Genome-wide identification and expression analysis of Na+/H+antiporter (NHX) genes in tomato under salt stress.

Plant Na ⁺/H ⁺ antiporter (NHX) genes enhance salt tolerance by preventing excessive Na⁺ accumulation in the cytosol through partitioning of Na⁺ ions into vacuoles or extracellular transport across the plasma membrane. However, there is limited detailed information regarding the salt stress responsive SlNHXs in the most recent tomato genome. We investigated the role of this gene family's expression patterns in the open flower tissues under salt shock in Solanum lycopersicum using a genome-wide approach. A total of seven putative SlNHX genes located on chromosomes 1, 4, 6, and 10 were identified, but no ortholog of the NHX5 gene was identified in the tomato genome. Phylogenetic analysis revealed that these genes are divided into three different groups. SlNHX proteins with 10-12 transmembrane domains were hypothetically localized in vacuoles or cell membranes. Promoter analysis revealed that SlNHX6 and SlNHX8 are involved with the stress-related MeJA hormone in response to salt stress signaling. The structural motif analysis of SlNHX1, -2, -3, -4, and -6 proteins showed that they have highly conserved amiloride binding sites. The protein-protein network revealed that SlNHX7 and SlNHX8 interact physically with Salt Overly Sensitive (SOS) pathway proteins. Transcriptome analysis demonstrated that the SlNHX2 and SlNHX6 genes were substantially expressed in the open flower tissues. Moreover, quantitative PCR analysis indicated that all SlNHX genes, particularly SlNHX6 and SlNHX8, are significantly upregulated by salt shock in the open flower tissues. Our results provide an updated framework for future genetic research and development of breeding strategies against salt stress in the tomato.