Halophyte species implement a "salt-including" strategy, sequestering significant amounts of Na + to cell vacuoles. This requires a reduction of passive Na + leak from the vacuole. In this work, we used quinoa (Chenopodium quinoa) to investigate the ability of halophytes to regulate Na + -permeable slow-activating (SV) and fast-activating (FV) tonoplast channels, linking it with Na + accumulation in mesophyll cells and salt bladders as well as leaf photosynthetic efficiency under salt stress. Our data indicate that young leaves rely on Na + exclusion to salt bladders, whereas old ones, possessing far fewer salt bladders, depend almost exclusively on Na + sequestration to mesophyll vacuoles. Moreover, although old leaves accumulate more Na + , this does not compromise their leaf photochemistry. FV and SV channels are slightly more permeable for K + than for Na + , and vacuoles in young leaves express less FV current and with a density unchanged in plants subjected to high (400 mM NaCl) salinity. In old leaves, with an intrinsically lower density of the FV current, FV channel density decreases about 2-fold in plants grown under high salinity. In contrast, intrinsic activity of SV channels in vacuoles from young leaves is unchanged under salt stress. In vacuoles of old leaves, however, it is 2-and 7-fold lower in older compared with young leaves in control-and saltgrown plants, respectively. We conclude that the negative control of SV and FV tonoplast channel activity in old leaves reduces Na + leak, thus enabling efficient sequestration of Na + to their vacuoles. This enables optimal photosynthetic performance, conferring salinity tolerance in quinoa species.The increasing problem of global land salinization (Flowers, 2004;Rengasamy, 2006) and its associated multibillion dollar losses in agricultural production require a better understanding of the key physiological mechanisms that confer salinity tolerance in crops. One effective way of gaining such knowledge comes from studying halophytes (Glenn et al., 1999;Flowers and Colmer, 2008;Shabala and Mackay, 2011).One of the prominent features of halophytes is their ability to efficiently sequester cytosolically toxic Na + to the cell vacuole. The classic view is that this sequestration is achieved by tonoplast Na + /H + antiporters (Barkla et al., 1995;Flowers and Colmer, 2008), a process energized by both vacuolar H + pumps: ATPase (Ayala et al., 1996;Vera-Estrella et al., 1999;Wang et al., 2001) and pyrophosphatase (Parks et al., 2002;Vera-Estrella et al., 2005;Guo et al., 2006;Krebs et al., 2010). However, recent studies have added more complexity to the relationship between Na + /H + antiporters and vacuolar Na + sequestration, assigning a role to the transporter in the regulation of K + and H + homeostasis (for review, see Rodríguez-Rosales et al., 2009;Jiang et al., 2010; Bassil et al., 2011 (Yamaguchi et al., 2005). Consequently, other transporters, in addition to and different from NHX, are likely to be involved in vacuolar Na + sequestration. In addition...