This study was financially supported by the National Natural Science Foundation of China (31801923, 31101596 and 31372130), the National Key Research and Development Program of China (2017YFD0200100 and 2017YFD0200103), the Hunan Provincial Recruitment Program of Foreign Experts, National Oilseed Rape Production Technology System of China,‘2011 Plan’ supported by the Ministry of Education of China, and Double First-class Construction Project of Hunan Agricultural University (kxk201801005).