This work was funded by the National Natural Science Foundation of China (Grant nos. 31101596 and 31372130), Talent Scholar of Hunan Agricultural University (11YJ21) P. R. China. FuRong Scholar Program of Hunan Province, P. R. China, The “Twelfth Five-Year” National Science and technology support program (2012BAD15BO4). Open project of National Key Laboratory of Plant Molecular Genetics. Open novel science foundation of Hunan province (13K062). We thank Tanya Streeter and Juanita Johns from the University of West Florida, United States of America for proofreading drafts of this manuscript.