国家自然科学基金项目31071358,30871476;国家"十二五"科技支撑计划项目2013BAD07B06-2;国家现代农业产业技术体系建设专项CARS-02-20;国家公益性行业农业科研专项HY20121203100,HY1203096;山东省财政支持农业重大应用技术创新课题2014资助. This study was supported by the National Natural Science Foundation of China31071358, 30871476;the National "12th Five-Year" Science and Technology Support Plan2013BAD07B06-2;the China Agriculture Research SystemCARS-02-20;the Special Fund for Agro-scien- tific Research in the Public InterestHY20121203100, HY1203096;the Funds for Financial Support to Agriculture in Shandong Pro-vince2014