国家自然科学基金31570063,31870020;山东省重点研发计划2017GSF17129;山东省重大科技创新工程2017CXGC0303;山东省农业科学院农业科技创新工程+CXGC2016B10Supported by the National Natural Science Foundation of China31570063,31870020;by the Shandong Key Research and Development Program2017GSF17129;by the Shandong Key Scientific and Technological Innovation Program2017CXGC0303;by the Agricultural Scientific and Technological Innovation Project of Shandong Academy of Agricultural Sciences+CXGC2016B10