Contamination of soils by heavy metals is of widespread occurrence as a result of human, agricultural and industrial activities. Among heavy metals, lead is a potential pollutant that readily accumulates in soils and sediments. Although lead is not an essential element for plants, it gets easily absorbed and accumulated in different plant parts. Uptake of Pb in plants is regulated by pH, particle size and cation exchange capacity of the soils as well as by root exudation and other physico-chemical parameters. Excess Pb causes a number of toxicity symptoms in plants e.g. stunted growth, chlorosis and blackening of root system. Pb inhibits photosynthesis, upsets mineral nutrition and water balance, changes hormonal status and affects membrane structure and permeability. This review addresses various morphological, physiological and biochemical effects of Pb toxicity and also strategies adopted by plants for Pb-detoxification and developing tolerance to Pb. Mechanisms of Pb-detoxification include sequestration of Pb in the vacuole, phytochelatin synthesis and binding to glutathione and aminoacids etc. Pb tolerance is associated with the capacity of plants to restrict Pb to the cell walls, synthesis of osmolytes and activation of antioxidant defense system. Remediation of soils contaminated with Pb using phytoremediation and rhizofiltration technologies appear to have great potential for cleaning of Pb-contaminated soils.

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Lead toxicity in plants

Effect of exogenous salicylic acid under changing environment: a review

Changes in the hormonal status of wheat seedlings induced by salicylic acid and salinity

Salicylic acid (SA) plays a key role in plant disease resistance and hypersensitive cell death but is also implicated in hardening responses to abiotic stressors. Cadmium (Cd) exposure increased the free SA contents of barley (Hordeum vulgare) roots by a factor of about 2. Cultivation of dry barley caryopses presoaked in SA-containing solution for only 6 h or single transient addition of SA at a 0.5 mM concentration to the hydroponics solution partially protected the seedlings from Cd toxicity during the following growth period. Both SA treatments had little effect on growth in the absence of Cd, but increased root and shoot length and fresh and dry weight and inhibited lipid peroxidation in roots, as indicated by malondialdehyde contents, in the presence of Cd. To test whether this protection was due to up-regulation of antioxidant enzymes, activities and transcript levels of the H(2)O(2)-metabolizing enzymes such as catalase and ascorbate peroxidase were measured in control and SA-treated seedlings in the presence or absence of 25 microM Cd. Cd stress increased the activity of these enzymes by variable extent. SA treatments strongly or completely suppressed the Cd-induced up-regulation of the antioxidant enzyme activities. Slices from leaves treated with SA for 24 h also showed an increased level of tolerance toward high Cd concentrations as indicated by chlorophyll a fluorescence parameters. The results support the conclusion that SA alleviates Cd toxicity not at the level of antioxidant defense but by affecting other mechanisms of Cd detoxification.

Salicylic Acid Alleviates the Cadmium Toxicity in Barley Seedlings

Salicylic acid induced changes on some physiological parameters symptomatic for oxidative stress and mineral nutrition in maize (Zea mays L.) grown under salinity.

Deterioration of membranes caused by lipoxygenase (LOX) activity under 10 µM PbCl2 or 10 µM HgCl2 was partially alleviated by the exogenous application of 100 µM salicylic acid (SA). In two cultivars of rice (Oryza sativa L. cvs. Ratna and IR 36), the presence of SA ameliorated the increased leakage of electrolytes, injury index, and the content of malondialdehyde caused by these heavy metals. Lead decreased H2O2 content whereas Hg increased it in both cultivars. Application of SA increased H2O2 in presence of Pb, while decreased it in presence of Hg. Both Pb and Hg decreased superoxide dismutase activity, while increased peroxidase activity. The activity of catalase was decreased by Hg but increased by Pb and SA reversed their effects. Thus, SA ameliorated the damaging effects of Pb and Hg on membranes.

Effects of Salicylic Acid on Heavy Metal-Induced Membrane Deterioration Mediated by Lipoxygenase in Rice

Germination of two rice (Oryza sativa L.) cultivars (Ratna and IR 36) in the presence of 10 µM PbCl2 and 10 µM HgCl2 decreased germination percentage, germination index, shoot/root length, tolerance index and dry mass of shoots and roots. Mercury was more toxic than lead. Reduced glutathione (GSH), cysteine (Cys), ascorbic acid, and α-tocopherol alleviated the adverse effects of these metals on plants in the order GSH > Cys > ascorbic acid > α-tocopherol. The effects were more pronounced in tolerant cultivar IR 36 than in the relatively susceptible cultivar Ratna.

A. Mishra

Papers

Effects of Salicylic Acid on Heavy Metal-Induced Membrane Deterioration Mediated by Lipoxygenase in Rice

Amelioration of lead and mercury effects on germination and rice seedling growth by antioxidants