国家重点基础研究发展计划973计划项目2014CB138200;北京市科技计划项目D141100005014003;中国农业科学院科技创新工程项目资助。 This study was supported by the National Basic Research Program of China2014CB138200;the Program of Beijing Municipal Science and TechnologyD141100005014003;the Agricultural Science and Technology Innovation Program ASTIP of CAAS