Mol Immunol"},{"key":"B15","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-294x.2011.05269.x","article-title":"Natural killer cells in free-living Mus musculus have a primed phenotype","volume":"20","author":"Boysen","year":"2011","journal-title":"Mol Ecol"},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.1111\/tbed.12614","article-title":"Biological characterization of African swine fever virus genotype II strains from north-eastern Estonia in European wild boar","volume":"64","author":"Nurmoja","year":"2017","journal-title":"Transbound Emerg Dis"},{"key":"B17","doi-asserted-by":"publisher","first-page":"6510","DOI":"10.1038\/s41598-018-24740-1","article-title":"Deletion at the 5'-end of Estonian ASFV strains associated with an attenuated phenotype","volume":"8","author":"Zani","year":"2018","journal-title":"Sci Rep"},{"key":"B18","doi-asserted-by":"publisher","DOI":"10.22541\/au.159986510.05288008","article-title":"T-cell responses in domestic pigs and wild boar upon infection with the moderately virulent African swine fever virus strain 'Estonia2014'","volume":"68","author":"Sch\u00e4fer","year":"2021","journal-title":"Transbound Emerg Dis"},{"key":"B19","doi-asserted-by":"publisher","DOI":"10.22541\/au.159986510.05288008","article-title":"Impaired T-cell responses in domestic pigs and wild boar upon infection with a highly virulent African swine fever virus strain","volume":"67","author":"H\u00fchr","year":"2020","journal-title":"Transbound Emerg Dis"},{"key":"B20","doi-asserted-by":"publisher","DOI":"10.3390\/pathogens9080662","article-title":"Comparative pathology of domestic pigs and wild boar infected with the moderately virulent African swine fever virus strain \"Estonia 2014","volume":"9","author":"Sehl","year":"2020","journal-title":"Pathogens"},{"key":"B21","doi-asserted-by":"publisher","DOI":"10.3390\/pathogens11020274","article-title":"Adaptive cellular immunity against African swine fever virus infections","volume":"11","author":"Sch\u00e4fer","year":"2022","journal-title":"Pathogens"},{"key":"B22","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.161.10.5338","article-title":"Differentiation of naive CTL to effector and memory CTL: correlation of effector function with phenotype and cell division","volume":"161","author":"Oehen","year":"1998","journal-title":"J Immunol"},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.1006\/cimm.1993.1141","article-title":"Expression and regulation of the porcine CD44 molecule","volume":"149","author":"Yang","year":"1993","journal-title":"Cell Immunol"},{"key":"B24","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.immunol.22.012703.104702","article-title":"Central memory and effector memory T cell subsets: function, generation, and maintenance","volume":"22","author":"Sallusto","year":"2004","journal-title":"Annu Rev Immunol"},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.1038\/35005534","article-title":"Two subsets of memory T lymphocytes with distinct homing potentials and effector functions","volume":"22","author":"Sallusto","year":"1999","journal-title":"Nature"},{"key":"B26","doi-asserted-by":"publisher","DOI":"10.1126\/science.abc8511","article-title":"Deep immune profiling of COVID-19 patients reveals distinct immunotypes with therapeutic implications","volume":"369","author":"Mathew","year":"2020","journal-title":"Science"},{"key":"B27","doi-asserted-by":"publisher","DOI":"10.1101\/2020.09.25.20201863","article-title":"Deep immune profiling of MIS-C demonstrates marked but transient immune activation compared to adult and pediatric COVID-19","volume":"6","author":"Vella","year":"2021","journal-title":"Sci Immunol"},{"key":"B28","doi-asserted-by":"publisher","DOI":"10.1002\/eji.201343751","article-title":"The who's who of T-cell differentiation: human memory T-cell subsets","volume":"43","author":"Mahnke","year":"2013","journal-title":"Eur J Immunol"},{"key":"B29","doi-asserted-by":"publisher","DOI":"10.1002\/jlb.5mr0321-145r","article-title":"Stem cell-like memory T cells: the generation and application","volume":"110","author":"Wang","year":"2021","journal-title":"J Leukoc Biol"},{"key":"B30","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1307572110","article-title":"Virtual memory CD8 T cells display unique functional properties","volume":"110","author":"Lee","year":"2013","journal-title":"Proc Natl Acad Sci USA"},{"key":"B31","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2022.849922","article-title":"Transcriptome profiling of porcine na\u00efve, intermediate and terminally differentiated CD8+ T cells","author":"Lagumdzic","year":"2022","journal-title":"Front Immunol"},{"key":"B32","doi-asserted-by":"crossref","DOI":"10.1016\/B978-0-12-374279-7.12019-3","article-title":"The immune system of swine","volume-title":"Encyclopedia of Immunology","author":"Saalm\u00fcller","year":"2016"},{"key":"B33","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1016\/j.molimm.2014.10.025","article-title":"Phenotypic and functional differentiation of porcine \u03b1\u03b2 T cells: current knowledge and available tools","volume":"66","author":"Gerner","year":"2015","journal-title":"Mol Immunol"},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.3390\/vaccines9020106","article-title":"Maternal autogenous inactivated virus vaccination boosts immunity to PRRSV in piglets","volume":"9","author":"Kick","year":"2021","journal-title":"Vaccine"},{"key":"B35","doi-asserted-by":"publisher","first-page":"20","DOI":"10.1186\/s13567-015-0157-4","article-title":"PCV2 vaccination induces IFN-\u03b3\/TNF-\u03b1 co-producing T cells with a potential role in protection","volume":"46","author":"Koinig","year":"2015","journal-title":"Vet Res"},{"key":"B36","doi-asserted-by":"publisher","first-page":"52","DOI":"10.1186\/s13567-015-0182-3","article-title":"Magnitude and kinetics of multifunctional CD4+ and CD8\u03b2+ T cells in pigs infected with swine influenza A virus","volume":"46","author":"Talker","year":"2015","journal-title":"Vet Res"},{"key":"B37","doi-asserted-by":"publisher","DOI":"10.1128\/jvi.01211-16","article-title":"Influenza A virus infection in pigs attracts multifunctional and cross-reactive T cells to the lung","volume":"90","author":"Talker","year":"2016","journal-title":"J Virol"},{"key":"B38","doi-asserted-by":"publisher","DOI":"10.1186\/1297-9716-44-18","article-title":"CD27 expression discriminates porcine T helper cells with functionally distinct properties","volume":"44","author":"Reutner","year":"2013","journal-title":"Vet Res"},{"key":"B39","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":"B40","doi-asserted-by":"publisher","first-page":"2890","DOI":"10.1038\/s41467-021-23126-8","article-title":"AutoSpill is a principled framework that simplifies the analysis of multichromatic flow cytometry data","volume":"12","author":"Roca","year":"2021","journal-title":"Nat Commun"},{"key":"B41","doi-asserted-by":"publisher","DOI":"10.1016\/j.gene.2023.148089","article-title":"From stability to reliability: unveiling the un-biased reference genes in porcine ovarian granulosa cells under different conditions","volume":"897","author":"Huo","year":"2024","journal-title":"Gene"},{"key":"B42","doi-asserted-by":"publisher","DOI":"10.1007\/BF03195623","article-title":"Verification of reference genes for relative quantification of gene expression by real-time reverse transcription PCR in the pig","volume":"49","author":"Svobodova","year":"2008","journal-title":"J Appl Genet"},{"key":"B43","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1600847","article-title":"T cell fates zipped up: how the Bach2 basic leucine zipper transcriptional repressor directs T cell differentiation and function","volume":"197","author":"Richer","year":"2016","journal-title":"J Immunol"},{"key":"B44","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1306691110","article-title":"Bach2 maintains T cells in a naive state by suppressing effector memory-related genes","volume":"110","author":"Tsukumo","year":"2013","journal-title":"Proc Natl Acad Sci USA"},{"key":"B45","doi-asserted-by":"publisher","first-page":"4706","DOI":"10.1038\/s41467-019-12464-3","article-title":"Single-cell transcriptomics of human T cells reveals tissue and activation signatures in health and disease","volume":"10","author":"Szabo","year":"2019","journal-title":"Nat Commun"},{"key":"B46","doi-asserted-by":"publisher","first-page":"54","DOI":"10.1002\/eji.200636251","article-title":"Functional expression of chemokine receptor CCR6 on human effector memory CD8+ T cells","volume":"37","author":"Kondo","year":"2007","journal-title":"Eur J Immunol"},{"key":"B47","doi-asserted-by":"publisher","first-page":"1473","DOI":"10.4049\/jimmunol.200.supp.51.12","article-title":"Unique phenotypes and clonal expansions of human CD4 effector memory T cells re-expressing CD45RA","volume":"8","author":"Tian","year":"2017","journal-title":"Nat Commun"},{"key":"B48","doi-asserted-by":"publisher","first-page":"21662","DOI":"10.1038\/s41598-021-01260-z","article-title":"Selection and stability validation of reference gene candidates for transcriptional analysis in Rousettus aEgyptiacus","volume":"11","author":"Friedrichs","year":"2021","journal-title":"Sci Rep"},{"key":"B49","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1201150","article-title":"Cutting edge: generation of memory precursors and functional memory CD8+ T cells depends on T cell factor-1 and lymphoid enhancer-binding factor-1","volume":"189","author":"Zhou","year":"2012","journal-title":"J Immunol"},{"key":"B50","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2009.06.021","article-title":"Blimp-1 transcription factor is required for the differentiation of effector CD8(+) T cells and memory responses","volume":"31","author":"Kallies","year":"2009","journal-title":"Immunity"},{"key":"B51","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2022.110858","article-title":"Transcriptional profiling of human V\u03b41 T cells reveals a pathogen-driven adaptive differentiation program","volume":"39","author":"McMurray","year":"2022","journal-title":"Cell Rep"},{"key":"B52","doi-asserted-by":"publisher","first-page":"296","DOI":"10.1016\/j.immuni.2009.05.014","article-title":"Transcriptional repressor Blimp-1 promotes CD8(+) T cell terminal differentiation and represses the acquisition of central memory T cell properties","volume":"31","author":"Rutishauser","year":"2009","journal-title":"Immunity"},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.1111\/imcb.12566","article-title":"SATB1 ensures appropriate transcriptional programs within naive CD8(+) T cells","volume":"100","author":"Nussing","year":"2022","journal-title":"Immunol Cell Biol"},{"key":"B54","doi-asserted-by":"publisher","DOI":"10.1016\/j.it.2019.03.001","article-title":"ZEBs: Novel players in immune cell development and function","volume":"40","author":"Scott","year":"2019","journal-title":"Trends Immunol"},{"key":"B55","doi-asserted-by":"publisher","DOI":"10.1083\/jcb.2113oia258","article-title":"The transcription factors ZEB2 and T-bet cooperate to program cytotoxic T cell terminal differentiation in response to LCMV viral infection","volume":"212","author":"Dominguez","year":"2015","journal-title":"J Exp Med"},{"key":"B56","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20150194","article-title":"Transcriptional repressor ZEB2 promotes terminal differentiation of CD8+ effector and memory T cell populations during infection","volume":"212","author":"Omilusik","year":"2015","journal-title":"J Exp Med"},{"key":"B57","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2017.00325","article-title":"The transcription factor Hobit identifies human cytotoxic CD4(+) T cells","volume":"8","author":"Oja","year":"2017","journal-title":"Front Immunol"},{"key":"B58","doi-asserted-by":"publisher","first-page":"235","DOI":"10.1038\/s41392-023-01471-y","article-title":"T cells in health and disease","volume":"8","author":"Sun","year":"2023","journal-title":"Signal Transduct Target Ther"},{"key":"B59","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2023.06.025","article-title":"Graded expression of the chemokine receptor CX3CR1 marks differentiation states of human and murine T cells and enables cross-species interpretation","volume":"56","author":"Zwijnenburg","year":"2023","journal-title":"Immunity"},{"key":"B60","doi-asserted-by":"publisher","DOI":"10.1016\/j.dci.2022.104462","article-title":"Identification of CD4(+) T cells with T follicular helper cell characteristics in the pig","volume":"134","author":"Hoog","year":"2022","journal-title":"Dev Comp Immunol"},{"key":"B61","doi-asserted-by":"publisher","DOI":"10.3389\/fvets.2024.1390486","article-title":"Investigation of activation-induced markers (AIM) in porcine T cells by flow cytometry","volume":"11","author":"Moorton","year":"2024","journal-title":"Front Vet Sci"},{"key":"B62","doi-asserted-by":"publisher","DOI":"10.1016\/s0165-2427(02)00045-4","article-title":"T-helper cells from naive to committed","volume":"87","author":"Saalm\u00fcller","year":"2002","journal-title":"Vet Immunol Immunopathol"},{"key":"B63","doi-asserted-by":"publisher","DOI":"10.1016\/j.dci.2012.11.010","article-title":"Analysis of chemokine receptor CCR7 expression on porcine blood T lymphocytes using a CCL19-Fc fusion protein","volume":"39","author":"Moreno","year":"2013","journal-title":"Dev Comp Immunol"},{"key":"B64","doi-asserted-by":"publisher","DOI":"10.1016\/j.dci.2016.02.022","article-title":"Expression of T -bet, Eomesodermin and GATA-3 in porcine \u03b1\u03b2 T cells","volume":"60","author":"Rodriguez-Gomez","year":"2016","journal-title":"Dev Comp Immunol"},{"key":"B65","doi-asserted-by":"publisher","first-page":"396","DOI":"10.3389\/fimmu.2019.00396","article-title":"Expression of T-Bet, Eomesodermin, and GATA-3 correlates with distinct phenotypes and functional properties in porcine \u03b3\u03b4 T cells","volume":"10","author":"Rodriguez-Gomez","year":"2019","journal-title":"Front Immunol"},{"key":"B66","doi-asserted-by":"publisher","DOI":"10.1016\/s0021-9258(18)34459-4","article-title":"Direct activation of calcium-activated, phospholipid-dependent protein kinase by tumor-promoting phorbol esters","volume":"257","author":"Castagna","year":"1982","journal-title":"J Biol Chem"},{"key":"B67","doi-asserted-by":"publisher","DOI":"10.1021\/ol025872f","article-title":"Total synthesis of ionomycin using ring-opening strategies","volume":"4","author":"Lautens","year":"2002","journal-title":"Org Lett"},{"key":"B68","article-title":"The use of concanavalin A to study the immunoregulation of human T cells","volume":"46","author":"Dwyer","year":"1981","journal-title":"Clin Exp Immunol"},{"key":"B69","doi-asserted-by":"publisher","DOI":"10.1016\/s0021-9258(18)66787-0","article-title":"The nucleotide and partial amino acid sequence of toxic shock syndrome toxin-1","volume":"261","author":"Blomster-Hautamaa","year":"1986","journal-title":"J Biol Chem"},{"key":"B70","doi-asserted-by":"publisher","DOI":"10.1007\/s00251-013-0718-5","article-title":"How immunogenetically different are domestic pigs from wild boars: a perspective from single-nucleotide polymorphisms of 19 immunity-related candidate genes","volume":"65","author":"Chen","year":"2013","journal-title":"Immunogenetics"},{"key":"B71","doi-asserted-by":"publisher","DOI":"10.1038\/nature11622","article-title":"Analyses of pig genomes provide insight into porcine demography and evolution","volume":"491","author":"Groenen","year":"2012","journal-title":"Nature"},{"key":"B72","doi-asserted-by":"publisher","DOI":"10.1080\/15592294.2020.1712814","article-title":"The emerging role of epigenetics in the immune response to vaccination and infection: a systematic review","volume":"15","author":"Bannister","year":"2020","journal-title":"Epigenetics"},{"key":"B73","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2656.2008.01413.x","article-title":"The dynamics of health in wild field vole populations: a haematological perspective","volume":"77","author":"Beldomenico","year":"2008","journal-title":"J Anim Ecol"},{"key":"B74","doi-asserted-by":"publisher","first-page":"893","DOI":"10.1111\/j.1365-294x.2010.04907.x","article-title":"The analysis of immunological profiles in wild animals: a case study on immunodynamics in the field vole, Microtus agrestis","volume":"20","author":"Jackson","year":"2011","journal-title":"Mol Ecol"},{"key":"B75","doi-asserted-by":"publisher","DOI":"10.2307\/1939397","article-title":"Temporal variation in humoral and cell-mediated immune response in a Sigmodon hispidus population","volume":"75","author":"Lochmiller","year":"1994","journal-title":"Ecology"},{"key":"B76","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2007.2142","article-title":"Seasonal changes in vertebrate immune activity: mediation by physiological trade-offs","volume":"363","author":"Martin","year":"2008","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"key":"B77","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2025.1530214","article-title":"Pig jejunal single-cell RNA landscapes revealing breed-specific immunology differentiation at various domestication stages","volume":"16","author":"Fu","year":"2025","journal-title":"Front Immunol"},{"key":"B78","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1186\/s12862-024-02340-0","article-title":"Inferring the energy cost of resistance to parasitic infection and its link to a trade-off","volume":"25","author":"Douhard","year":"2025","journal-title":"BMC Ecol Evol"},{"key":"B79","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.2018.0489","article-title":"The energetic cost of parasitism in a wild population","volume":"285","author":"Hicks","year":"2018","journal-title":"Proc Biol Sci"},{"key":"B80","doi-asserted-by":"publisher","DOI":"10.1111\/mam.12015","article-title":"A review of wild boar us scrofa diet and factors affecting food selection in native and introduced ranges","volume":"44","author":"Ballari","year":"2014","journal-title":"Mammal Rev"},{"key":"B81","doi-asserted-by":"publisher","DOI":"10.25225\/fozo.v67.i3-4.a3.2018","article-title":"The importance of natural food in wild boar (Sus scrofa) diet during autumn and winter","volume":"67","author":"Ond\u0159ej","year":"2018","journal-title":"Folia Zool"},{"key":"B82","doi-asserted-by":"publisher","DOI":"10.1093\/jn\/136.5.1141","article-title":"Malnutrition and energy restriction differentially affect viral immunity","volume":"136","author":"Ritz","year":"2006","journal-title":"J Nutr"},{"key":"B83","doi-asserted-by":"publisher","DOI":"10.1046\/j.1365-2567.1999.00694.x","article-title":"T cells with a quiescent phenotype (CD45RA+) are overabundant in the blood and involuted lymphoid tissues in wasting protein and energy deficiencies","volume":"96","author":"Woodward","year":"1999","journal-title":"Immunology"},{"key":"B84","doi-asserted-by":"publisher","first-page":"5592","DOI":"10.1038\/ncomms6592","article-title":"DNA methylation signatures link prenatal famine exposure to growth and metabolism","volume":"5","author":"Tobi","year":"2014","journal-title":"Nat Commun"},{"key":"B85","doi-asserted-by":"publisher","DOI":"10.1126\/science.1076064","article-title":"Direct link between mhc polymorphism, T cell avidity, and diversity in immune defense","volume":"298","author":"Messaoudi","year":"2002","journal-title":"Science"},{"key":"B86","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2007.07.007","article-title":"Naive CD4(+) T cell frequency varies for different epitopes and predicts repertoire diversity and response magnitude","volume":"27","author":"Moon","year":"2007","journal-title":"Immunity"},{"key":"B87","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1007\/bf01311692","article-title":"Comparative evaluation of disease dynamics in wild boar and domestic pigs experimentally inoculated intranasally with the European highly virulent African swine fever virus genotype II strain \"Armenia 2007","volume":"55","author":"Sanchez-Cordon","year":"2024","journal-title":"Vet Res"},{"key":"B88","doi-asserted-by":"publisher","DOI":"10.1002\/eji.202250069","article-title":"Immune-mediated pathology as a consequence of impaired immune reactions: the IMPATH paradox","volume":"52","author":"Ehl","year":"2022","journal-title":"Eur J Immunol"},{"key":"B89","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20030235","article-title":"Differentiation of effector\/memory V\u03b42 T cells and migratory routes in lymph nodes or inflammatory sites","volume":"198","author":"Dieli","year":"2003","journal-title":"J Exp Med"},{"key":"B90","doi-asserted-by":"publisher","DOI":"10.1182\/blood-2008-01-136713","article-title":"Long-term expansion of effector\/memory V\u03b42-\u03b3\u03b4 T cells is a specific blood signature of CMV infection","volume":"112","author":"Pitard","year":"2008","journal-title":"Blood"},{"key":"B91","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/s0008-8749(02)00519-1","article-title":"Differentiation of human \u03b3\u03b4 T cells towards distinct memory phenotypes","volume":"218","author":"Eberl","year":"2002","journal-title":"Cell Immunol"},{"key":"B92","doi-asserted-by":"publisher","DOI":"10.1007\/s12250-018-0073-7","article-title":"Phenotypic characterization of porcine IFN\u03b3-producing lymphocytes in porcine reproductive and respiratory syndrome virus vaccinated and challenged pigs","volume":"33","author":"Li","year":"2018","journal-title":"Virol Sin"},{"key":"B93","doi-asserted-by":"publisher","DOI":"10.1089\/vim.2005.18.490","article-title":"\u03b3\u03b4 lymphocyte response to porcine reproductive and respiratory syndrome virus","volume":"18","author":"Olin","year":"2005","journal-title":"Viral Immunol"},{"key":"B94","doi-asserted-by":"publisher","DOI":"10.1016\/j.vaccine.2011.10.098","article-title":"Vaccination with NS1-truncated H3N2 swine influenza virus primes T cells and confers cross-protection against an H1N1 heterosubtypic challenge in pigs","volume":"30","author":"Kappes","year":"2012","journal-title":"Vaccine"},{"key":"B95","doi-asserted-by":"publisher","DOI":"10.3390\/ani11082165","article-title":"Comprehensive transcriptomic comparison between porcine CD8(-) and CD8(+) \u03b3\u03b4 T cells revealed distinct immune phenotype","volume":"11","author":"Kim","year":"2021","journal-title":"Anim (Basel)"},{"key":"B96","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.jim.2004.10.002","article-title":"\u03b3\u03b4 T-lymphocyte cytotoxic activity against Mycobacterium bovis analyzed by flow cytometry","volume":"297","author":"Olin","year":"2005","journal-title":"J Immunol Methods"},{"key":"B97","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1016\/j.dci.2014.02.008","article-title":"CD2 and CD8\u03b1 define porcine \u03b3\u03b4 T cells with distinct cytokine production profiles","volume":"45","author":"Sedlak","year":"2014","journal-title":"Dev Comp Immunol"},{"key":"B98","doi-asserted-by":"publisher","first-page":"45","DOI":"10.1046\/j.1365-2567.1997.00308.x","article-title":"Differential expression of CD8 epitopes amongst porcine CD8-positive functional lymphocyte subsets","volume":"92","author":"Yang","year":"1997","journal-title":"Immunology"},{"key":"B99","doi-asserted-by":"publisher","DOI":"10.1038\/ni.3483","article-title":"Human memory T cells with a naive phenotype accumulate with aging and respond to persistent viruses","volume":"17","author":"Pulko","year":"2016","journal-title":"Nat Immunol"},{"key":"B100","doi-asserted-by":"publisher","DOI":"10.3390\/vaccines6040084","article-title":"T cell memory to vaccination","volume":"6","author":"Todryk","year":"2018","journal-title":"Vaccines (Basel)"},{"key":"B101","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0323823","article-title":"Rabies vaccination induces a CD4+ TEM and CD4+CD8+ TEMRA TH1 phenotype in dogs","volume":"20","author":"Lang","year":"2025","journal-title":"PloS One"},{"key":"B102","doi-asserted-by":"publisher","DOI":"10.3390\/pathogens14020119","article-title":"Evaluation of the dose of African Swine Fever Virus required to establish infection in pigs following oral uptake","volume":"14","author":"Olesen","year":"2025","journal-title":"Pathogens"},{"key":"B103","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2018.02.010","article-title":"Understanding subset diversity in T cell memory","volume":"48","author":"Jameson","year":"2018","journal-title":"Immunity"},{"key":"B104","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2016.10.018","article-title":"The chemokine receptor CX3CR1 defines three antigen-experienced CD8 T cell subsets with distinct roles in immune surveillance and homeostasis","volume":"45","author":"Gerlach","year":"2016","journal-title":"Immunity"},{"key":"B105","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2013.00455","article-title":"Inhibitory receptor expression depends more dominantly on differentiation and activation than \"exhaustion\" of human CD8 T cells","volume":"4","author":"Legat","year":"2013","journal-title":"Front Immunol"},{"key":"B106","doi-asserted-by":"publisher","DOI":"10.1111\/acel.12675","article-title":"Human CD8(+) EMRA T cells display a senescence-associated secretory phenotype regulated by p38 MAPK","volume":"17","author":"Callender","year":"2018","journal-title":"Aging Cell"},{"key":"B107","doi-asserted-by":"publisher","DOI":"10.1016\/j.vetimm.2025.110983","article-title":"Breed-specific humoral immune responses to lumpy skin disease vaccination and its associated factors in cattle","volume":"287","author":"Bok","year":"2025","journal-title":"Vet Immunol Immunopathol"},{"key":"B108","doi-asserted-by":"publisher","DOI":"10.3390\/ani13101642","article-title":"Comparison of immune-related gene expression in two chicken breeds following infectious bronchitis virus vaccination","volume":"13","author":"Chuwatthanakhajorn","year":"2023","journal-title":"Anim (Basel)"},{"key":"B109","doi-asserted-by":"publisher","DOI":"10.1016\/j.psj.2025.105565","article-title":"Baseline immune profiles of local chicken breeds: linking biodiversity, animal health, and vaccination response","volume":"104","author":"Freier","year":"2025","journal-title":"Poult Sci"},{"key":"B110","doi-asserted-by":"publisher","DOI":"10.1093\/jas\/sky288","article-title":"Breed-specific differences in the immune response to lipopolysaccharide in ewes","volume":"96","author":"Hadfield","year":"2018","journal-title":"J Anim Sci"},{"key":"B111","doi-asserted-by":"publisher","DOI":"10.3390\/genes15020205","article-title":"Differences in immune characteristics and related gene expression in spleen among Ningxiang, Berkshire breeds and their hybrid pigs","volume":"15","author":"Song","year":"2024","journal-title":"Genes (Basel)"},{"key":"B112","doi-asserted-by":"publisher","DOI":"10.3389\/fgene.2022.800178","article-title":"Integration of transcriptome and proteome in lymph nodes reveal the different immune responses to PRRSV between PRRSV-resistant Tongcheng pigs and PRRSV-susceptible Large White pigs","volume":"13","author":"Liang","year":"2022","journal-title":"Front Genet"},{"key":"B113","doi-asserted-by":"publisher","DOI":"10.3389\/fvets.2023.1289570","article-title":"Genetic background influences pig responses to porcine reproductive and respiratory syndrome virus","volume":"10","author":"Pei","year":"2023","journal-title":"Front Vet Sci"},{"key":"B114","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2022.960709","article-title":"Integrated time-series transcriptomic and metabolomic analyses reveal different inflammatory and adaptive immune responses contributing to host resistance to PRRSV","volume":"13","author":"Wu","year":"2022","journal-title":"Front Immunol"}],"container-title":["Frontiers in Immunology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2026.1844742\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,18]],"date-time":"2026-06-18T04:37:26Z","timestamp":1781757446000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2026.1844742\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6,18]]},"references-count":114,"alternative-id":["10.3389\/fimmu.2026.1844742"],"URL":"https:\/\/doi.org\/10.3389\/fimmu.2026.1844742","relation":{},"ISSN":["1664-3224"],"issn-type":[{"value":"1664-3224","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,6,18]]},"article-number":"1844742"}}