30. Wu, C. Y., Monie, A., Pang, X., Hung, C. F. & Wu, T. C. Improving therapeutic HPV peptide-based vaccine potency by enhancing CD4 +T help and dendritic cell activation. J. Biomed. Sci. 17, 88 (2010). ACKNOWLEDGEMENTS We would like to thank all the patients and donors who provided the tissues and clinical information for this study. AUTHOR CONTRIBUTIONS The authors con ﬁrm contributions to the paper as follows: study concept and design: W.-Y.P. and D.-S.L.; acquisition of data: S.-M.K., B.L., S.H.K., S.K., and G.-H.N.; statisticalanalysis and interpretation of data: S.-M.K., B.L., S.H.K., S.K., M.K., H.-S.K., Y.C., K.Y.S., andD.-S.L.; drafting of the manuscript: S.-M.K., S.H.K., and K.Y.S; critical revision of themanuscript: S.H.K., K.Y.S., D.-S.L., Y.C.; study supervision; W.-Y.P., D.-S.L, Y.C., K.Y.S., G.-H.N., and I.-S.K. All authors reviewed the results and approved the ﬁnal version of the manuscript. FUNDING This work was supported by the Bio & Medical Technology Development Program ofthe National Research Foundation of Korea (NRF), funded by the Korean government(MSIT) (2017M3A9A7050803 to W.-Y.P.), the Korea Drug Development Fund fundedby the Ministry of Science and ICT, Ministry of Trade, Industry, and Energy, andMinistry of Health and Welfare (RS-2023-00284167 to W.-Y.P.), the Basic ScienceResearch Program through the NRF of Korea (NRF-2021R1A2C1011920 to D.-S.L.), andthe Korea Health Technology R&D Project through the Korea Health Industry Development Institute (KHIDI), funded by the Ministry of Health & Welfare, Korea (HV22C0228). Y.C. was supported by the Medical Research Center Program (NRF-2020R1A5A2031185), the National Immunotherapy Innovation Center (NRF-2020M3A9G3080281), and the Basic Science Research Program (NRF-2021R1F1A1063769). COMPETING INTERESTS The authors declare no competing interests. ETHICS APPROVAL AND CONSENT TO PARTICIPATE This study was approved by the institutional review board (IRB) of Asan MedicalCenter (IRB number: 2022-0263), and informed consent was obtained from allparticipants. All animal experiments were performed in accordance with theguidelines of the Institutional Animal Care and Use Committee (IACUC) of KIST (KIST-IACUC-2022-018-2). ADDITIONAL INFORMATION Supplementary information The online version contains supplementary material available at https://doi.org/10.1038/s12276-024-01259-2 . Correspondence and requests for materials should be addressed to Yoonjoo Choi, Dong-Sup Lee or Woong-Yang Park. Reprints and permission information is available at http://www.nature.com/ reprints Publisher ’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional af ﬁliations. Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third partymaterial in this article are included in the article ’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in thearticle ’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directlyfrom the copyright holder. To view a copy of this licence, visit http:// creativecommons.org/licenses/by/4.0/ . © The Author(s) 2024S.H. Kim et al. 1471 Experimental & Molecular Medicine (2024) 56:1461 – 1471