We are grateful to Lei Bai, Ruijun Bai, Qun Chu, Shikun Ma, Jing Lu, Luyang Tian and Ying Yang for their administrative assistance. We thank Mengyi Sun, Jiayin Wang from Prof. Keqiong Ye’s lab (IBP, CAS) for their technical supports on RNA electrophoresis, Xueying Wang (Tsinghua University), Jifeng Wang (IBP, CAS), Zhensheng Xie (IBP, CAS) for their help in LC-MS/MS assays, Junying Jia (IBP, CAS), Shu Meng (IBP, CAS) and Shuang Sun (IBP, CAS) for their assistance in flow cytometry analyses. We also thank Chuqian Liang and Wang Kang for their technical supports. This work was supported by the National Key Research and Development Program of China (2019YFA0110100), the Strategic Priority Research Program of the Chinese Academy of Sciences (XDA16010000), the National Natural Science Foundation of China (Grant Nos. 31900524, 31970597, 81921006, 81625009, 91749202, 81861168034, 91949209, 92049304, 82125011, 81822018, 92049116, 82071588, 82122024, 32100937, 92149301, and 92168201), the National Key Research and Development Program of China (2018YFC2000100, 2020YFA0804000, 2017YFA0103300, 2017YFA0102800, 2018YFA0107200, 2019YFA0110900, 2020YFA0112200, 2020YFA0803401, and 2019YFA0802202), Beijing Natural Science Foundation (Z190019), the Key Research Program of the Chinese Academy of Sciences (KFZD-SW-221), the 14th Five-year Network Security and Informatization Plan of Chinese Academy of Sciences (WX145XQ07-18), K. C. Wong Education Foundation (GJTD-2019-06, GJTD-2019-08), Youth Innovation Promotion Association of CAS (E1CAZW0401), CAS Special Research Assistant (SRA) Program, the State Key Laboratory of Membrane Biology, the State Key Laboratory of Stem Cell and Reproductive Biology, the State Key Laboratory of Brain and Cognitive Science, the Tencent Foundation (2021-1045), and the Milky Way Research Foundation (MWRF).

Sheng Zhang, Zeming Wu, Yue Shi, Si Wang, Jie Ren, Zihui Yu, Daoyuan Huang, Kaowen Yan, Yifang He, Xiaoqian Liu, Qianzhao Ji, Beibei Liu, Zunpeng Liu, Jing Qu, Guang-Hui Liu, Weimin Ci, Xiaoqun Wang and Weiqi Zhang declared no conflict of interest.