The salt tolerance locus SOS1 from Arabidopsis has been shown to encode a putative plasma membrane Na ؉ /H ؉ antiporter. In this study, we examined the tissue-specific pattern of gene expression as well as the Na ؉ transport activity and subcellular localization of SOS1. When expressed in a yeast mutant deficient in endogenous Na ϩ transporters, SOS1 was able to reduce Na ؉ accumulation and improve salt tolerance of the mutant cells. Confocal imaging of a SOS1-green fluorescent protein fusion protein in transgenic Arabidopsis plants indicated that SOS1 is localized in the plasma membrane. Analysis of SOS1 promoter- -glucuronidase transgenic Arabidopsis plants revealed preferential expression of SOS1 in epidermal cells at the root tip and in parenchyma cells at the xylem/symplast boundary of roots, stems, and leaves. Under mild salt stress (25 mM NaCl), sos1 mutant shoot accumulated less Na ؉ than did the wildtype shoot. However, under severe salt stress (100 mM NaCl), sos1 mutant plants accumulated more Na ؉ than did the wild type. There also was greater Na ؉ content in the xylem sap of sos1 mutant plants exposed to 100 mM NaCl. These results suggest that SOS1 is critical for controlling long-distance Na ؉ transport from root to shoot. We present a model in which SOS1 functions in retrieving Na ؉ from the xylem stream under severe salt stress, whereas under mild salt stress it may function in loading Na ؉ into the xylem.
INTRODUCTIONPlant growth depends on mineral nutrients absorbed from the soil by roots. Although Na ϩ is a major cation present in soil solutions, Na ϩ is not considered an essential mineral for most plants. In saline soils, high concentrations of Na ϩ disrupt K ϩ and other mineral nutrition, create hyperosmotic stress, and cause secondary problems such as oxidative stress (Zhu, 2001). These adverse effects contribute to plant growth inhibition and even plant death.Many cytosolic enzymes are activated by K ϩ and inhibited by Na ϩ (Flowers et al., 1977). Three mechanisms are available to plant cells to prevent excessive accumulation of Na ϩ in the cytosol (Niu et al., 1995;Blumwald et al., 2000; Zhu, 2001). First, Na ϩ entry to plant cells may be restricted by selective ion uptake. Nonselective cation channels have been proposed to mediate substantial Na ϩ entry into plant roots, but genes encoding these channels have yet to be identified (Amtmann and Sanders, 1999; Tyerman and Skerrett, 1999). The cloned transporters HKT1 and LCT1 have Na ϩ permeability when expressed in yeast or oocytes, suggesting that they also may be candidate Na ϩ influx transporters (Rubio et al., 1995;Schachtman et al., 1997). Recently, studies in yeast demonstrated that the magnitude of the membrane potential affects net Na ϩ influx into cells. Mutations in the yeast PMP3 gene lead to membrane hyperpolarization, increased Na ϩ influx, and salt sensitivity (Navarre and Goffeau, 2000).Second, internalized Na ϩ can be stored in vacuoles. Vacuolar compartmentation is an efficient strategy for plant cells to deal with salt s...