This work was financially supported by China National Natural Science Foundation (31571608), Natural Science Fund for Colleges and Universities in Jiangsu Province (15KJA210003), Jiangsu Provincial Natural Science Foundation (BK20151311), Jiangsu Provincial Agriculture Innovation Program (CX(16)1001), Jiangsu Provincial Fund for Application of Research Achievement (BA2016173) and the open funds of the Key Laboratory of Plant Functional Genomics of the Ministry of education (ML201903).