Plasma Membrane-Localized Transporter MdSWEET12 Is Involved in Sucrose Unloading in Apple Fruit

Sugar content is an important factor determining the flavor in apple fruit. Sugar unloading is a prerequisite step for sugar accumulation. However, little is known about sugar unloading mechanisms in apple. Transcriptomic sequencing of two apple varieties, "Envy" and "Pacific Rose," with significantly different sugar content was performed. MdSWEET12a from the SWEET transporter family was differentially expressed. Further study of the MdSWEET12a showed that this plasma membrane-localized transporter protein-encoding gene was mainly expressed in sieve element-companion cells (SE-CC) in the fruit, which was positively correlated with the sucrose accumulation during the development of "Envy" apple. Consistently manipulating the gene expression through either transient overexpression or silencing significantly increased or decreased the sugar content in apple fruit, respectively. Complementary growth experiments in mutant yeast cells indicated that MdSWEET12a transported sucrose. Heterologous expression of MdSWEET12a in tomato increased the expression of genes related to sugar metabolism and transport, leading to increased sugar content. These findings underpin the involvement of MdSWEET12a in sugar unloading in apple fruit.