国家自然科学基金项目31271635;山东省自主创新重大关键技术计划项目2014GJJS0201-1;国家现代农业产业技术体系建设专项CARS-03-1-8;山东省农业科学院青年英才培养计划资助。This study was supported by the National Natural Science Founding of China31271635;Shandong Provincial Innovation Project for Key Technologies2014GJJS0201-1;China Agriculture Research SystemCARS-03-1-8;the Yong Talents Training Program of Shandong Academy of Agricultural Sciences