supported by the National Key R&D Program of China(2022YFA1304000);the National Natural Science Foundation of China(81972823);the Foundation for Innovative Research Groups of the National Natural Science Foundation of China(32121004);the Guangdong Special Support Program innovation team(2019BT02N054);the Open Research Fund of State Key Laboratory of Cellular Stress Biology,Xiamen University(SKLCSB2019KF013);the Fundamental Research Funds for the Central University(20720220003)。