Soil contamination with toxic environmental pollutants [such as cadmium (Cd)] is becoming a serious global problem due to rapid development of social economy. To improve the growth and yield of a plant, various chelating agents, such as ethylenediaminetetraacetic acid (EDTA) and citric acid (CA), can be applied to the soil; such application not only increases plant uptake of metals from the soil but also promotes plant absorption of micronutrient fertilizers from the medium. For this purpose, we have conducted a pot experiment using the exogenous application of CA (2.5 mM) and EDTA (2.5 mM) in pepper (Capsicum annuum L.) seedlings grown under the varying levels of Cd (0, 50 and 100 µM) in the soil. M]. Our results depicted that Cd addition to the soil significantly (P < 0.05) decreased plant growth and biomass, gas exchange attributes, and mineral uptake by C. annuum when compared to the plants grown without the addition of Cd. However, Cd toxicity boosted the production of reactive oxygen species (ROS) by increasing the content of malondialdehyde (MDA), which is the indication of oxidative stress in C. annuum, and was also manifested by hydrogen peroxide (H2O2) content and electrolyte leakage to the membrane-bound organelles. The results showed that the activities of various antioxidative enzymes, such as superoxidase dismutase (SOD), peroxidase (POD), catalase (CAT), and ascorbate peroxidase (APX), and their specific gene expression and also the content of non-enzymatic antioxidants, such as phenolic, flavonoid, ascorbic acid, and anthocyanin, initially increased with an increase in the Cd concentration in the soil. The results also revealed that the levels of soluble sugar, reducing sugar, and non-reducing sugar were decreased in plants grown under elevating Cd levels, but the accumulation of the metal in the roots and shoots of C. annuum, was found to be increased. The negative impacts of Cd injury were reduced by the application of EDTA and CA, which increased plant growth and biomass, improved photosynthetic apparatus, antioxidant enzymes and their gene expression, and mineral uptake, as well as diminished the exudation of organic acids and oxidative stress indicators in C. annuum by decreasing Cd toxicity. Here, we conclude that the application of EDTA and CA under the exposure to Cd stress significantly improved plant growth and biomass, photosynthetic pigments, and gas exchange characteristics; regulated antioxidant defense system and essential nutrient uptake; and balanced organic acid exudation pattern in C. annuum.
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