本文由国家自然科学基金项目41301312,41371296,41571219、公益性行业农业科研专项201303126,201203030和国家重点研发计划项目2016YFD0200309-6,2016YFD0300806-1资助 This work was supported by the National Natural Science Foundation of China 41301312,41371296,41571219,the Special Fund for Agro-scientific Research in the Public Interest 201303126,201203030,and the National Key Research and Development Program of China 2016YFD0200309-6,2016YFD0300806-1.