Effect of divalent cations on ion fluxes and leaf photochemistry in salinized barley leaves

Photosynthetic characteristics, leaf ionic content, and net fluxes of Na⁺, K⁺, and Cl^- were studied in barley (Hordeum vulgare L) plants grown hydroponically at various Na/Ca ratios. Five weeks of moderate (50 mM) or high (100 mM) NaCl stress caused a significant decline in chlorophyll content, chlorophyll fluorescence characteristics, and stomatal conductance (g_s) in plant leaves grown at low calcium level. Supplemental Ca²⁺ enabled normal photochemical efficiency of PSII (F_v/F_m around 0.83), restored chlorophyll content to 80-90% of control, but had a much smaller (50% of control) effect on g _s. In experiments on excised leaves, not only Ca²⁺, but also other divalent cations (in particular, Ba²⁺ and Mg ²⁺), significantly ameliorated the otherwise toxic effect of NaCl on leaf photochemistry, thus attributing potential targets for such amelioration to leaf tissues. To study the underlying ionic mechanisms of this process, the MIFE technique was used to measure the kinetics of net Na⁺, K ⁺, and Cl^- fluxes from salinized barley leaf mesophyll in response to physiological concentrations of Ca²⁺,Ba²⁺, Mg²⁺, and Zn²⁺. Addition of 20 mM Na⁺ as NaCl or Na₂SO₄ to the bath caused significant uptake of Na ⁺ and efflux of K⁺. These effects were reversed by adding 1 mM divalent cations to the bath solution, with the relative efficiency Ba ²⁺>Zn²⁺=Ca²⁺>Mg²⁺. Effect of divalent cations on Na⁺ efflux was transient, while their application caused a prolonged shift towards K⁺ uptake. This suggests that, in addition to their known ability to block non-selective cation channels (NSCC) responsible for Na⁺ entry, divalent cations also control the activity or gating properties of K⁺ transporters at the mesophyll cell plasma membrane, thereby assisting in maintaining the high K/Na ratio required for optimal leaf photosynthesis.