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Human papillomavirus (HPV) infection constitutes a high-risk factor for HNSCC carcinogenesis. However, its role in TLS formation has yet to be elucidated. Herein, immunohistochemical (IHC) analysis from 59 HNSCC patients revealed a higher prevalence of mature TLS in HPV-positive (HPV\n                    <jats:sup>+<\/jats:sup>\n                    ) HNSCC compared to HPV-negative (HPV\n                    <jats:sup>-<\/jats:sup>\n                    ) cases. Furthermore, integrated analysis of single-cell RNA sequencing, spatial transcriptomics, and RNA-seq data indicated that TLS-positive tumors were characterized by an expanded population of KRT15\n                    <jats:sup>high<\/jats:sup>\n                    tumor cells in HNSCC. IHC and cytological experiments confirmed upregulation of KRT15 in HPV\n                    <jats:sup>+<\/jats:sup>\n                    HNSCC tumor cells, which also showed high expression of cancer stem cell marker genes. These KRT15\n                    <jats:sup>high<\/jats:sup>\n                    stem-like tumor cells specifically secreted CCL20, which was related to the infiltration of TLS-associated immune cells in HPV\n                    <jats:sup>+<\/jats:sup>\n                    HNSCC. Murine models confirmed that CCL20 treatment promoted TLS formation and enhanced the efficacy of anti-PD-1 therapy. Multiplex immunofluorescence showed that TLS provided specialized microenvironments that supported the proliferation of CD39\n                    <jats:sup>+<\/jats:sup>\n                    PD-1\n                    <jats:sup>+<\/jats:sup>\n                    CD8\n                    <jats:sup>+<\/jats:sup>\n                    T cells. Collectively, our findings proposed that CCL20 secreted by HPV-infected KRT15\n                    <jats:sup>high<\/jats:sup>\n                    tumor cells promoted TLS formation, thereby enhancing anti-PD-1 therapy responses in HPV\n                    <jats:sup>+<\/jats:sup>\n                    HNSCC. This study provides mechanistic insights into HPV-mediated TLS development and supports precision immunotherapeutic strategies for HNSCC.\n                  <\/jats:p>","DOI":"10.1038\/s41419-025-08359-5","type":"journal-article","created":{"date-parts":[[2025,12,29]],"date-time":"2025-12-29T07:56:29Z","timestamp":1766994989000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":4,"title":["CCL20 secreted by KRT15high tumor Cells promotes tertiary lymphoid structure formation and enhances anti-PD-1 therapy response in HPV+HNSCC"],"prefix":"10.1038","volume":"17","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-2615-5943","authenticated-orcid":false,"given":"Siwei","family":"Zhang","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Huan","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaoxing","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yourong","family":"Jiang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lu","family":"Tang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tianyang","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rui","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zengchen","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Minghui","family":"Wei","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jingchun","family":"Sun","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhuledesi","family":"Hahan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Heng","family":"Ma","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9472-1965","authenticated-orcid":false,"given":"Lanlan","family":"Wei","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,12,29]]},"reference":[{"key":"8359_CR1","doi-asserted-by":"publisher","first-page":"92","DOI":"10.1038\/s41572-020-00224-3","volume":"6","author":"DE Johnson","year":"2020","unstructured":"Johnson DE, Burtness B, Leemans CR, Lui VWY, Bauman JE, Grandis JR. 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