国家自然科学基金项目31501713;国家重点研发计划项目2016YFD0201100;国家现代农业产业技术体系建设资金项目CARS-28;山东省自然科学基金项目ZR2015PC001资助 This work was supported by the National Natural Science Foundation of China31501713;the National Key Research and Development Program of China2016YFD0201100;the China Modem Agriculture Industry System Construction FoundationCARS-28;the Natural Science Foundation of Shandong Province,ChinaZR2015PC001