We are grateful to Lei Bai, Luyang Tian, Ruijun Bai, Jing Lu, Ying Yang, Xiuping Li, Shikun Ma and Lei Zhang for administrative assistance, to Wang Kang (BIG, CAS) for his help in RNA-seq analysis, to Wei Li, Jingyi Jia (Xuanwu Hospital Capital Medical University) and Lukai Huang (IOZ, CAS) for their help in the management of laboratory animals, and to Xixia Li and Xueke Tan (IBP, CAS) for their help in transmission electron microscopy. This work was supported by the National Key Research and Development Program of China (2018YFA0107203), the Strategic Priority Research Program of the Chinese Academy of Sciences (XDA16010000), the National Key Research and Development Program of China (2020YFA0804000, 2018YFC2000100, 2020YFA0112201, 2017YFA0103304, 2017YFA0102802, 2020YFA0113400, 2019YFA0110100), the National Natural Science Foundation of China (Grant Nos. 81901433, 81921006, 81625009, 91749202, 81861168034, 91949209, 92049304, 81822018, 92049116, 82071588, 32000500, 81922027, 81870228, 82125011, 82122024, 32100937, 92149301, 92168201), the Key Research Program of the Chinese Academy of Sciences (KFZD-SW-221), the Program of Beijing Municipal Science and Technology Commission (Z191100001519005), the Program of the Beijing Natural Science Foundation (Z190019, JQ20031), K. C. Wong Education Foundation (GJTD-2019-06, GJTD-2019-08), Beijing Hospitals Authority Youth Programme (QML20200802), Youth Innovation Promotion Association of CAS (2021078, E1CAZW0401), the 14th Five-year Network Security and Informatization Plan of Chinese Academy of Sciences (WX145XQ07-18), the Non-profit Central Research Institute Fund of Chinese Academy of Medical Sciences (2020-JKCS-011), the State Key Laboratory of Stem Cell and Reproductive Biology, the State Key Laboratory of Membrane Biology, and the Milky Way Research Foundation (MWRF).

Qun Chu, Feifei Liu, Yifang He, Xiaoyu Jiang, Yusheng Cai, Zeming Wu, Kaowen Yan, Lingling Geng, Yichen Zhang, Huyi Feng, Kaixin Zhou, Si Wang, Weiqi Zhang, Guang-Hui Liu, Shuai Ma, Jing Qu and Moshi Song declared no conflict of interest. All institutional and national guidelines for the care and use of laboratory animals were followed.