Membrane fluxes, bypass flows, and sodium stress in rice: the influence of silicon.

Provision of silicon (Si) to roots of rice (Oryza sativa L.) can alleviate salt stress by blocking apoplastic, transpirational bypass flow of Na+ from root to shoot. However, little is known about how Si affects Na+ fluxes across cell membranes. Here, we measured radiotracer fluxes of 24Na+, plasma membrane depolarization, tissue ion accumulation, and transpirational bypass flow, to examine the influence of Si on Na+ transport patterns in hydroponically grown, salt-sensitive (cv. IR29) and salt-tolerant (cv. Pokkali) rice. Si increased growth and lowered [Na+] in shoots of both cultivars, with minor effects in roots; neither root nor shoot [K+] were affected. In IR29, Si lowered shoot [Na+] via a large reduction in bypass flow, while in Pokkali, where bypass flow was small and not affected by Si, this was achieved mainly via a growth dilution of shoot Na+. Si had no effect on unidirectional 24Na+ fluxes (influx and efflux), or on Na+-stimulated plasma-membrane depolarization, in either IR29 or Pokkali. We conclude that, while Si can reduce Na+ translocation via bypass flow in some (but not all) rice cultivars, it does not affect unidirectional Na+ transport or Na+ cycling in roots, either across root cell membranes or within the bulk root apoplast.

Measurement of Differential Na(+) Efflux from Apical and Bulk Root Zones of Intact Barley and Arabidopsis Plants.

Rapid sodium cycling across the plasma membrane of root cells is widely thought to be associated with Na(+) toxicity in plants. However, the efflux component of this cycling is not well understood. Efflux of Na(+) from root cells is believed to be mediated by Salt Overly-Sensitive-1, although expression of this Na(+)/H(+) antiporter has been localized to the vascular tissue and root meristem. Here, we used a chambered cuvette system in which the distal root of intact salinized barley and Arabidopsis thaliana plants (wild-type and sos1) were isolated from the bulk of the root by a silicone-acrylic barrier, so that we could compare patterns of (24)Na(+) efflux in these two regions of root. In barley, steady-state release of (24)Na(+) was about four times higher from the distal root than from the bulk roots. In the distal root, (24)Na(+) release was pronouncedly decreased by elevated pH (9.2), while the bulk-root release was not significantly affected. In A. thaliana, tracer efflux was about three times higher from the wild-type distal root than from the wild-type bulk root and also three to four times higher than both distal- and bulk-root fluxes of Atsos1 mutants. Elevated pH also greatly reduced the efflux from wild-type roots. These findings support a significant role of SOS1-mediated Na(+) efflux in the distal root, but not in the bulk root.