Rare-earth element scandium improves stomatal regulation and enhances salt and drought stress tolerance by up-regulating antioxidant responses of Oryza sativa

Oryza sativa L. cv. Gönen grown in hydroponic culture was treated with scandium (Sc; 25 and 50 μM) alone or in combination with salt (100 mM NaCl) and/or drought (5% PEG-6000). Stress caused a decrease in growth (RGR), water content (RWC), osmotic potential (ΨΠ), chlorophyll fluorescence (Fv/Fm) and potential photochemical efficiency (Fv/Fo). Sc application prevented the decreases of these parameters. Sc also alleviated the changes on gas exchange parameters (carbon assimilation rate (A), stomatal conductance (gs), intercellular CO2 concentrations (Ci), transpiration rate (E) and stomatal limitation (Ls)). Stress caused no increase in superoxide dismutase (SOD) activity. After induvial applied NaCl or PEG, catalase (CAT) and ascorbate peroxidase (APX) showed an enhancement in activation and tried to scavenge of hydrogen peroxide (H2O2). On the other hand, in plants with the combination form of NaCl and PEG, only CAT activity was induced. Sc applications to NaCl-treated rice led to an increase of SOD, APX, glutathione reductase (GR), monodehydroascorbate reductase (MDHAR) and dehydroascorbate reductase (DHAR) as well as peroxidase (POX). Sc under NaCl could be maintained both ascorbate (AsA) and glutathione (GSH) regeneration. Despite of induction of MDHAR and DHAR under Sc plus PEG, Sc did not maintain AsA redox state because of no induction in APX activity. However, GSH pool could be regenerated by induction in DHAR and GR in this group. Sc application (especially for 25 μM) in rice exposed to NaCl + PEG resulted an enhancement in APX and MDHAR and so Sc could be partially provided AsA regeneration. Since no increases in DHAR and GR were observed, GSH pool was reduced. Due to this activation of antioxidant enzymes, stress-induced H2O2 and TBARS content (lipid peroxidation) significantly decreased in rice with Sc applications. Sc in plants with stress also increased the transcript levels of OsCDPK7 and OsBG1 related to stomatal movement and signaling pathway. Consequently, Sc protected the rice plants by minimizing disturbances caused by NaCl or PEG exposure via the AsA-GSH redox-based systems.