This research was supported by the Hunan Provincial Department of Education Outstanding Youth Project (18B176), the 13th Five-Year National Key R & D Program Project (2016YFD0600102), the Hunan Key R & D Program Project (2016NK2143), the Hunan University Students Innovation and Entrepreneurship Training Project (S202010538068), and the Project of Hunan Provincial Key Laboratory of Crop Germplasm Innovation and Utilization (Grant No.19KFXM13).