Silicon Alleviates Copper Toxicity in Flax Plants by Up-Regulating Antioxidant Defense and Secondary Metabolites and Decreasing Oxidative Damage

In recent years, nutrient management has gained much attention as a way to mitigate heavy metal stress. Silicon (Si) promotes plant defense responses against toxic metal stresses. In this study, we evaluated the effects of silicon (Si) on copper (Cu) toxicity in two flax genotypes (Sakha 1 and Sakha 2) as it relates to plant growth, yield attributes, total chlorophyll, nucleic acid content, enzymatic and non-enzymatic antioxidants, oxidative damage, lipid peroxidation, copper and silicon content, and fatty acid composition. The results showed that Cu (100 and 200 µM) inhibited plant growth and increased Cu accumulation in soil, roots, and shoots. Cu significantly decreased the yield attributes, total chlorophyll by 9.5% and 22% in Sakha 1 and by 22.5% and 29% in Sakha 2, and enhanced the accumulation of non-enzymatic (tocopherol), enzymatic antioxidants such as superoxide dismnutase, peroxidase, ascorbate peroxidase and catalase) and secondary metabolites (phenol and flavonoids). The DNA content significantly decreased in stressed plants with 100 and 200 µM Cu about 22% and 44%, respectively, in Sakha 1 and about 21.6% and 34.7% in Sakha 2, and RNA content also decreased by about 20% and 29%, respectively, in Sakha 1 and by about 2% and 13% in Sakha 2 compared to the control plant. Furthermore, Cu stress accelerated the generation of reactive oxygen species (ROS), such as hydrogen peroxide (H 2 O 2 ) and induced cellular oxidative injury caused by lipid peroxidation. In parallel, Cu induced a change in the composition of fatty acids, resulting in lower unsaturated fatty acid levels and increased saturated fatty acids (increased saturation/unsaturation ratio for both genotypes). Treating the flax plants with irrigation three times with Si protected the plants from Cu toxicity. Si treatment decreased the uptake and the transport of Cu to the shoots and harvested seeds and promoted plant growth, yield attributes, and antioxidant defense systems by reducing Cu accumulation, lipid peroxidation, and the generation of H 2 O 2 . In addition, the alleviation of Cu toxicity correlated with increased Si accumulation in the roots and shoots. In conclusion, Si can be used to improve the resistance of flax plants to Cu toxicity by up-regulating the antioxidant defense system such as superoxide dismutase (SOD), peroxidase (POD), ascorbate peroxidase (APX) and catalase (CAT) and decreasing the oxidative damage caused by reactive oxygen species (ROS).