国家自然科学基金项目31502191;公益性行业农业科研专项201503108;现代农业产业技术体系建设专项CARS-49;中央级公益性科研院所基本科研业务费专项资金项目2016C009共同资助[This work was supported by the National Natural Science Foundation of China31502191;the Special Fund for Agro-scientific Research in the Public Interest201503108;Modern Agricultural Industry Technology SystemCARS-49;the Special Fund of Chinese Central Government for Basic Scientific Research Operations in Commonweal Research Institutes 2016C009]