基金This work was supported by the National Natural Science Foundation of China(31730110,31661143031,91940303,and 91753135)the Science and Technology Commission of Shanghai Municipality grant(17JC1404900,18XD1404400,and 20ZR1467300)a Joint Research grant with State Key Laboratory of Microbial Metabolism,School of Life Science and Biotechnology,Shanghai Jiao Tong University(MMLKF16-11).