1State Key Laboratory of Cell Biology, Shanghai Key Laboratory of Molecular Andrology, CAS Center for Excellence in Molecular Cell Science, Shanghai Institute of Biochemistry and Cell Biology, University of Chinese Academy of Sciences, Chinese Academy of Sciences, Shanghai 200031, China †These authors contributed equally to this work *Correspondence to:Shifeng Li , Email:sfli01@sibcb.ac.cn Jinsong Li , Email:jsli@sibcb.ac.cn Yiping Li , Email:yipingli@sibcb.ac.cn