This work was performed at the Key Laboratory of Agriculture, Beijing, China, and was supported by Natural Science Foundation of China (31872102), the Chinese Academy of Agricultural Sciences Innovation Project (CAAS-ASTIP-IVFCAAS), Beijing Scientific Program of Municipal Commission of Science and Technology (Z171100001517014), The National Key Research and Development Program of China (2018YFD0100805), and Key Laboratory of Biology and Genetic Improvement of Horticultural Crops, Ministry of Agriculture, Beijing, China.