We thank the Center of Cryo-Electron Microscopy, and Core Facility of Shanghai Medical College, Fudan University for technical support. This work was supported by the National Key R&D Program of China (2023YFA0915000 to Q.Q. & J.H.), the National Science Fund for Distinguished Young Scholars (82425015 to J.H.), the National Natural Science Foundation of China (32171194 & 32371256 to Q.Q., 82151215 to D.L., 82171102 to J.H., 32401007 to Z.Z.), the National Science and Technology Major Project of the Ministry of Science and Technology of China (2023ZD0503203 to Q.Q.), the Strategic Priority Research Program of CAS (XDB37020204 to D.L.), Science and Technology Commission of Shanghai Municipality (22ZR1468300 to D.L.), the Shanghai Medical Innovation Research Program (22Y21900900 to J.H.), Shanghai Municipal Education Commission China (24SG11 to J.H.), Shanghai Science and Technology Innovation Action Plan for Cell and Gene Therapy (24J22800500 to J.H.), Shanghai Science and Technology Innovation Action Plan for Advanced Materials (24CL2900800 to J.H.), and the China Postdoctoral Science Foundation (2022M720805 to Z.Z.).