国家自然科学基金项目31271660;国家公益性行业农业科研专项201503130;山东省农业重大应用技术创新项目2014-2016资助。This work was supported by the National Natural Science Foundation of China31271660;the Special Fund for Agro-scientific Research in the Public Interest of China201503130;the Key Innovation Project on Agriculture Applied Technology from Shandong Provincial Government2014-2016