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

Arsenic (As) is an environmental and food chain contaminant. Excessive accumulation of As, particularly inorganic arsenic (As(i)), in rice (Oryza sativa) poses a potential health risk to populations with high rice consumption. Rice is efficient at As accumulation owing to flooded paddy cultivation that leads to arsenite mobilization, and the inadvertent yet efficient uptake of arsenite through the silicon transport pathway. Iron, phosphorus, sulfur, and silicon interact strongly with As during its route from soil to plants. Plants take up arsenate through the phosphate transporters, and arsenite and undissociated methylated As species through the nodulin 26-like intrinsic (NIP) aquaporin channels. Arsenate is readily reduced to arsenite in planta, which is detoxified by complexation with thiol-rich peptides such as phytochelatins and/or vacuolar sequestration. A range of mitigation methods, from agronomic measures and plant breeding to genetic modification, may be employed to reduce As uptake by food crops.

Arsenic as a Food Chain Contaminant: Mechanisms of Plant Uptake and Metabolism and Mitigation Strategies

Arsenic (As) is an element that is nonessential for and toxic to plants. Arsenic contamination in the environment occurs in many regions, and, depending on environmental factors, its accumulation in food crops may pose a health risk to humans.Recent progress in understanding the mechanisms of As uptake and metabolism in plants is reviewed here. Arsenate is taken up by phosphate transporters. A number of the aquaporin nodulin26-like intrinsic proteins (NIPs) are able to transport arsenite,the predominant form of As in reducing environments. In rice (Oryza sativa), arsenite uptake shares the highly efficient silicon (Si) pathway of entry to root cells and efflux towards the xylem. In root cells arsenate is rapidly reduced to arsenite, which is effluxed to the external medium, complexed by thiol peptides or translocated to shoots. One type of arsenate reductase has been identified, but its in planta functions remain to be investigated. Some fern species in the Pteridaceae family are able to hyperaccumulate As in above-ground tissues. Hyperaccumulation appears to involve enhanced arsenate uptake, decreased arsenite-thiol complexation and arsenite efflux to the external medium, greatly enhanced xylem translocation of arsenite, and vacuolar sequestration of arsenite in fronds. Current knowledge gaps and future research directions are also identified.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1111/j.1469-8137.2008.02716.x

Arsenic uptake and metabolism in plants.

Heavy metal (HM) toxicity is one of the major abiotic stresses leading to hazardous effects in plants. A common consequence of HM toxicity is the excessive accumulation of reactive oxygen species (ROS) and methylglyoxal (MG), both of which can cause peroxidation of lipids, oxidation of protein, inactivation of enzymes, DNA damage and/or interact with other vital constituents of plant cells. Higher plants have evolved a sophisticated antioxidant defense system and a glyoxalase system to scavenge ROS and MG. In addition, HMs that enter the cell may be sequestered by amino acids, organic acids, glutathione (GSH), or by specific metal-binding ligands. Being a central molecule of both the antioxidant defense system and the glyoxalase system, GSH is involved in both direct and indirect control of ROS and MG and their reaction products in plant cells, thus protecting the plant from HM-induced oxidative damage. Recent plant molecular studies have shown that GSH by itself and its metabolizing enzymes—notably glutathione S-transferase, glutathione peroxidase, dehydroascorbate reductase, glutathione reductase, glyoxalase I and glyoxalase II—act additively and coordinately for efficient protection against ROS- and MG-induced damage in addition to detoxification, complexation, chelation and compartmentation of HMs. The aim of this review is to integrate a recent understanding of physiological and biochemical mechanisms of HM-induced plant stress response and tolerance based on the findings of current plant molecular biology research.

https://downloads.hindawi.com/archive/2012/872875.pdf

Molecular Mechanism of Heavy Metal Toxicity and Tolerance in Plants: Central Role of Glutathione in Detoxification of Reactive Oxygen Species and Methylglyoxal and in Heavy Metal Chelation

The biochemistry of environmental heavy metal uptake by plants: implications for the food chain.

Toxic heavy metals entering the plant tissues inhibit most physiological processes at all levels of metabolism. The inhibition of growth, photosynthesis, ion and water uptake and nitrate assimilation etc. have been described in many papers. The extent of these effects is greatly dependent on the concentration of the metal ions and the sensitivity or tolerance of the plant. Therefore the concentration is a key factor in interpreting metal actions. Another widely discussed question is the specific symptoms of heavy metal toxicity which is extremely hard to separate from those of other simultaneously affecting or consequently induced stresses.

Physiological Responses of Vascular Plants to Heavy Metals

Flow injection analysis (FIA) and high-performance liquid chromatography double-focusing sector field inductively coupled plasma mass spectrometry (HPLC-DF-ICP-MS) were used for total arsenic determination and arsenic speciation of xylem sap of cucumber plants (Cucumis sativus L.) grown in hydroponics containing 2 μmol dm−3 arsenate or arsenite, respectively. Arsenite [As(III)], arsenate [As(V)] and dimethylarsinic acid (DMA) were identified in the sap of the plants. Arsenite was the predominant arsenic species in the xylem saps regardless of the type of arsenic treatment, and the following concentration order was determined: As(III) > As(V) > DMA. The amount of total As, calculated taking into consideration the mass of xylem sap collected, was almost equal for both treatments. Arsenite was taken up more easily by cucumber than arsenate. Partial oxidation of arsenite to arsenate (<10% in 48 h) was observed in the case of arsenite-containing nutrient solutions, which may explain the detection of arsenate in the saps of plants treated with arsenite.

Arsenic speciation in xylem sap of cucumber (Cucumis sativus L.)

Effects of Pb and Cd on cucumber depending on the Fe-complex in the culture solution

This work was financed by an INCO-COPERNICUS (IC15-CT98-0126) grant, by Hungarian grants from OTKA (F-033076, T043646) and FKFP (0056/2001) and Spanish grants BOS2001-2343 and AGL2003-1999 from the MCE (co-financed with FEDER).

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Effects of two iron sources on iron and cadmium allocation in poplar (Populus alba) plants exposed to cadmium.

Mobility and distribution of lead (Pb) were investigated in cucumber (Cucumis sativus L.) grown in solution culture. Based on the observation that Pb uptake is influenced by the chemical form of iron (Fe) (complexed or ionic) in the solution, Fe distribution was also determined. Iron concentration was even in the stem (separated to internodes) and petioles but slightly increased at the shoot tip while Pb concentration decreased upwards. In the leaves Fe concentration decreased upwards (whereas in the control it increased a little at the youngest leaves) while Pb concentration increased to the largest leaves then decreased towards the shoot tip. It is suggested that the distribution of Pb in cucumber is passive process and it accumulates in the apoplast while Fe distribution is determined by the requirements of synthetic processes in young or photosynthesizing tissues. Lead probably lowers Fe transport into the symplast in the leaves causing slight chlorosis at the youngest leaves. In the plants t...

Ferenc Fodor

Papers

Physiological Responses of Vascular Plants to Heavy Metals

Arsenic speciation in xylem sap of cucumber (Cucumis sativus L.)

Effects of Pb and Cd on cucumber depending on the Fe-complex in the culture solution

Effects of two iron sources on iron and cadmium allocation in poplar (Populus alba) plants exposed to cadmium.

Lead uptake, distribution, and remobilization in cucumber