Introduction Plants, which cannot change their locations, are inevitably affected by environmental factors (Krasensky and Jonak, 2012). One of the major environmental factors is heavy metal toxicity. Heavy metals usually cause phytotoxicity and decrease of growth, yield, and quality in agricultural crops, even leading to deterioration of human health by entering the food chain (Yang et al., 2011). Due to its widespread distribution and high toxic potential for all living organisms, lead has been accepted as one of the most dangerous substances by the Agency for Toxic Substances and Disease (Fahr et al., 2013). Today, smelting of lead ores, mining, burning of fossil fuels, fertilizers, pesticides, and metal plating plants are the most important sources of lead pollution (Basharat et al., 2014). Although lead is not essential for plant nutrition, it is accumulated in plant organs by being taken up by the roots. It negatively affects plant metabolic processes such as seed germination, photosynthesis, and carbohydrate metabolism, and it can cause many physiological, biochemical, and structural disorders such as declines in chlorophyll contents, photosynthetic rate, and biomass as well as inhibition of the root and shoot growth (Verma and Dubey, 2003; Lamhamdi et al., 2011). Lead and other heavy metals also lead to the deterioration of nutrient balance in plants. Lead can inhibit the exchange of cations such as potassium, calcium, magnesium, iron, and zinc in roots (Fahr et al., 2013; Basharat et al., 2014). Unlike redox-active metals such as iron and copper, lead does not cause direct generation of reactive oxygen species (ROS) like superoxide anion (O 2-), hydroxyl radical (•OH), and hydrogen peroxide (H 2 O 2) by contributing to an increase in biological redox reactions such as Haber-Weiss and Fenton reactions. However, it leads to indirect increase in ROS formation by enhancing the activity of NADPH oxidases, displacing essential cations from specific binding sites of enzymes, and inhibiting the enzymatic activities due to the lead affinity of SH groups