Genome-wide identification and expression profile analysis of SWEET genes in Chinese jujube

The novel sugar transporter known as SWEET (sugars will eventually be exported transporter) is involved in the transport and distribution of photosynthesis products in plants. The SWEET protein is also involved in pollen development, nectar secretion, stress responses, and other important physiological processes. Although SWEET genes have been characterized and identified in model plants, such as Arabidopsis and rice, little is known about them in jujube. In this study, the molecular characteristics of the SWEET gene family in the Chinese jujube (Ziziphus jujuba Mill.) and their expression patterns in different organs, at different fruit developmental stages, and under abiotic stress were analyzed. A total of 19 ZjSWEET genes were identified in jujube through a genome-wide study; these were classified into four sub-groups based on their phylogenic relationships. The gene structure analysis of ZjSWEET genes showed that all the members had introns. The expression patterns of different ZjSWEET genes varied significantly in different organs (root, shoot, leave, flower, fruit), which indicated that ZjSWEETs play different roles in multiple organs. According to the expression profiles by quantitative real-time PCR analysis during fruit development, the expression levels of the two genes (ZjSWEET11, ZjSWEET18) gradually increased with the development of the fruit and reached a high level at the full-red fruit stage. A prediction of the cis-acting regulatory elements indicated that the promoter sequences of ZjSWEETs contained nine types of phytohormone-responsive cis-regulatory elements and six environmental factors. In addition, the expression profiles by quantitative real-time PCR analysis showed that some of the ZjSWEETs responded to environmental changes; ZjSWEET2 was highly induced in response to cold stress, and ZjSWEET8 was significantly up-regulated in response to alkali and salt stresses. This study showed that the functions of the ZjSWEET family members of jujube are different, and some may play an important role in sugar accumulation and abiotic stress in jujube.