{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,5]],"date-time":"2026-05-05T02:56:00Z","timestamp":1777949760531,"version":"3.51.4"},"reference-count":75,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,5,4]],"date-time":"2026-05-04T00:00:00Z","timestamp":1777852800000},"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>\n                    In enzootic bovine leukosis (EBL), a B-cell lymphoma caused by bovine leukemia virus (BLV) infection, immune remodeling within tumor-affected lymph nodes is currently poorly understood. Here, we analyzed chemokine production and macrophage phenotypes in tumor-affected lymph nodes in EBL, focusing on CCL4. Intracellular cytokine staining revealed that, compared with healthy lymph nodes, EBL tumor-affected lymph nodes showed increased proportions of CCL4 expressing cells, particularly among CD4\n                    <jats:sup>+<\/jats:sup>\n                    T cells and B cells. Functional migration assays showed that recombinant CCL4 induced robust monocyte migration, with a comparatively modest T-cell migration. To identify lymph node macrophages, we characterized CD11b\n                    <jats:sup>hi<\/jats:sup>\n                    CD172a\n                    <jats:sup>+<\/jats:sup>\n                    cells and confirmed their macrophage identity based on high expression levels of CD11c, CD14, CD16, and CD68. Using this definition, we found that the CD11b\n                    <jats:sup>hi<\/jats:sup>\n                    CD172a\n                    <jats:sup>+<\/jats:sup>\n                    population was markedly increased in tumor-affected lymph nodes. Phenotypic analysis revealed an increased proportion of CD163\n                    <jats:sup>+<\/jats:sup>\n                    and CD163\n                    <jats:sup>+<\/jats:sup>\n                    PD-L1\n                    <jats:sup>+<\/jats:sup>\n                    macrophages, consistent with the acquisition of an immunosuppressive phenotype. In parallel, macrophages in tumor-affected lymph nodes showed a shift from MHC class I\/II\n                    <jats:sup>hi<\/jats:sup>\n                    to MHC I\/II\n                    <jats:sup>lo<\/jats:sup>\n                    subsets and reduced co-expression of MHC molecules with CD80 and CD86, indicating impaired antigen-presenting features. Collectively, these findings suggest that elevated CCL4 production in tumor lesions induced by EBL is associated with monocyte recruitment and the accumulation of phenotypically altered macrophages, highlighting a chemokine-driven remodeling of the tumor immune microenvironment.\n                  <\/jats:p>","DOI":"10.3389\/fimmu.2026.1808815","type":"journal-article","created":{"date-parts":[[2026,5,4]],"date-time":"2026-05-04T04:25:31Z","timestamp":1777868731000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Exhausted CD4+T cells are associated with CCL4-driven immunosuppressive macrophage accumulation in enzootic bovine leukosis"],"prefix":"10.3389","volume":"17","author":[{"given":"Hayato","family":"Nakamura","sequence":"first","affiliation":[{"name":"Department of Disease Control, Faculty of Veterinary Medicine, Hokkaido University","place":["Sapporo, 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Zoonosis Control, Hokkaido University","place":["Sapporo, Japan"]},{"name":"Institute for Vaccine Research and Development (HU-IVReD), Hokkaido University","place":["Sapporo, Japan"]},{"name":"One Health Research Center, Hokkaido University","place":["Sapporo, Japan"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yukinari","family":"Kato","sequence":"additional","affiliation":[{"name":"Department of Antibody Drug Development, Tohoku University Graduate School of Medicine.","place":["Sendai, Japan"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yasuhiko","family":"Suzuki","sequence":"additional","affiliation":[{"name":"Institute for Vaccine Research and Development (HU-IVReD), Hokkaido University","place":["Sapporo, Japan"]},{"name":"Division of Bioresources, International Institute for Zoonosis Control, Hokkaido University","place":["Sapporo, Japan"]},{"name":"Global Station for Zoonosis Control, Global Institution for Collaborative Research and Education (GI-CoRE), Hokkaido University","place":["Sapporo, Japan"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Naoya","family":"Maekawa","sequence":"additional","affiliation":[{"name":"Department of Disease Control, Faculty of Veterinary Medicine, Hokkaido University","place":["Sapporo, Japan"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Shiro","family":"Murata","sequence":"additional","affiliation":[{"name":"Department of Disease Control, Faculty of Veterinary Medicine, Hokkaido University","place":["Sapporo, Japan"]},{"name":"Veterinary Research Unit, International Institute for Zoonosis Control, Hokkaido University","place":["Sapporo, Japan"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Kazuhiko","family":"Ohashi","sequence":"additional","affiliation":[{"name":"Department of Disease Control, Faculty of Veterinary Medicine, Hokkaido University","place":["Sapporo, Japan"]},{"name":"Veterinary Research Unit, International 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