国家重点研发计划项目七大作物育种试点专项2016YFD101802;北京市自然科学基金重点项目6151002;中国农业科学院科技创新工程项目的资助。 This study was supported by the National Key R&D Program of China Pilot Project on Seven Main Crop Breeding2016YFD101802;the Natural Science Foundation of Beijing Key Program6151002;the Agricultural Science and Technology Innovation Program of CAAS