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Acknowledgements This work was supported by National Natural Science Foundation of China (82388201 and 82350111), National Key Research and Development Program of China (2022YFA1206000), Chinese Academy of Medical Sciences Innovation Fund for Medical Sciences (2021-I2M-1-021 and 2023-I2M-2-005), Noncommunicable Chronic Diseases National Science and Technology Major Project (2025ZD0552500), China Postdoctoral Science Foundation (2024M750249 and 2025T180405) and Haihe Laboratory of Cell Ecosystem Innovation Fund (22HHXBSS00009)