SlWHY2 interacts with SlRECA2 to maintain mitochondrial function under drought stress in tomato.

Drought stress in plants leads to inhibition of photosynthesis and respiration, accumulation of reactive oxygen species (ROS), and reprogramming of gene expression. Here, we established that the tomato (Solanum lycopersicum) WHIRLY2 (SlWHY2) gene, which encodes a mitochondrial single-stranded DNA-binding protein, was significantly induced by drought stress. Under drought conditions, SlWHY2 RNAi plants showed more wilting and lower fresh weight, chlorophyll content, quantum yield of photosystem I (PSI; YI), and maximal photochemical efficiency of PSII (Fv/Fm) than the wild type (WT). Drought treatment also caused the SlWHY2 RNAi lines to accumulate more ROS than the WT, and the silenced lines had lower AOX (alternative oxidase) activity. As expected, the mitochondrial membrane potential (MMP) was less stable in the SlWHY2 RNAi lines. The expression levels of seven genes in the mitochondrial genome (SYCF15, NAD7, NAD4, COS2, COX1, COX2, and COX3) were decreased even more in the SlWHY2 RNAi lines than they were in the WT under drought stress. SlWHY2 interacted directly in vivo and in vitro with SlRECA2, a mitochondrial recombinase A that is important for mitochondrial DNA recombination and repair. These results suggest that SlWHY2 plays an essential role in maintaining mitochondrial function and enhancing drought tolerance in tomato.