本文由国家重点基础研究发展计划项目2015CB453303、公益性行业农业科研专项201303050和山东省“泰山学者”工程专项2008-67资助;This work was supported by the National Basic Research Program of China 2015CB453303,the Special Fund for Agro-scientific Research in the Public Interest 201303050,and the Taishan Scholar Program of Shandong Province 2008-67.