Cadmium (Cd) is one of the main pollutants in paddy fields, and its accumulation in rice (Oryza sativa L.) and subsequent transfer to food chain is a global environmental issue. This paper reviews the toxic effects, tolerance mechanisms, and management of Cd in a rice paddy. Cadmium toxicity decreases seed germination, growth, mineral nutrients, photosynthesis, and grain yield. It also causes oxidative stress and genotoxicity in rice. Plant response to Cd toxicity varies with cultivars, growth condition, and duration of Cd exposure. Under Cd stress, stimulation of antioxidant defense system, osmoregulation, ion homeostasis, and over production of signaling molecules are important tolerance mechanisms in rice. Several strategies have been proposed for the management of Cd-contaminated paddy soils. One such approach is the exogenous application of hormones, osmolytes, and signaling molecules. Moreover, Cd uptake and toxicity in rice can be decreased by proper application of essential nutrients such as nitrogen, zinc, iron, and selenium in Cd-contaminated soils. In addition, several inorganic (liming and silicon) and organic (compost and biochar) amendments have been applied in the soils to reduce Cd stress in rice. Selection of low Cd-accumulating rice cultivars, crop rotation, water management, and exogenous application of microbes could be a reasonable approach to alleviate Cd toxicity in rice. To draw a sound conclusion, long-term field trials are still required, including risks and benefit analysis for various management strategies.

Cadmium stress in rice: toxic effects, tolerance mechanisms, and management: a critical review.

A critical review of selenium biogeochemical behavior in soil-plant system with an inference to human health.

Increasing industrialization and urbanization result in emission of pollutants in the environment including toxic heavy metals, as cadmium and lead. Among the different heavy metals contaminating the environment, cadmium raises great concern, as it is ecotoxic and as such can heavily impact ecosystems. The cell wall is the first structure of plant cells to come in contact with heavy metals. Its composition, characterized by proteins, polysaccharides and in some instances lignin and other phenolic compounds, confers the ability to bind non-covalently and/or covalently heavy metals via functional groups. A strong body of evidence in the literature has shown the role of the cell wall in heavy metal response: it sequesters heavy metals, but at the same time its synthesis and composition can be severely affected. The present review analyzes the dual property of plant cell walls, i.e., barrier and target of heavy metals, by taking Cd toxicity as example. Following a summary of the known physiological and biochemical responses of plants to Cd, the review compares the wall-related mechanisms in early- and later-diverging land plants, by considering the diversity in cell wall composition. By doing so, common as well as unique response mechanisms to metal/cadmium toxicity are identified among plant phyla and discussed. After discussing the role of hyperaccumulators’ cell walls as a particular case, the review concludes by considering important aspects for plant engineering.

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Target or barrier? The cell wall of early- and later-diverging plants vs cadmium toxicity: differences in the response mechanisms

Cadmium stress in cotton seedlings: Physiological, photosynthesis and oxidative damages alleviated by glycinebetaine

Abiotic stresses, such as temperature extremes, drought, salinity, and heavy metals are major factors limiting crop productivity and sustainability worldwide. Abiotic stresses disturb plant growth and yield formation. Several chemical compounds, known as plant growth regulators (PGRs), modulate plant responses to biotic and abiotic stresses at the cellular, tissue, and organ levels. Thiourea (TU) is an important synthetic PGR containing nitrogen (36%) and sulfur (42%) that has gained wide attention for its role in plant stress tolerance. Tolerance against abiotic stresses is a complex phenomenon involving an array of mechanisms, and TU may modulate several of these. An understanding of TU-induced tolerance mechanisms may help improve crop yield under stress conditions. However, the potential mechanisms involved in TU-induced plant stress tolerance are still elusive. In this review, we discuss the essential role of TU-induced tolerance in improving performance of plants growing under abiotic stresses and potential mechanisms underlying TU-induced stress tolerance. We also highlight exploitation of new avenues critical in TU-induced stress tolerance.

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Potential Mechanisms of Abiotic Stress Tolerance in Crop Plants Induced by Thiourea.

Heat exposure alters the expression of SOD, POD, APX and CAT isozymes and mitigates low cadmium toxicity in seedlings of sensitive and tolerant rice cultivars.

Effect of Cd2+ toxicity and heat stress in sensitive rice cv. DR-92 and tolerant rice cv. Bh-1 grown in North East region of India were studied in sand cultures. Increasing levels of 0–500 μM Cd2+ alone and/or heat stress showed increased activities of superoxide dismutase, guaiacol peroxidase, ascorbate peroxidase and glutathione reductase enzymes which were associated with induced oxidative stress and altered enzyme activities. The values for SOD and POD activities were always more in cv. DR-92 whereas CAT and GR activities were higher in cv. Bh-1 in roots and shoots under Cd2+ or heat stress alone in sensitive cv. DR-92. Upon imposition of a combination of Cd2+ + heat the activities of SOD and POD decreased significantly in root/shoot of both the sensitive and tolerant rice varieties. A nine fold increase in GR activity under combination of heat + 100 μM Cd2+ stress in shoots of cv. Bh-1 at day 15 was noted when compared to controls. The dual stress combination of Cd2+ + heat did not alter catalase activity in vivo in both the rice varieties. Results suggest a time-specific and varietal distribution of the antioxidant enzymes in rice plants subjected to Cd2+ and/or heat stress. Tolerant cv. Bh-1 has better survival to combined stressors like Cd2+ and heat than sensitive rice cv. DR-92 and heat stress when given in combination with Cd2+ toxicity seem to mitigate the effect of Cd2+ stress alone in rice. The study indicates individual Cd2+ toxicity and heat stress and a combination of the two stresses to have separate implications on antioxidative defense mechanism in rice plants. Among enzymes of the defense apparatus ascorbate peroxidase and glutathione reductase appear to serve as an important component for better survival of rice plants under combination of Cd2+ + heat stress.

Expression of key antioxidant enzymes under combined effect of heat and cadmium toxicity in growing rice seedlings

Excess cadmium (Cd2+) in the soil environment is taken up by plants and can cause phytotoxicity. Elevated temperatures also lead to deleterious effects on plants. Plants are very often exposed to a combination of stresses rather than a single stress. The effect of Cd2+ and heat stress (HS) on the growth, root ultrastructure, lipid peroxidation (MDA), hydrogen peroxide accumulation and the activities of antioxidant enzymes peroxidase (POD), catalase (CAT) and ascorbate peroxidase (APX) of rice roots from sensitive cv. DR-92 and tolerant cv. Bh-1 were investigated at 10 and 20 day of growth under controlled conditions. At day 10 under all Cd2+ treatments, the Cd2+ content between the two rice cultivars were almost similar. Application of 500 μM Cd2+ significantly increased metal concentrations at day 20 in the roots of rice seedlings resulting in a maximum accumulation of 44.25 μg Cd2+ g-1 dry wt in cv. DR-92 and 30 μg Cd2+ g-1 dry wt in cv. Bh-1 with a ~25 % decline in Relative Growth Index (RGI) in cv. DR-92. TEM studies revealed slight disorganization with cell wall ingrowths in root tissues from cv. DR-92 grown in 100 μM Cd2+ + HS. Uptake and accumulation of Cd2+ increased upon heat treatment in parenchyma, vacuoles and vascular cylinder of root tissues. Peroxidase primarily located in cell walls, the intensity being higher in sensitive cv. DR-92. Under Cd2+ stress alone, plants of sensitive cv. DR-92 significantly increased the H2O2 and MDA levels together with increased activities of the enzymes POD, CAT and APX at day 10 but remained almost stable at day 20. A strong increase in MDA levels was noted at day 20 in tolerant cv. Bh-1. Cd2+ + HS treatments in tolerant cv.Bh-1 led to a decreased H2O2 and MDA levels and decreased activities of the enzymes POD, CAT and APX. Results suggest stimulation of root antioxidant system under combination of two stresses and that heat stress seem to have a direct protective role by mitigating the effect of mild Cd2+ toxicity largely by enhanced Cd2+-MT formation contributing thereby towards the management of Cd2+ toxicity at cellular level that confers Cd2+ tolerance to rice cv. Bh-1.

Effect of cadmium uptake and heat stress on root ultrastructure, membrane damage and antioxidative response in rice seedlings

The direct immobilization of soluble peroxidase isolated and partially purified from shoots of rice seedlings in calcium alginate beads and in calcium agarose gel was carried out. Peroxidase was assayed for guaiacol oxidation products in presence of hydrogen peroxide. The maximum specific activity and immobilization yield of the calcium agarose immobilized peroxidase reached 2,200 U mg−1 protein (540 mU cm−3 gel) and 82%, respectively. In calcium alginate the maximum activity of peroxidase upon immobilization was 210 mU g−1 bead with 46% yield. The optimal pH for agarose immobilized peroxidase was 7.0 which differed from the pH 6.0 for soluble peroxidase. The optimum temperature for the agarose immobilized peroxidase however was 30°C, which was similar to that of soluble peroxidase. The thermal stability of calcium agarose immobilized peroxidase significantly enhanced over a temperature range of 30∼60°C upon immobilization. The operational stability of peroxidase was examined with repeated hydrogen peroxide oxidation at varying time intervals. Based on 50% conversion of hydrogen peroxide and four times reuse of immobilized gel, the specific degradation of guaiacol for the agarose immobilized peroxidase increased three folds compared to that of soluble peroxidase. Nearly 165% increase in the enzyme protein binding to agarose in presence of calcium was noted. The results suggest that the presence of calcium, ions help in the immobilization process of peroxidase from rice shoots and mediates the direct binding of the enzyme to the agarose gel and that agarose seems to be a better immobilization matrix for peroxidase compared to sodium alginate.

Effect of calcium on immobilization of rice ( Oryza sativa L.) peroxidase for bioassays in sodium alginate and Agarose gel

The continuous addition of the heavy metal cadmium (Cd) in the biosphere results from industrial processes, from use of fertilizers or other crop protection agents, from municipal waste, and through atmospheric fallout. To ensure the protection of crops from this dangerous abiotic stress, it is important to understand and assess the availability, accumulation, uptake, and localization of cadmium in crop plants, in addition to its toxic responses. The uptake of Cd by plants results in various anatomical, histochemical, and biochemical alterations inside the plant cells. The Cd-induced toxicity resulting from its uptake and distribution in plants is dependent largely upon its uptake concentration in the plant and plant species, and even among varieties of the same species. Cadmium localization varies in different plants, causing anatomical abnormalities within the plant cells. This chapter is an update on the uptake, localization, and distribution of cadmium and the corresponding anatomical changes observed in crops grown in Cd-rich soil. This chapter also discusses the major abnormalities in different plant species based on the amount of Cd uptake and localization, in particular.

Sareeta Nahakpam

Papers

Heat exposure alters the expression of SOD, POD, APX and CAT isozymes and mitigates low cadmium toxicity in seedlings of sensitive and tolerant rice cultivars.

Expression of key antioxidant enzymes under combined effect of heat and cadmium toxicity in growing rice seedlings

Effect of cadmium uptake and heat stress on root ultrastructure, membrane damage and antioxidative response in rice seedlings

Effect of calcium on immobilization of rice ( Oryza sativa L.) peroxidase for bioassays in sodium alginate and Agarose gel

Cadmium-Induced Anatomical Abnormalities in Plants