This study aims to explain the effects of silicon (Si) foliar application on gas exchange characteristics, photosynthetic pigments, membrane stability and leaf relative water content of different wheat cultivars in the field under drought stress conditions. The experiment was arranged as a split-split plot based on randomized complete block design with three replications. Irrigation regime (100%, 60%, and 40% F.C.), silicon (control and Si application) and wheat cultivars (Shiraz, Marvdasht, Chamran, and Sirvan) were considered as main, sub and sub-sub plots, respectively. This study was carried out at the Research Farm of the Collage of Agriculture, Shiraz University, Iran, during 2012–2013 growing season. The results showed that foliar application of silicon increased the leaf relative water content, photosynthesis pigments (chlorophyll a, b and total chl and carotenoids), chlorophyll stability index (CSI) and membrane stability index (MSI) in all wheat cultivars, especially in Sirvan and Chamran (drought tolerant cultivars), under both stress and non-stress conditions. However, more improvement was observed under drought stress as compared to the non-stress condition. In contrast, these parameters decreased under drought stress. Si significantly decreased electrolyte leakage in all four cultivars under drought stress conditions. Furthermore, the intercellular carbon dioxide (CO₂) concentration (C_i) increased under drought stress. Si application decreased C_i especially under drought stress conditions. Net photosynthesis rate (A), transpiration rate (E) and stomatal conductance (g_s) were significantly decreased under drought conditions. Under drought, Si applied plants showed significantly higher leaf photosynthesis rate, transpiration rate, and stomatal conductance. Intrinsic water use efficiency (WUEi) and carboxylation efficiency (C_E) decreased in all cultivars under drought stress. However, the silicon-applied plants had greater WUEi and C_E under drought stress. The stomatal limitation was found to be higher in stressed plants compared to the control. Exogenously applied silicon also decreased stomatal limitation. Overall, application of Si was found beneficial for improving drought tolerance of wheat plants.

Environmental stresses present major challenges in our quest to achieve sustainable food production. The reactions of plants to environmental stresses are complex and involve many kinds of physiological and biochemical responses. Stress causes multifarious adverse effects in plants. Production of a family of reactive oxygen species (ROS) is a common phenomenon. When plants are subjected to environmental stress, the balance between the production of ROS and the quenching activity of antioxidants is upset, often resulting in an oxidative damage. Plants with high levels of antioxidant enzyme activity are reported to have greater resistance to this oxidative damage. The activities of component enzymes or the antioxidant levels are usually only double in response to many stress situations. This rather moderate response might be understood if we consider that the system is geared to self-destruction when it comes under threat. Understanding the mechanisms by which plants perceive environmental signals and transmit the signals to cellular machinery to activate adaptive responses is of fundamental importance to biology. The present review is focusing on ROS generation and plant defenses to them.

The effects of silicon (Si) and salicylic acid (SA) applications on proline content and expression of Δ¹-pyrrolin-5-carboxylate synthetase (P5CS) were examined under different drought levels and different drought exposure times. Two wheat cultivars, a drought tolerant and a drought sensitive were used. The experiment was a factorial based on completely randomized design with three replicates. Expression analysis by the quantitative real time PCR showed that the tolerant cultivar had significantly higher P5CS expressions compared to the sensitive one under drought stress. In sampling time points, the maximum level of mRNA was observed at 48 h after stress was applied. At 48 h after stress induction, the expression of P5CS was almost 3.1 fold higher in the tolerant cultivar compared to the sensitive one. In both cultivars, gene expression decreased from 48 to 72 h. The stressed plants treated with Si + SA showed a higher expression. Proline content started to increase by Si and SA treatments and the maximum proline content was obtained at simultaneous application of Si + SA. Drought stress significantly reduced chlorophyll content, relative water content and leaf water potential of both cultivars, while increased electrolyte leakage (EL) of the leaves. In contrast, foliar-applied Si and SA significantly increased these parameters and reduced EL, and the effect of simultaneous application of Si and SA was greater. The results suggest that the P5CS is a stress inducible gene. This gene has the potential to be used for improvement of drought stress tolerance in wheat. Network analysis highlighted positive interaction of osmotic stress, drought and cold stress on P5CS1 and the regulatory role of MYB2, ERF-1, and EIN3 transcription factors. In conclusion, alleviation of drought stress by application of Si and SA was associated partially with enhanced expression of P5CS gene and following proline accumulation.