(1. Hubei Key Laboratory of Food Crop Germplasms and Genetic Improvement, Institute of Food Crops, Hubei Academy of Agricultural Sciences, Wuhan 430064, China; 2. Hubei Hongshan Laboratory, Wuhan 430070, China; 3. College of Life Sciences, Wuhan University, Wuhan 430072, China; 4. Key Laboratory of Crop Molecular Breeding, Ministry of Agriculture and Rural Affairs, Wuhan 430072, China)