1MOE Key Laboratory for Cellular Dynamics, Hefei National Research Center for Cross-disciplinary Sciences at the Microscale, University of Science & Technology of China School of Life Sciences, Hefei 230027, China 2Frontier Science Center for Immunology and Metabolism, Medical Research Institute, Wuhan University, Wuhan 430071, China 3School of Basic Medical Sciences, Tianjin Medical University, Tianjin 300070, China †These authors contributed equally to this work *Correspondence to:Xing Liu , Email:xing1017@ustc.edu.cn Xiaoyu Song , Email:sonxiaoyu@ustc.edu.cn