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Plants 2022, 11, 2379 12 of 18 salinity stress. Comparatively, the non-enzymatic antioxidant activities were higher in B- 10008 as compared to B-14011, when compared to their respective controls under salt stress. It has been observed that many plants enhance the activities of non-enzymatic antioxidants to safeguard the cellular structures from ROS-induced oxidative damage under salinity stress [24,48,49]. The application of Si through external means boosted the salinity-induced scavenging of ROS through antioxidant defense system in both barley genotypes. The possible mecha- nism of Si-induced boosted defense system under salinity stress is due to reduction in Na+ uptake; increased K+ absorption, improved water status, enhanced water retention capacity and finally limiting ROS production [50]. The similar results were previously reported in millet [51], maize [52], alfalfa [53] and wheat [54]. This study also uncovered the fact that Si application enhanced the activities of enzymatic (CAT, POD, SOD, APX and GR) and non-enzymatic (AsA, α-tocopherol, total phenolics, glutathione and proline) antioxidants under salt stress. The sensitive barley genotype (B-14011) showed a comparatively better antioxidant response analogous to a tolerant genotype (B-10008) under salinity stress. The Si-mediated enhancement in antioxidant activities reduced the oxidative damages posed by ROS, thus lowering lipid peroxidation and conserving membrane permeabil- ity [31,55]. Although results clearly demonstrate that there is a clear difference in antioxi- dant response in Si-treated and non-Si treated barley plants, a gap still exists in clarifying the interaction of exogenously applied Si and the antioxidant battery system of plants. Under salinity stress, Si application through external means reduced the uptake of Na+ by stimulating the root plasma membrane H+-ATPase activity, which can possibly lower ROS thus enhancing salt tolerance aided by efficient antioxidant defense system [56,57]. Enhanced proline accumulation under stressful conditions is regarded as a defense response of plants to a specific stress [58]. In this experiment the proline contents was considerably enhanced in both barley genotypes, however the response of B-14011 was exponentially high as compared to B-10008 genotypes, whereas presence of Si in media further boosted proline synthesis under salinity stress. It has been generally accepted that proline accumulation is a stress adoptive strategy in tolerant plant species [59]. Yet, proline over-accumulation under salt stress cannot be regarded as a permanent bench-mark for salt tolerance as there are many reports that concluded that sensitive genotypes/varieties had accumulated much higher proline when compared to tolerant ones [60,61]. This proline over accumulation in sensitive genotypes is an indication of salt injury [62]. Results of this experiment showed that Si application lowered proline contents of both barley genotypes. Previously, Tuna et al. [63] in wheat, Soylemezoglu et al. [64] in grape, Yin et al. [65] in sorghum and Gunes et al. [66] in barley have reported similar results showing that Si application can reduce the proline contents under salt stress. 4. Materials and Methods The experiment was designed and conducted at Bio-Park of Institute of Pure and Applied Biology, Bahauddin Zakariya University, Multan, Pakistan during two successive seasons 2017–2018 and 2018–2019. Surface sterilized (Sodium hypochlorite solution) seeds of two barley genotypes; B-14011 and B-10008 were grown in pots weighing 8 kg of river sand arranged in a completely randomized design (CRD) with four replicates of each treatment. The pots were arranged in two sets; 1st non-saline (irrigated with water + Hoagland nutrient solution) and second saline (irrigated with 200 mM NaCl + Hoagland nutrient solution). The 50% pots from non-saline and 50% from saline set were irrigated with 200 ppm Si (K2SiO3.2H2O) solution through rooting media when seedlings were two weeks old. During the third week of germination, 50% pots of each genotype were irrigated with 200 mM NaCl solution (saline), while the remaining 50% were irrigated with tap water (non-saline). Hoagland and Arnon [67] nutrient solution was supplied to plants to fulfill their nutrient requirements.PDF Image | Silicon-Induced Mitigation of NaCl Stress in Barley
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