国家重点研发计划项目2016YFD0300602;国家自然科学基金项目61661136003, 41471285, 41271345;北京市农林科学院科技创新能力建设项目KJCX20170423资助. This study was supported by the National Key Research and Development Program of China2016YFD0300602;the National Natural Sci-ence Foundation of China61661136003, 41471285, 41271345;the Innovation Capacity Building Project of Beijing Academy of Agri-culture and Forestry SciencesKJCX20170423