supported by the National Natural Science Foundation of China(32088102,31730103,31825003,32050081,and 31870218);the CAS Project for Young Scientists in Basic Research(YSBR-011);the Strategic Priority Research Program “Molecular Mechanism of Plant Growth and Development”of the Chinese Academy of Sciences(XDB27040207);the National Key R&D Program of China(2019YFA0904703 and 2016YFA0500502);the Young Elite Scientists Sponsorship Program by CAST(20202022QNRC001/2/3)。