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Plants 2022, 11, 2379 11 of 18 that a strong negative correlation exists. The representation of RMDA showed that it had a little contribution in both PCAs (Figure 6A,B). 3. Discussion Salinity stress seriously hampers plant’s growth and development through a reduc- tion in the photosynthetic process, upsetting ionic equilibrium and enhancing oxidative damage [31]. The impact of salinity on a plant’s growth is a time dependent process, summarized in a two-phase model: (i) rapid phase ascribed to water deficit conditions (os- motic); and (ii) slow phase caused by the accumulation of ions to toxic levels (ion-specific) causing negative effects on plant’s physio-biochemical activities [32]. Salinity-induced reduction in biomass accumulation and plant height was observed in this experiment, however, this effect was more pronounced on salt sensitive barley genotype (B-14011) as compared to a salt tolerant one (B-10008). The decrease in biomass and lengths of plants was mainly the result of an excessive buildup of Na+ and Cl− [4] due to a disturbance in nutrient uptake [32] and a reduction in te photosynthetic process, which is directly related to biomass accumulation [33]. Salinity-induced reduction in morphological attributes has been previously observed in maize [34], wheat [35], canola [36], and barley [37]. The application of Si through rooting media showed a positive effect on growth of barley plants. Si-mediated increase in biomass and plant height under salinity is attributed to improved nutrient balance, reduced uptake of Na+, higher photosynthetic rates and efficient detoxification of excessive ROS via enhanced antioxidant activity [18,34]. The Si-mediated enhancement in growth under salinity stress was previously reported by Yan et al. [38] in wheat, Ahmad et al. [12] in mung bean, Laifa et al. [39] in barley and Raza et al. [34] in maize. The over-accumulation of H2O2 in cellular system under salinity results in fatty acid oxidation leading to membrane damage and electrolyte leakage, as was observed as enhanced MDA levels under salt stress, especially in B-14011. This salinity-induced production of ROS (oxidative stress) harms lipids, proteins, carbohydrates and nucleic acid leady to cell death [40]. On the other hand, Si application through external means in salt treated plants ameliorated extra ROS (H2O2) and caused reduction in lipid peroxidation (MDA). The main source of ROS is mitochondria and chloroplasts where it is accumulated during electron transport processes [41], which can start lipid peroxidation in the cell [42]. AbdElgawad et al. [43] reported that 150 mM NaCl enhanced the production of H2O2 and enhanced lipid peroxidation (higher MDA level) in maize. The Si application tends to maintain the metabolism of plant to an optimum level by decreasing ROS production, lowering lipid peroxidation, maintaining integrity of membranes and reducing leakage of electrolyte from cytosol in many crops like wheat [29], rice [38], Basil [24] and sunflower [44] and maize [30]. The immediate response of plants to the overproduction of ROS is enhancement in the synthesis of TSPs, TFAAs. Similarly, the activation of enzymatic and non-enzymatic antiox- idants is also enhanced in order to efficiently scavenge excessive ROS [12,43]. As compared to B-14011 the genotype B-10008 synthesized higher TSP contents under salt stress, while the accumulation of TFAA was higher in B-14011 than B-10008. However, imposition of Si augmented TSP contents in leaves and roots of barley plants. This enhancement in TSP accumulation indicate that plant’s endogenous defense system was boosted under salinity stress [45,46]. The enzymatic antioxidant (CAT, POD, SOD, APX, and GR) activities were boosted under salinity stress. The accumulation of CAT and POD was higher in B-14011, and displaying that increase in H2O2 levels enhanced these antioxidants. The salinity induced enhancement in the activities of enzymatic antioxidant, i.e., CAT, POD, SOD, APX and GR is reported in maize [30], sunflower [44], wheat [29] and alfalfa [47]. The stimulation in the synthesis of non-enzymatic antioxidants (AsA, α-tocopherol, total phenolics, glutathione and proline) in different plant tissues of barley genotypes was significantly enhanced underPDF Image | Silicon-Induced Mitigation of NaCl Stress in Barley
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