1Unit of Respiratory Infection and Immunity, Institut Pasteur of Shanghai, Chinese Academy of Sciences, Shanghai 200031, China 2CAS Key Laboratory of Molecular Virology and Immunology, Institut Pasteur of Shanghai, Chinese Academy of Sciences, Shanghai 200031, China 3University of Chinese Academy of Sciences, Beijing 100049, China 4State Key Laboratory of Respiratory Disease, National Clinical Research Centre for Respiratory Disease, Guangzhou Institute of Respiratory Health, The First Affiliated Hospital of Guangzhou Medical University, Guangzhou 510120, China †These authors contributed equally to this work. *Correspondence to:Jincun Zhao , Email:zhaojincun@gird.cn Xiao Su , Email:xsu@ips.ac.cn