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Arsenic contamination of soils threatens the health of millions globally through accumulation in crops. While plants detoxify arsenic via phytochelatin (PC) complexation and efflux of arsenite from roots, arsenite efflux mechanisms are not fully understood. Here, white lupin (Lupinus albus) was grown in semi-hydroponics and exudation of glutathione (GSH) derivatives and PCs in response to arsenic was scrutinised using LC-MS/MS. Inhibiting synthesis of PC precursor GSH with L-buthionine sulfoximine (BSO) or ABC transporters with vanadate drastically reduced (>22%) GSH-derivative and PC2 exudation, but not PC3 exudation. This was accompanied by arsenic hypersensitivity in plants treated with BSO and moderate sensitivity with vanadate treatment. Investigating arsenic-phytochelatin (As-PC) complexation revealed two distinct As-PC complexes, As bound to GSH and PC2 (GS-As-PC2) and As bound to PC3 (As-PC3), in exudates of As-treated lupin. Vanadate inhibited As-PC exudation, while BSO inhibited both the synthesis and exudation of As-PC complexes. These results demonstrate a role of GSH-derivatives and PC exudation in lupin arsenic tolerance and reveal As-PC exudation as a new potential mechanism contributing to active arsenic efflux in plants. Overall, this study uncovers insight into rhizosphere arsenic detoxification with potential to help mitigate pollution and reduce arsenic accumulation in crops.

Arbuscular Mycorrhizal (AM) fungi have substantial involvement in the existence of plants under heavy metal (HM)-stressed conditions. An overwhelming number of studies are there which advocate for the AM fungi as a future tool for remediation and revegetation of HM-polluted soils. One of the major complications associated with AM fungi facilitated phytoremediation is that the AM association is very much host as well as HM specific. Diverse strains of AM fungi behave differently with diverse hosts and HMs. AM fungi in association with host plants enhance the tolerance of HMs to the host. It enhances the absorption of nutrients in deficient soil, whereas it decreases the accumulation of HMs in polluted soils. AM fungi enhance the biomass production and, thus, dilute the HM concentration in plants. The association of AM fungi also protects the host roots from exposure of HMs by trapping them in the extracellular polymeric substances and the glomalin secreted by the AM fungal hyphae. The hyphal components also contain hydroxyl and carboxyl ligands that can bind the positively charged HMs and immobilize them outside the soil. In hyperaccumulator plants, AM fungi contribute differently as the transporters of HMs increase the uptake of HMs from the substrates. In this way, it enhances the accumulation of HMs beyond the permissible level. Inside the AM fungi as well as in the host cell, these HMs are either converted to the less toxic forms or conjugated with Metallothionein (MT), Glutathione (GSH), and Phytochelatin (PC) and safely stored in the vacuole. All these functions are specifically controlled by a number of transporters of HMs localized in the AM fungal hyphae, inside the AM fungi and inside the host cell. GintABC1 is one of the most studied transporters in AM fungi regulating the Zn. In this review, a deeper insight into the all-possible mechanisms of AM fungi facilitated HM stress alleviation in plants is summarized and discussed. [ABSTRACT FROM AUTHOR]

Rice is prone to take up the toxic elements arsenic (As) and cadmium (Cd) from paddy soil through the transporters for other essential elements. Disruption of these essential transporters usually adversely affects the normal growth of rice and the homeostasis of essential elements. Here we report on developing low-As and low-Cd rice grain through the co-overexpression of OsPCS1 , OsABCC1 , and OsHMA3 genes under the control of the rice OsActin1 promoter. Co-overexpression of OsPCS1 and OsABCC1 synergistically decreased As concentration in the grain. Overexpression of OsPCS1 also decreased Cd concentration in the grain by restricting the xylem-to-phloem Cd transport in node I, but paradoxically caused Cd hypersensitivity as the overproduced phytochelatins in OsPCS1 -overexpressing plants suppressed OsHMA3-dependent Cd sequestration in vacuoles and promoted Cd transport from root to shoot. Co-overexpression of OsHAM3 and OsPCS1 overcame this suppression and complemented the Cd hypersensitivity. Compared with non-transgenic rice control, co-overexpression of OsABCC1 , OsPCS1 , and OsHMA3 in rice decreased As and Cd concentrations in grain by 92.1% and 98%, respectively, without causing any defect in plant growth and reproduction or of mineral nutrients in grain. Our research provides an effective approach and useful genetic materials for developing low-As and low-Cd rice grain. [ABSTRACT FROM AUTHOR]

Agricultural soil salinization, which is often combined with heavy-metal contamination, is an ever-growing problem in the current era of global change. Legumes have a high potential for nitrogen fixation and are ideal crops for the reclamation of degraded soils. Alfalfa (Medicago sativa) is a valuable forage crop cultivated worldwide. Alfalfa plants fertilized with nitrogen or inoculated with a salt- and cadmium-tolerant Sinorhizobium meliloti strain were subjected to combined NaCl and CdCl2 stresses. Our results showed that inoculated plants presented higher aerial biomass than nitrogen-fertilized plants when they were exposed to salinity and cadmium together. To assess the mechanisms involved in the plant response to the combined stresses, superoxide dismutase and catalase antioxidant enzymatic activities were determined. Both increased upon stress; however, the increase in catalase activity was significantly less marked for inoculated plants, suggesting that other tolerance mechanisms might be active. Cd accumulation was lower in inoculated plants than in fertilized plants, which appears to imply that inoculation somehow prevented cadmium uptake by the plant roots. Expression analyses of several involved genes suggested that inoculation stimulated the biosynthesis of proline, phytochelatins, and homophytochelatins, together indicating that inoculated plants might be better suited to withstand combined salinity and cadmium stress effects. [ABSTRACT FROM AUTHOR]

Plants develop several external and internal mechanisms to increase their tolerance to heavy metals (HMs) toxicity including cadmium (Cd). Symbiosis with arbuscular mycorrhizae fungi (AMF) is one of the plants' strategies to tolerate HMs toxicity. Nitric oxide (NO), as a signaling molecule, is also involved in physiological responses of plants to various stresses. The present work was conducted as a factorial completely randomized design with three replications to study the effects of Funneliformis mosseae fungi and Sodium nitroprusside (SNP, 100 mM) as a donor of NO alone, in combination (AMF + SNP) on corn plant growth, and internal detoxification mechanisms of Cd toxicity in a Cd-contaminated calcareous soil (0, 25, 50, and 100 mg Cd kg−1). The results showed that under Cd stress, AMF inoculation and/or foliar application of SNP significantly increased plant growth (32% to 103% for shoot and 44% to 84% for root) by decreasing Cd concentration in corn plant tissues (23% to 46% for shoot and 19% to 40% for root). Cd-induced oxidative stress was mitigated by AMF and/or SNP by enhancing the activities of antioxidant enzymes, including superoxide dismutase (SOD) and catalase (CAT), and concentration of non-enzymatic antioxidants such as glutathione (GSH) and phytochelatin (PC). Increasing the tolerance index (TI) and decreasing the transfer factor (TF) in the corn plants treated with AMF and/or SNP, confirm the efficient role of SNP and AMF in stimulating the detoxification mechanisms of Cd within the plant cells, which was more pronounced at the lowest Cd level (25 mg Cd kg−1). In conclusion, symbiotic associations of corn plants with AMF alone or in combination with SNP mitigated the detrimental effect of Cd toxicity in corn grown in Cd-contaminated calcareous soil. The corn's internal detoxification mechanisms lowered the Cd concentration in plant tissue which resulted in the improvement of the corn's growth parameters. [ABSTRACT FROM AUTHOR]

Phytochelatins play an important role in plant resistance to cadmium. The pph6his gene encoding a peptide analog of natural phytochelatin with an α-peptide bond MetHys6[α-Glu(Cis)]6Gly was synthesized and used for genetic transformation of tobacco (Nicotiana tabacum L., variety Petit Havana, line SR1). The effect of cadmium on the accumulation of biomass and the content of glutathione, ascorbate, pigments (chlorophyll a, b and carotenoids) and malondialdehyde (MDA) in transgenic plants was evaluated. It was shown that plants of line 76 are able to grow and develop stably in the presence of up to 100 µmol of cadmium. These plants have also accumulated enough cadmium in the aerial parts to be used in phytoremediation. The created genetic construct can be used for transformation of various plant species. [ABSTRACT FROM AUTHOR]

Cadmium (Cd) uptake from polluted soils inhibits plant growth and disturbs physiological processes, at least partly due to disturbances in the cellular redox environment. Although the sulfur-containing antioxidant glutathione is important in maintaining redox homeostasis, its role as an antioxidant can be overruled by its involvement in Cd chelation as a phytochelatin precursor. Following Cd exposure, plants rapidly invest in phytochelatin production, thereby disturbing the redox environment by transiently depleting glutathione concentrations. Consequently, a network of signalling responses is initiated, in which the phytohormone ethylene is an important player involved in the recovery of glutathione levels. Furthermore, these responses are intricately connected to organellar stress signalling and autophagy, and contribute to cell fate determination. In general, this may pave the way for acclimation (e.g. restoration of glutathione levels and organellar homeostasis) and plant tolerance in the case of mild stress conditions. This review addresses connections between these players and discusses the possible involvement of the gasotransmitter hydrogen sulfide in plant acclimation to Cd exposure. [ABSTRACT FROM AUTHOR]

The endogenous stimulating molecule melatonin (N-acetyl-5-methoxytryptamine, MT) has an important function in mitigating the impact of multiple abiotic stressors. However, the ameliorating effect of MT on chromium (Cr) stress and its mechanisms remains unclear. Therefore, the present study aimed to clarify the mitigating effect of exogenous MT (0 μM and 100 μM) on wheat seedlings under Cr (0 μM and 50 μM) stress stemming from the growth and physiological characteristics, phytochelatin (PC) biosynthesis, Cr subcellular distribution, and antioxidant system of the plants in these treatments. The results showed that endogenous MT application significantly promoted plant growth and improved root morphology of wheat seedlings under Cr stress due to decreased Cr and reactive oxygen species (ROS) accumulation in both roots and leaves. Accumulation and transport of Cr from roots to leaves were reduced by MT, because enhanced vacuolar sequestration via upregulated PC accumulation, took place, derived from the fact that MT upregulated the expression of key genes for PC synthesis (TaPCS and Taγ-ECS). Furthermore, MT pre-treatment alleviated Cr-induced oxidative damage by diminishing lipid peroxidation and cell apoptosis, profiting from the enhanced scavenging ability of ROS as a result of the MT-induced increase in the activities of superoxide dismutase, catalase, ascorbate peroxidase, and glutathione reductase, and the related encoding gene expression levels of TaSOD2, TaCAT, TaAPX, and TaGR. In conclusion, endogenous MT application improved the growth traits, antioxidant system, and decreased Cr accumulation especially at the leaf level in wheat seedlings under Cr stress mainly through enhancing antioxidant enzyme activities and altering Cr subcellular distribution via strengthening PC biosynthesis. The mechanisms of MT-induced plant tolerance to Cr stress could help develop new strategies for secure crop production in Cr-polluted soils. [ABSTRACT FROM AUTHOR]

In highly intensive greenhouse vegetable production, soil acidification was caused by excessive fertilization, increasing cadmium (Cd) concentrations in the vegetables, which bears environmental hazards and is a negative influence on vegetables and humans. Transglutaminases (TGases), a central mediator for certain physiological effects of polyamines (PAs) in the plant kingdom, play important roles in plant development and stress response. Despite increased research on the crucial role of TGase in protecting against environmental stresses, relatively little is known about the mechanisms of Cd tolerance. In this study, we found, TGase activity and transcript level, which was upregulated by Cd, and TGase-induced Cd tolerance related to endogenous bound PAs increase and formation of nitric oxide (NO). Plant growth of tgase mutants was hypersensitive to Cd, chemical complementation by putrescine, sodium nitroprusside (SNP, nitric oxide donor) or gain of function TGase experiments restore Cd tolerance. α-diflouromethylornithine (DFMO, a selective ODC inhibitor) and 2–4-carboxyphenyl-4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide (cPTIO, NO scavenger), were respectively found declined drastically endogenous bound PA and NO content in TGase overexpression plants. Likewise, we reported that TGase interacted with polyamine uptake protein 3 (Put3), and the silencing of Put3 largely reduced TGase-induced Cd tolerance and bound PAs formation. This salvage strategy depends on TGase-regulated synthesis of bound PAs and NO that is able to positively increase the concentration of thiol and phytochelatins, elevate Cd in the cell wall, as well as induce the levels of expression Cd uptake and transport genes. Collectively, these findings indicate that TGase-mediated enhanced levels of bound PA and NO acts as a vital mechanism to protect the plant from Cd-caused toxicity.

With progress in technology, soaring demand for lithium (Li) has led to its release into the environment. This study demonstrated the mitigation of the adverse effects of Li stress on tomato (Solanum lycopersicum L.) by the application of waste materials, namely coconut shell biochar (CBC) and steel slag (SS). To explore the impact of Li treatment on tomato plants different morphological, biochemical parameters and plant defense system were analyzed. Tomato plants exposed to Li had shorter roots and shoots, lower biomass and relative water contents, and showed decreases in physiological variables, as well as increases in electrolyte leakage and lipid peroxidation. However, the application of CBC and SS as passivators, either singly or in combination, increased growth variables of tomato and relieved Li-induced oxidative stress responses. The combined CBC and SS amendments reduced Li accumulation 82 and 90% in tomato roots and shoots, respectively, thereby minimizing the negative impacts of Li. Antioxidant enzymes SOD, CAT, APX and GR reflected 4, 5, 30, and 52% and glyoxalase enzymes I and II 7 and 250% enhancement in presence of both CBC and SS in Li treated soil, with a concurrent decrease in methylglyoxal content. Lithium treatment triggered oxidative stress, increased enzymatic and non-enzymatic antioxidant levels, and induced the synthesis of thiols and phytochelatins in roots and shoots. Hence, co-amendment with CBC and SS protected tomato plants from Li-induced oxidative damage by increasing antioxidant defenses and glyoxalase system activity. Both CBC, generated from agricultural waste, and SS, an industrial waste, are environmentally benign, safe, economical, and non-hazardous materials that can be easily applied on a large scale for crop production in Li-polluted soils. The present findings highlight the novel reutilization of waste materials as renewable assets to overcome soil Li problems and emphasize the conversion of waste into wealth and its potential for practical applications. [Display omitted] • Coconut shell biochar and steel slag increased tomato plant growth under Li stress. • Accumulation of osmolytes under combined CBC and SS treatment, strengthened ROS scavenging mechanisms. • Li stress increased thiol and phytochelatin contents in roots and shoots. • Biochar and steel slag reduced oxidative damage by altering antioxidant defense. [ABSTRACT FROM AUTHOR]

Globally, metal/metalloid(s) soil contamination is a persistent issue that affects the atmosphere, soil, water and plant health in today's industrialised world. However, an overabundance of these transition ions promotes the excessive buildup of reactive oxygen species (ROS) and ion imbalance, which harms agricultural productivity. Plants employ several strategies to overcome their negative effects, including hyperaccumulation, tolerance, exclusion, and chelation with organic molecules. Polyamines (PAs) are the organic compounds that act as chelating agents and modulate various physiological, biochemical, and molecular processes under metal/metalloid(s) stress. Their catabolic products, including H 2 O 2 and gamma amino butyric acid (GABA), are also crucial signalling molecules in abiotic stress situations, particularly under metal/metalloid(s) stress. In this review, we explained how PAs regulate genes and enzymes, particularly under metal/metalloid(s) stress with a specific focus on arsenic (As), boron (B), cadmium (Cd), chromium (Cr), and zinc (Zn). The PAs regulate various plant stress responses by crosstalking with other plant hormones, upregulating phytochelatin, and metallothionein synthesis, modulating stomatal closure and antioxidant capacity. This review presents valuable insights into how PAs use a variety of tactics to reduce the harmful effects of metal/metalloid(s) through multifaceted strategies. • Metal/metalloid(s) soil pollution impacts the atmosphere, plant health, and crop production. • Polyamines (PAs) induce significant stress tolerance in plants. • PAs regulate physiological and metabolic responses in plants. • PAs promote secondary metabolite production under stress conditions. [ABSTRACT FROM AUTHOR]

• Pb toxicity disturbed the homeostasis between ROS and antioxidants. • Exposure to nSiO 2 induced a transient upregulation of RBOH gene in early hours that did not cause oxidative damage. • Foliar spray of nSiO 2 improved redox homeostasis and upregulated PCS1 transcripts. • ROS scavenging by DMTU and the inhibition of RBOH-dependent signaling by DPI aggravated Pb phytotoxicity and impaired the beneficial effects of nSiO 2. • H 2 O 2 signaling acts in downstream of nSiO 2 to mediate systemic acquired acclimation in Pb-stressed quinoa plants. The role of silicon dioxide nanoparticles (nSiO 2) and hydrogen peroxide (H 2 O 2) in heavy metal tolerance of plants has been well defined; however, their interaction together remains largely unknown. Therefore, two experiments were conducted to determine whether RBOH-dependent signaling is involved in nSiO 2 -induced systemic lead (Pb) acclimation in Chenopodium quinoa. In the first experiment, quinoa leaves were sprayed with nSiO 2 (0, 2, 4 mM) under various Pb concentrations (0, 200 and 400 µM). After 12 h, an H 2 O 2 burst, and higher transcription levels of RBOHD and PCS1 genes (encoding NADPH oxidase and phytochelatin synthase) were recorded in leaves of nSiO 2 -treated plants. After 15 days, Pb toxicity reduced growth parameters, and Chl a , Chl b contents and increased electrolyte leakage, and foliar spray with nSiO 2 reversed this tendency. In the second experiment, foliar application of 4 mM nSiO 2 minimized intracellular H 2 O 2 and O 2 ˙¯ and reduced malondialdehyde content and leaf Pb concentration under 400 µM Pb. While, this treatment increased the content of reduced ascorbate, and thiol compounds and enhanced the activities of superoxide dismutase and ascorbate peroxidase not only in leaves but also in untreated roots. However, H 2 O 2 scavenging with DMTU and the inhibition of RBOH-dependent signaling by DPI aggravated Pb phytotoxicity and impaired the beneficial effects of nSiO 2 on aforementioned attributes; thereby reduced tolerance index in both leaves and roots. Together, our pharmacological and molecular evidence for the first time unraveled that RBOH-dependent H 2 O 2 burst plays vital role in triggering nSiO 2 -induced systemic acquired acclimation in Pb-stressed plants. [Display omitted] [ABSTRACT FROM AUTHOR]

Accumulation of arsenic (As) and cadmium (Cd) in wheat grain is a serious threat to human health. Sulfur (S) can simultaneously decrease wheat grain As and Cd concentrations by decreasing their translocation in wheat; however, the mechanisms are unclear. We conducted hydroponic experiments to explore the mechanisms by which S modulates As and Cd translocation and their toxicity in wheat. Wheat seedlings were grown in deficient sulfate (2.5 μM) or sufficient sulfate (1.0 mM) nutrient solutions for 6 days and then exposed to zero (control), low As+Cd (1 μM As plus 0.5 μM Cd), or high As+Cd (50 μM As plus 30 μM Cd) for another 6 days. Compared with the control, plant growth was not affected by low As+Cd, but was significantly inhibited by high As+Cd. In the low As+Cd treatment, S supply had no significant effect on plant growth or root-to-shoot As and Cd translocation. In the high As+Cd treatment, sufficient S supply significantly alleviated As and Cd toxicity and their translocation by increasing phytochelatin (PC) synthesis and the subsequent vacuolar sequestration of As and Cd in roots, compared with deficient S supply. The use of L-buthionine sulfoximine (a specific inhibitor of g-glutamylcysteine synthetase) confirmed that the alleviation of As and Cd translocation and toxicity in wheat by S is mediated by increased PC production. Also, TaHMA3 gene expression in wheat root was not affected by the As+Cd and S treatments, but the expression of TaABCC1 was upregulated by the high As+Cd treatment and further increased by sufficient S supply and high As+Cd treatment. These results indicate that S-induced As and Cd subcellular changes affect As and Cd translocation mainly by regulating thiol metabolism and ABCC1 expression in wheat under As and Cd stress. [ABSTRACT FROM AUTHOR]

The dieback of Picea rubens Sarg. (red spruce) in the Appalachian Mountains of New England has been correlated with emissions transported from the Great Lakes region, including acids, metals, and oxidants. In 1994–1995, metal stress in red spruce foliage evidenced by phytochelatin concentrations increased with red spruce damage index in spruce-fir dominated stands in high elevation forests. In this study, we revisited those same forests after two decades to examine metal stress impacts on high-elevation forests following reductions in atmospheric pollutant loading. We measured metal concentrations in soils, lichens, and foliage, and concentrations of phytochelatin and its precursors in foliage of red spruce trees at 1000 m along a west–east transect from New York to New Hampshire, and along an 800–1000 m elevational transect on Whiteface Mountain, NY. Path analysis showed that foliar Hg had a direct positive effect on foliar phytochelatins, metal-binding peptides produced by the metal stress response in plants. Essential metals Cu and Zn decreased the concentration of Hg in foliage. However, we could not determine the relative importance of atmospheric vs soil pathways for metal exposure. While metal stress was still occurring on Whiteface Mountain in 2013, the overall visual health of red spruce trees across the region was significantly improved compared to 1993–1995. Thus, although metal stress is still measurable in red spruce, the physiological impact may be lessened by decreases in the deposition of metals and acids, thus providing evidence of positive forest health outcomes from improvements in regional air quality in the Northeastern US. [ABSTRACT FROM AUTHOR]

Arsenic (As) contamination of soils threatens the health of millions globally through accumulation in crops. While plants detoxify As via phytochelatin (PC) complexation and efflux of arsenite from roots, arsenite efflux mechanisms are not fully understood. Here, white lupin (Lupinus albus) was grown in semi-hydroponics, and exudation of glutathione (GSH) derivatives and PCs in response to As was measured using LC-MS/MS. Inhibiting synthesis of the PC precursor GSH with l-buthionine sulfoximine (BSO) or ABC transporters with vanadate drastically reduced (>22%) GSH derivative and PC2 exudation, but not PC3 exudation. This was accompanied by As hypersensitivity in plants treated with BSO and moderate sensitivity with vanadate treatment. Investigating As-PC complexation revealed two distinct As-PC complexes, As bound to GSH and PC2 (GS-As-PC2) and As bound to PC3 (As-PC3), in exudates of As-treated lupin plants. Vanadate inhibited As-PC exudation, while BSO inhibited both the synthesis and exudation of As-PC complexes. These results demonstrate a role for GSH derivatives and PC exudation in lupin As tolerance and reveal As-PC exudation as a new potential mechanism contributing to active As efflux in plants. Overall, this study uncovers insights into rhizosphere As detoxification with potential to help mitigate pollution and reduce As accumulation in crops., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Lines of transgenic plants Nicotiana tabacum L. expressing artificially synthesized gene PPH6HIS were studied in order to evaluate the contribution of the product, the metal binding peptide encoded by this gene, to the realization of the resistance of transformants to the effects of cadmium stress. In the course of the work, eight lines, expressing PPH6HIS that proven by Western blot analysis using His6-tag antibodies were received. An analysis of cadmium accumulation in explants and an assessment of the degree of damage to regenerants under the action of cadmium acetate showed that the lines of transgenic tobacco plants 13, 32, and 76 accumulated more cadmium than other transformed lines and control plants. In these three lines, signs of resistance to cadmium stress were observed, which was expressed in the preservation of the regenerative potential and a significantly lower level of leaf chlorosis in comparison with other lines and controls. These data allow us to conclude that the gene expression product of PPH6HIS, which is metal binding peptide, may be a promising candidate for increasing the accumulative properties of potential phytoremediant plants. [ABSTRACT FROM AUTHOR]

Rutin is a flavonoid with strong antioxidative effects on plant metabolism that facilitates resistance to environmental stress. The effect of foliar rutin on cadmium (Cd) uptake in Amaranthus hypochondriacus (K472) was studied. The results showed that a foliar spray of rutin alleviated Cd toxicity, promoted plant growth, improved Cd transfer to and storage in aerial plant parts and Cd accumulation with positive effects over time. A rutin concentration of 1.5 mg/mL showed the strongest promotion effect: the biomass and Cd content were increased at 13 days by 68.62% and 405.54% compared to 3 days, respectively, whereas a high concentration of rutin (5 mg/mL) inhibited plant growth and hindered Cd absorption. Two stages of Cd detoxification were identified in K472 after appropriate rutin application. First, an antioxidant system including an enzymatic antioxidant (superoxide dismutase [SOD]) and nonenzymatic antioxidants (glutathione [GSH] and flavonoids) was activated to enhance plant stress resistance. Quercetin and phytochelatin (PC) synthesis were then enhanced to perform detoxification synergistically with the antioxidant system to improve stress tolerance and achieve stable Cd detoxification. The results demonstrated that appropriately prolonging the application time of exogenous rutin to K472 is an effective way to improve the Cd remediation efficiency. The application of exogenous rutin to regulate the growth and Cd absorption of grain amaranth is reported for the first time. A foliar spray of rutin enriches Cd by regulating the metabolism of flavonoids and enhancing antioxidation and phytochelatin detoxification under Cd stress. Properly prolonging the harvest time after rutin treatment can greatly improve the Cd remediation efficiency of soil. The findings of the present study would be helpful for the remediation of Cd-contaminated soils. Rutin promotes the growth and Cd uptake of Amaranthus hypochondriacus. Extending the treatment time of rutin can improve Cd remediation efficiency. Rutin achieves stress detoxification for Cd by activating antioxidation. The synthesis and transfer of flavonoids and phytochelatin can reduce Cd toxicity. [ABSTRACT FROM AUTHOR]

Key message: An organomercurial phenylmercury activates AtPCS1, an enzyme known for detoxification of inorganic metal(loid) ions in Arabidopsis and the induced metal-chelating peptides phytochelatins are essential for detoxification of phenylmercury. Small thiol-rich peptides phytochelatins (PCs) and their synthases (PCSs) are crucial for plants to mitigate the stress derived from various metal(loid) ions in their inorganic form including inorganic mercury [Hg(II)]. However, the possible roles of the PC/PCS system in organic mercury detoxification in plants remain elusive. We found that an organomercury phenylmercury (PheHg) induced PC synthesis in Arabidopsis thaliana plants as Hg(II), whereas methylmercury did not. The analyses of AtPCS1 mutant plants and in vitro assays using the AtPCS1-recombinant protein demonstrated that AtPCS1, the major PCS in A. thaliana, was responsible for the PheHg-responsive PC synthesis. AtPCS1 mutants cad1-3 and cad1-6, and the double mutant of PC-metal(loid) complex transporters AtABCC1 and AtABCC2 showed enhanced sensitivity to PheHg as well as to Hg(II). The hypersensitivity of cad1-3 to PheHg stress was complemented by the own-promoter-driven expression of AtPCS1-GFP. The confocal microscopy of the complementation lines showed that the AtPCS1-GFP was preferentially expressed in epidermal cells of the mature and elongation zones, and the outer-most layer of the lateral root cap cells in the meristematic zone. Moreover, in vitro PC-metal binding assay demonstrated that binding affinity between PC and PheHg was comparable to Hg(II). However, plant ionomic profiles, as well as root morphology under PheHg and Hg(II) stress, were divergent. These results suggest that PheHg phytotoxicity is different from Hg(II), but AtPCS1-mediated PC synthesis, complex formation, and vacuolar sequestration by AtABCC1 and AtABCC2 are similarly functional for both PheHg and Hg(II) detoxification in root surficial cell types. [ABSTRACT FROM AUTHOR]

Globally, many people have been suffering from arsenic poisoning. Arsenate (AsV) exposure to twelve rice cultivars caused growth retardation, triggered production of As-chelatin biopeptides and altered activities of antioxidants along with increase in ascorbate (AsA)–glutathione (GSH) contents as a protective measure. The effects were more conspicuous in cvs. Swarnadhan, Tulaipanji, Pusa basmati, Badshabhog, Tulsibhog and IR-20 to attenuate oxidative-overload mediated adversities. Contrastingly, in cvs. Bhutmuri, Kumargore, Binni, Vijaya, TN-1 and IR-64, effects were less conspicuous in terms of alterations in the said variables due to reduced generation of oxidative stress. Under As(V) imposition, the protective role of phytochelatins (PCs) were recorded where peaks height and levels of PCs (PC2, PC3 and PC4) were elevated significantly in the test seedlings with an endeavour to detoxify cells by sequestering arsenic–phytochelatin (As–PC) complex into vacuole that resulted in reprogramming of antioxidants network. Additionally, scatter plot correlation matrices, color-coded heat map analysis and regression slopes demonstrated varied adaptive responses of test cultivars, where cvs. Bhutmuri, Kumargore, Binni, Vijaya, TN-1 and IR-64 found tolerant against As(V) toxicity. Results were further justified by hierarchical clustering. These findings could help to grow identified tolerant rice cultivars in As-prone soil with sustainable growth and productivity after proper agricultural execution. [ABSTRACT FROM AUTHOR]

Bermudagrass (Cynodon) species has been identified as a dominant plant species in cadmium (Cd)-contaminated soils. Objectives of current study were to evaluate Cd tolerance of two hybrid bermudagrass (Cynodon transvaalensis × Cynodon dactylon) cultivars Tifdwarf and Chuannong-3 for urban greening and slope ecological restoration and further to explore the difference in Cd tolerance associated with changes in ion absorption and distribution, antioxidant metabolism, and accumulations of phytochelatins (PCs) and metallothioneins (MTs) in roots, stems, and leaves. After exposure to equal concentration and duration of Cd stress, growth, chlorophyll content, and photochemical efficiency of both cultivars reduced significantly, but Cd-induced these inhibitory effects were more pronounced in Chuannong-3. As compared to Chuannong-3, Tifdwarf maintained significantly higher Cd concentration in roots and lower Cd concentration, bioconcentration factor, and translocation factor in aboveground parts in response to Cd stress. Each plant of Tifdwarf also accumulated more Cd in leaves, stems, and roots than each plant of Chuannong-3 due to larger biomass under Cd stress. Cd stress also significantly inhibited uptake and partitioning of iron, sodium, and potassium in two cultivars. In addition, Tifdwarf exhibited better antioxidant defense for reactive oxygen species (ROS) scavenging than Chuannong-3 under Cd stress, as reflected by higher antioxidant enzyme activities and antioxidant metabolites involved in ascorbic acid (ASA)-glutathione (GSH) cycle. Tifdwarf also accumulated more PCs in leaves, stems, and roots and more MTs in leaves and stems than Chuannong-3, which could help to chelate Cd to reduce cytotoxicity of Cd. Moreover, strong antioxidant property of MTs was beneficial for maintaining ROS homeostasis in plants. Therefore, better Cd tolerance of Tifdwarf could be mainly due to the inhibition of Cd uptake and partitioning, enhanced ASA-GSH cycle, and more accumulations of PCs and MTs. Tifdwarf showed better potential for Cd remediation or urban greening in Cd-polluted soils. • Tifdwarf accumulated more Cd in leaves, stems, and roots due to larger biomass. • Cd tolerance of Tifdwarf was related to inhibition of Cd uptake and partitioning. • Tifdwarf enhanced ASA-GSH cycle for reactive oxygen species scavenging. • Improved sulfur metabolism for GSH, PCs, and MTs to reduce cytotoxicity of Cd. • Tifdwarf showed better potential for phytoremediation in Cd-polluted soils. [ABSTRACT FROM AUTHOR]

Soil contamination with toxic metalloids, such as arsenic, can represent a substantial human health and environmental risk. Some plants are thought to tolerate soil toxicity using root exudation, however, the nature of this response to arsenic remains largely unknown. Here, white lupin plants were exposed to arsenic in a semi‐hydroponic system and their exudates were profiled using untargeted liquid chromatography‐tandem mass spectrometry. Arsenic concentrations up to 1 ppm were tolerated and led to the accumulation of 12.9 μg As g−1 dry weight (DW) and 411 μg As g−1 DW in above‐ground and belowground tissues, respectively. From 193 exuded metabolites, 34 were significantly differentially abundant due to 1 ppm arsenic, including depletion of glutathione disulphide and enrichment of phytochelatins and coumarins. Significant enrichment of phytochelatins in exudates of arsenic‐treated plants was further confirmed using exudate sampling with strict root exclusion. The chemical tolerance toolkit in white lupin included nutrient acquisition metabolites as well as phytochelatins, the major intracellular metal‐binding detoxification oligopeptides which have not been previously reported as having an extracellular role. These findings highlight the value of untargeted metabolite profiling approaches to reveal the unexpected and inform strategies to mitigate anthropogenic pollution in soils around the world. Root exudation response of white lupin to arsenic contamination was tested in a semi‐hydroponic system using untargeted tandem mass spectrometry. Significant enrichment of phytochelatins in exudates represents a potential mechanism which can explain the successful tolerance of metalloid excluder species. [ABSTRACT FROM AUTHOR]

Phytochelatins are plants’ small metal-binding peptides which chelate internal heavy metals to form nontoxic complexes. Detecting the complexes in plants would simplify identification of cultivars with both high tolerance and enrichment capabilities for heavy metals which represent phytoextraction performance. Thus, a terahertz spectroscopy combined with density functional theory, chemometrics and circular dichroism was used for characterization of phytochelatin2 (PC2), Cd-PC2 mixture standards, and pak choi (Brassica chinensis) leaves as a plant model. Results showed PC2 chelates Cd2+ in a 2:1 ratio to form Cd(PC2)2 complex; Cd connected to thoils of PC2 and changed β-turn and random coil of PC2 peptide chain to β-Sheet which presented as terahertz vibrations of PC2 around 1.03 and 1.71 THz being suppressed; the best models for detecting the complex in pak choi were obtained by partial least squares regression modeling combined with successive projections algorithm selection; the models used PC2 as a natural probe for visualizing and quantifying chelated Cd in pak choi leaf and achieved a limit of detection up to 1.151 ppm. This study suggested that terahertz information of the heavy metal-PCs complexes is qualified for representing a simpler alternative to classical index for evaluating phytoextraction performance of plant; it provided a general protocol for structure analysis and detection of heavy metal-PCs complexes in plant by terahertz absorbance.

Ethylenediamine tetraacetic acid (EDTA) is a chelating agent used worldwide for the phytoextraction of heavy metals using hyperaccumulating plants as well as in household and industrial purposes and persists in cultivated land for a long time without any known impact on nonhyperaccumulating plants. In this study, the effects of EDTA in the presence and absence of chromium on the hydroponic solution were evaluated at the cellular and molecular levels in nonhyperaccumulating rice plants (Oryza sativa L.). EDTA application enhanced phytochelatin (PC) and metallothionein (MT) synthesis, which was confirmed by the upregulation of phytochelatin and metallothionein genes in rice roots. However, a lower quantity of EDTA (25 µM) along with chromium enhanced chromium uptake (22.5%) in the root but inhibited its translocation to the shoot, indicating vacuolar sequestration of excessive chromium in the root. Conversely, in the absence of chromium, the H2O2 concentration was found to be increased, which significantly enhanced electrolyte leakage (98.5%) and lipid peroxidation (33.3%) compared with control plants. Our data indicate that excessive PCs and MTs without vacuolar sequestration enhance reactive oxygen species (ROS) generation. The findings of this investigation show that enhanced PCs and MTs owing to EDTA sequester excessive chromium in root vacuoles but, in the absence of sequestration due to a lack of chromium, enhance reactive oxygen species generation, resulting in toxicity. The outcomes of this investigation will open a new avenue of research in phytoextraction or phytoremediation activities.

Plants do not always have the genetic capacity to tolerate high levels of arsenic (As), which may not only arrest their growth but pose potential health risks through dietary bioaccumulation. Meanwhile, the interplay between the tomato plants and As–NO-driven molecular cell dynamics is obscure. Accordingly, seedlings were treated with As (10 mg/L) alone or in combination with 100 μM sodium nitroprusside (SNP, NO donor) and 200 μM 2-(4-carboxyphenyl)-4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide (cPTIO, NO scavenger). Sodium nitroprusside immobilized As in the roots and reduced the shoot translocation by up-regulating the transcriptional expression of the PCS , GSH1 , MT2, and ABC1. SNP further restored the growth retardation through modulating the chlorophyll and proline metabolism, increasing NO accumulation and stomatal conductance along with clear crosstalk between the antioxidant activity as well as glyoxalase I and II leading to endogenous H 2 O 2 and MG reduction. Higher PCs and glutathione accumulation helped protect photosynthetic apparatus; however, cPTIO reversed the protective effects of SNP, confirming the role of NO in the As toxicity alleviation. • SNP blocked As mobility in root and reduced shoot translocation by governing gene transcripts in As-sequestration network. • The expression of PCS , GSH1 , MT2 and ABC1 genes were up-regulated in As-stressed tomato. • SNP guided the proline and chlorophyll metabolism thereby improved tomato tolerance under As stress. • SNP decreased the H 2 O 2 and MG genesis by modulating overall anti-oxidant machinery, as well as the glyoxalase I and II. [ABSTRACT FROM AUTHOR]

Soil and water are among the most valuable resources on earth. Unfortunately, their contamination with heavy metals has become a global problem. Heavy metals are not biodegradable and cannot be chemically degraded; therefore, they tend to accumulate in soils or to be transported by streaming water and contaminate both surface and groundwater. Cadmium (Cd) has no known biological function but is one of the most toxic metals. It represents a serious environmental concern since its accumulation in soils is associated with health risks to plants, animals and humans. On the other hand, copper (Cu) and zinc (Zn) are heavy metals that are indispensable to plants but become toxic when their concentration in soils exceeds a certain optimal level. Plants have evolved many mechanisms to cope with heavy metal toxicity; vacuolar sequestration is one of them. Vacuolar sequestration can be achieved through either phytochelatin‐dependent or phytochelatin‐independent pathways. Most of the transgenic plants meant for phytoremediation described in the literature result from the manipulation of genes involved in the phytochelatin‐dependent pathway. However, recent evidence has emerged to support the importance of the phytochelatin‐independent pathway in heavy metal sequestration into the vacuole, with metal/H+ antiporters and proton pumps playing an important role. In this review, the importance of vacuolar proton pumps and metal/H+ antiporters transporting Cd, Cu, and Zn is discussed. In addition, the recent advances in the production of transgenic plants with potential application in phytoremediation and food safety through the manipulation of genes encoding V‐PPase proton pumps is described. [ABSTRACT FROM AUTHOR]

Pollution and poisoning with carcinogenic arsenic (As) is of major concern globally. Interestingly, there are ferns that can naturally tolerate remarkably high As concentrations in soils while hyperaccumulating this metalloid in their fronds. Besides Pteris vittata in which As-related traits and molecular determinants have been studied in detail, the As hyperaccumulation status has been attributed also to Pteris cretica. We thus inspected two P. cretica cultivars, Parkerii and Albo-lineata, for As hyperaccumulation traits. The cultivars were grown in soils supplemented with 20, 100, and 250 mg kg−1 of inorganic arsenate (iAsV). Unlike Parkerii, Albo-lineata was confirmed to be As tolerant and hyperaccumulating, with up to 1.3 and 6.4 g As kg−1 dry weight in roots and fronds, respectively, from soils amended with 250 mg iAsV kg−1. As speciation analyses rejected that organoarsenical species and binding with phytochelatins and other proteinaceous ligands would play any significant role in the biology of As in either cultivar. While in Parkerii, the dominating As species, particularly in roots, occurred as iAsV, in Albo-lineata the majority of the root and frond As was apparently converted to iAsIII. Parkerii markedly accumulated iAsIII in its fronds when grown on As spiked soils. Considering the roles iAsV reductase ACR2 and iAsIII transporter ACR3 may have in the handling of iAs, we isolated Albo-lineata PcACR2 and PcACR3 genes closely related to P. vittata PvACR2 and PvACR3. The gene expression analysis in Albo-lineata fronds revealed that the transcription of PcACR2 and PcACR3 was clearly As responsive (up to 6.5- and 45-times increase in transcript levels compared to control soil conditions, respectively). The tolerance and uptake assays in yeasts showed that PcACRs can complement corresponding As-sensitive mutations, indicating that PcACR2 and PcACR3 encode functional proteins that can perform, respectively, iAsV reduction and membrane iAsIII transport tasks in As-hyperaccumulating Albo-lineata.

Key message: Melatonin enhanced arsenic (As) tolerance by inhibiting As bioaccumulation, modulating the expression of As transporters and phytohormone homeostasis, leading to efficient utilization of thiol machinery for sequestration and detoxification of this toxic metalloid. The present study was aimed at investigating the influence of exogenous melatonin on the regulation of endogenous plant growth regulators and their cumulative effects on metal(loid)-binding ligands in two contrasting indica rice cultivars, viz., Khitish (arsenic sensitive) and Muktashri (arsenic tolerant) under arsenic stress. Melatonin supplementation ameliorated arsenic-induced perturbations by triggering endogenous levels of gibberellic acid and melatonin, via up-regulating the expression of key biosynthetic genes like GA3ox, TDC, SNAT and ASMT. The endogenous abscisic acid content was also enhanced upon melatonin treatment by induced expression of the key anabolic gene, NCED3 and concomitant suppression of ABA8ox1. Enhanced melatonin content induced accumulation of higher polyamines (spermidine and spermine), together with up-regulation of SPDS and SPMS in Khitish, thereby modulating stress condition. On the contrary, melatonin escalated putrescine and spermidine levels in Muktashri, via enhanced expression of ADC and SAMDC. The role of melatonin appeared to be more prominent in Khitish, as evident from better utilization of thiol components like cysteine, GSH, non-protein thiols and phytochelatins, with higher GSH/GSSG ratio, despite down-regulated expression of corresponding thiol-metabolic genes (OsMT2 and OsPCS1) to deal with arsenic toxicity. The extent of arsenic bioaccumulation, which was magnified several folds, particularly in Khitish, was decreased upon melatonin application. Overall, our observation highlighted the fact that melatonin enhanced arsenic tolerance by inhibiting arsenic bioaccumulation, via modulating the expression levels of selected arsenic transporters (OsNramp1, OsPT2, OsPT8, OsLsi1) and controlling endogenous phytohormone homeostasis, leading to efficient utilization of thiol machinery for sequestration and detoxification of this toxic metalloid. [ABSTRACT FROM AUTHOR]

Plants employ various defence strategies to ameliorate the effects of heavy metal exposures, leading to re-establishment of metal homeostasis. One of the strategies includes the biosynthesis of main heavy metal detoxifying peptides phytochelatins (PCs) by phytochelatin synthase (PCS). In the present study, 14 PCS homologues were identified in the genomes of 10 selected plants. The size of these PCSs was 452–545 amino acid residues, with characteristic phytochelatin and phytochelatin_C domains. The N-terminal site of the proteins is highly conserved, whereas the C-terminal site is less conserved. Further, the present study also identified fully conserved Cys residues involved in heavy metal binding reported earlier. In addition, other preserved cysteines, with minor substitutions Cys(C)→Ser(S) or Tyr(Y) or Trp(W), were also identified in the PCS sequences that might be associated with metal binding. The reported catalytic triad residues from Arabidopsis, Cys56, His162 and Asp180, are all conserved at the respective sites of PCSs. A clear monocot/dicot separation was revealed by phylogenetic analysis and was further corroborated by the exon–intron organisations of the PCS genes. Moreover, gene ontology terms, co-expression network, cis-regulatory motif and miRNA analyses indicated that the complex as well as dynamic regulation of PCSs has significant involvement in different metabolic pathways associated with signalling, defence, stress and phytohormone, in addition to metal detoxification. Moreover, variations in protein structure are suggested to confer the functional divergence in PCS proteins.

Aims: The contribution of sulphur (S)-induced responses to chromium (Cr) tolerance of rice plants is not yet fully elucidated. It is hypothesised that S nutrition mitigates the accumulation and toxicity of Cr through enhanced formation of iron plaque (IP) and S-containing chelators. This study aimed to investigate the responses of iron (Fe) and Cr availability and transfer in the hydroponic rice system to added S levels. Methods: We explored the influence of S nutrition on Cr accumulation in rice under a combination of Cr (VI) (+Cr, –Cr) and S (0, 1.75, 3.5, 7 mM) treatments. Results: S additions at rates of 1.75 and 3.5 mM gave the least decline in root and shoot growth of rice seedlings under Cr stress. Fe concentration in shoots was consistent with the level of Cr uptake. The subcellular distribution of Cr in roots and shoots differed with varying S supply levels. Our results also revealed that S treatment at a moderate level (3.5 mM) was more effective in suppressing the bioavailability of Cr in rice shoots than were the other levels. Conclusions: S-induced reduction in shoot Cr concentration, particularly from 1.75 to 3.5 mM, was likely attributed to the enhanced biosynthesis of glutathione (GSH) and phytochelatins (PCs) in roots than the enhanced physical resistance of IP induced by S. The poor barrier capacity of IP to Cr absorption in rice plants primarily ascribed to the level of applied Cr concentration and partly to the competition between Cr and S at the absorbing sites. [ABSTRACT FROM AUTHOR]

Lead (Pb) not only negatively alters plant growth and yield but may also have potentially toxic risks to human health. Nevertheless, the interaction between rice (Oryza sativa L.) plants and the molecular cell dynamics induced by lead-methyl jasmonate (MJ) remains unknown. Here, plants were hydroponically exposed to Pb (150 and 300 µM) alone or in combination with 0.5 and 1 µM MJ. The application of MJ modulated the expression of the HMAs, PCS1, PCS2 and ABCC1 genes, thereby immobilizing the Pb in the roots and lessening its translocation to the aerial parts of the rice plant. The supplementation of MJ improved the growth and yield of Pb-stressed rice by adjusting the proline and chlorophyll metabolism, increasing the phytochelatins (PCs) accumulation and diminishing the accumulation of Pb in the shoots. the application of MJ alleviated the oxidative stress of rice plants exposed to Pb toxicity by enhancing the activity of antioxidant enzymes and enzymes of the glyoxalase system (glyoxalase I and II) and decreasing the endogenous levels of malondialdehyde (MDA), hydrogen peroxide (H2O2) and methylglyoxal (MG). Therefore, the results of the present study could provide a molecular insight and cellular interplay scheme for the development of a promising strategy in Pb-contaminated areas to produce healthy food. [ABSTRACT FROM AUTHOR]

In this study E. coli recombinant clones that express the EC20 synthetic phytochelatin intracellularly were constructed. The increasement of Cd2+ biosorption capacity, and, also, the increasement of resistance to this toxic metal were analyzed. A gene that encodes the synthetic phytochelatin EC20 was synthesized in vitro. The EC20 synthetic gene was amplified by PCR, inserted into the DNA cloning vectors pBluescript®KS+ and pGEM®-TEasy, and also into the expression vectors pTE [pET-28(a)® derivative] and pGEX-T4-2®. The obtained recombinant plasmids were employed for genetic transformation of E. coli: pBsKS-EC20 and pGEM-EC20, they were introduced into DH10B and DH5α strains, similarly to pTE-EC20 and pGEX-EC20 that were introduced into BL21 strain. The EC20 expression was confirmed by SDS-PAGE analysis. The recombinant clones’ resistances to Cd2+ were determined by MIC analyses. The MIC for Cd2+ of DH10B/pBKS-EC20 and DH10B/pGEM-EC20 were 2.5 mM (EC20 induced), and 0.312 mM (EC20 repressed); respectively, 16 and 2 times higher than the control DH10B/pBsKS (0.156 mM). The MIC for Cd2+ of BL21/pTE-EC20 was 10.0 mM (EC20 induced) and 2.5 mM (EC20 repressed), compared with the control BL21/pTE which was only 1.25 mM. Analysis of ICP-AES showed that BL21/pGEX-EC20, after growth on the condition of EC20 expression, absorbed 37.5% of Cd2+, and even when cultured into the non-induction condition of EC20 expression, it absorbed 11.5%. These results allow the conclusion that recombinant E. coli clones expressing the synthetic phytochelatin EC20 show increased capacity for Cd2+ biosorption and enhanced resistance to this toxic ion.

The capacity of plants to accumulate cadmium (Cd) is significant for phytoremediation of Cd-polluted soils. Turnips cultivated in China include species featuring high Cd accumulation and some of these plants act as Cd hyperaccumulator landraces. These plants can accumulate over 100 mg Cd kg−1 dry weight in leaves without injury. Hence, studies that explore mechanisms underlying Cd detoxification and transport in turnip plants are essential. In the present study, we compared physiological and biochemical changes in turnip leaves treated with two Cd concentrations to controls. We discovered that Cd stress significantly increased the enzymatic activities or compound contents in the antioxidant system, including members of the glutathione-ascorbic acid cycle, whereas oxidation of reactive oxygen species (ROS) remained stable. Cd treatments also increased the contents of phytochelatins as well as a number of amino acids. Based on these results, we conclude that turnips initiate a series of response processes to manage Cd treatment. First, the antioxidant system maintaining ROS homeostasis and osmotic adjustment is excited to maintain stability of cell osmotic potential. Cd is chelated into its stable form to reduce its toxicity. Cd is possibly transported to vacuoles or non-protoplasts for isolation. Amino acid synthesis may directly and indirectly play an important role in these processes. This study partly revealed physiological and biochemical mechanisms underlying turnip response to Cd stress and provides information on artificially increasing or decreasing Cd accumulation in turnips and other plants.

Phytochelatin synthases (PCSs) play essential roles in detoxification of a broad range of heavy metals in plants and other organisms. Until now, however, no PCS gene from liverworts, the earliest branch of land plants and possibly the first one to acquire a PCS with a C-terminal domain, has been characterized. In this study, we isolated and functionally characterized the first PCS gene from a liverwort, Marchantia polymorpha (MpPCS). MpPCS is constitutively expressed in all organs examined, with stronger expression in thallus midrib. The gene expression is repressed by Cd2+ and Zn2+. The ability of MpPCS to increase heavy metal resistance in yeast and to complement cad1-3 (the null mutant of the Arabidopsis ortholog AtPCS1) proves its function as the only PCS from M. polymorpha. Site-directed mutagenesis of the most conserved cysteines of the C-terminus of the enzyme further uncovered that two twin-cysteine motifs repress, to different extents, enzyme activation by heavy metal exposure. These results highlight an ancestral function of the PCS elusive C-terminus as a regulatory domain inhibiting enzyme overactivation by essential and non-essential heavy metals. The latter finding may be relevant for obtaining crops with decreased root to shoot mobility of cadmium, thus preventing its accumulation in the food chain. [ABSTRACT FROM AUTHOR]

Chickpea (Cicer arietinum) belonging to the Fabaceae family is a major legume crop and is a good source of protein and carbohydrates. Industrialization has resulted in soil contamination with heavy metals such as cadmium. Adsorption of cadmium by plants can lead to reduced yields and heavy metal toxicity. In the current study, changes in the anatomical, morphological features and biochemical properties of the chickpea plant were evaluated. Two indexes DWSTI and PHSTI were determined. Anatomically, there was a change in the number of xylem poles within the root structure which was most significant at treatments of 125 μg cadmium. There was also a noticeable change in leaf pigmentation, the total phenolics and soluble protein in the plant. Cadmium levels were elevated attaining concentrations of 0.21, 0.40 and 0.52 mg per gram dry weight in plants exposed to 62, 125 and 250 μg/g Perlit cadmium after a period of 30 days. A noticeable increase in the level of cadmium in the plant was observed. Two PCS genes, glutathione gamma-glutamylcysteinyltransferase 1 and glutathione gamma-glutamylcysteinyltransferase and four FC genes, 4 proteins and 4 mRNA were detected in chickpeas. Bioinformatics tools were utilized to predict enzyme structure and binding sites. Chickpea may be classified as a cadmium hyperaccumulator and may be considered for use in phytoremediation. This study provides a better understanding with regards to the response of chickpeas to cadmium and the genetic mechanism by which the plant regulates heavy metal toxicity. [ABSTRACT FROM AUTHOR]

Cadmium is a non-essential trace metal, which is highly toxic to nearly all living organisms. Soil pollution causes Cd contamination of crops, thereby rendering plant products responsible for the chronic low level Cd over-exposure of numerous populations in the world. For this reason, Cd accumulation in plants has been studied for about five decades now. The research first focused on the relationships between plant and soil Cd levels, on the factors of the metal availability in soil, as well as the root uptake processes. Cd distribution in plant organs was also investigated, first using a macroscopic and eco-physiological approach, and then with the help of molecular biology tools, at both tissue and cell scales. Cadmium has no biological function and hijacks the transport pathways of micronutrients such as Fe, Mn, or Zn, in order to enter the plant through the roots and be distributed to all its organs. The study of the genes that control the influx and efflux of the Cd2+ ion in the cytosol, vacuoles, and vascular tissues has significantly contributed to the understanding of the metal root uptake and of its transfer to the aerial parts. However, the mechanisms responsible for its distribution to the different above-ground tissues and specially to fruits and seeds have yet to be clarified. This review summarizes current knowledge in order to present a detailed overview of Cd transport and storage, from the rhizosphere to the different organs and tissues of the plant. [ABSTRACT FROM AUTHOR]

Arsenic (As) contamination in paddy soil can cause phytotoxicity and elevated As accumulation in rice grains. Arsenic detoxification is closely linked to sulfur assimilation, but the genes involved have not been described in rice. In this study, we characterize the function of OASTL-A1, an O -acetylserine(thiol) lyase, in cysteine biosynthesis and detoxification of As in rice. Tissue expression analysis revealed that OsOASTL-A1 is mainly expressed in roots at the vegetative growth stage and in nodes at the reproductive stage. Furthermore, the expression of OsOASTL-A1 in roots was strongly induced by As exposure. Transgenic rice plants expressing pOsOASTL-A1::GUS (β-glucuronidase) indicated that OsOASTL-A1 was strongly expressed in the outer cortex and the vascular cylinder in the root mature zone. Subcellular localization using OsOASTL-A1:eGFP (enhanced green fluorescent protein) fusion protein showed that OsOASTL-A1 was localized to the cytosol. In vivo and in vitro enzyme activity assays showed that OsOASTL-A1 possessed the O -acetylserine(thiol) lyase activity. Knockout of OsOASTL-A1 led to significantly lower levels of cysteine, glutathione, and phytochelatins in roots and increased sensitivity to arsenate stress. Furthermore, the osoastl-a1 knockout mutants reduced As accumulation in the roots, but increased As accumulation in shoots. We conclude that OsOASTL-A1 is the cytosolic O -acetylserine(thiol) lyase that plays an important role in non-protein thiol biosynthesis in roots for As detoxification. [ABSTRACT FROM AUTHOR]

Cadmium (Cd) is detrimental to crops and the environment. This work examines the natural mechanisms underlying silicon‐ (Si‐)directed Cd detoxification in rice plants.The addition of Si to plants under Cd stress caused significant improvements in morphological parameters, chlorophyll score, Fv/Fm and total soluble protein concentration compared to controls, confirming that Si is able to ameliorate Cd‐induced damage in rice plants. This morpho‐physiological evidence was correlated with decreased cell death and electrolyte leakage after Si application.The results showed no critical changes in root Cd concentration, while shoot Cd decreased significantly after Si supplementation in comparison with Cd‐stressed rice. Additionally, expression of Cd transporters (OsNRAMP5 and OsHMA2) was significantly down‐regulated while the concentration of phytochelatin, cysteine and glutathione, together with expression of OsPCS1 (phytochelatin synthase) in roots of Cd‐stressed rice was significantly induced when subjected to Si treatment. This confirms that the alleviation of Cd stress is not only limited to the down‐regulation of Cd transporters but also closely related to the phytochelatin‐driven vacuolar storage of Cd in rice roots.The enzymatic analysis further revealed the role of SOD and GR enzymes in protecting rice plants from Cd‐induced oxidative harm. These findings suggest a mechanistic basis in rice plants for Si‐mediated mitigation of Cd stress. [ABSTRACT FROM AUTHOR]

Phytoremediation is a cost-effective and environment-friendly strategy for decontaminating heavy-metal-contaminated soil. However, the practical use of phytoremediation is constrained by the low biomass of plants and low bioavailability of heavy metals in soil. A pot experiment was conducted to investigate the effects of the metal chelator ethylenediaminetetraacetic acid (EDTA) and EDTA in combination with plant growth-promoting rhizobacteria (Burkholderia sp. D54 or Burkholderia sp. D416) on the growth and metal uptake of the hyperaccumulator Sedum alfredii Hance. According to the results, EDTA application decreased shoot and root biomass by 50% and 43%, respectively. The soil respiration and Cd, Pb, Zn uptake were depressed, while the photosynthetic rate, glutathione and phytochelatin (PC) contents were increased by EDTA application. Interestingly, Burkholderia sp. D54 and Burkholderia sp. D416 inoculation significantly relieved the inhibitory effects of EDTA on plant growth and soil respiration. Compared with the control, EDTA + D416 treatment increased the Cd concentration in shoots and decreased the Pb concentration in shoots and roots, but did not change the Zn concentration in S. alfredii plants. Furthermore, EDTA, EDTA + D54 and EDTA + D416 application increased the cysteine and PC contents in S. alfredii (p < 0.05); among all tested PCs, the most abundant species was PC2, and compared with the control, the PC2 content was increased by 371.0%, 1158.6% and 815.6%, respectively. These results will provide some insights into the practical use of EDTA and PGPR in the phytoremediation of heavy-metal-contaminated soil by S. alfredii. Image 1 [ABSTRACT FROM AUTHOR]

Plants employ various defence strategies to ameliorate the effects of heavy metal exposures, leading to re-establishment of metal homeostasis. One of the strategies includes the biosynthesis of main heavy metal detoxifying peptides phytochelatins (PCs) by phytochelatin synthase (PCS). In the present study, 14 PCS homologues were identified in the genomes of 10 selected plants. The size of these PCSs was 452–545 amino acid residues, with characteristic phytochelatin and phytochelatin_C domains. The N-terminal site of the proteins is highly conserved, whereas the C-terminal site is less conserved. Further, the present study also identified fully conserved Cys residues involved in heavy metal binding reported earlier. In addition, other preserved cysteines, with minor substitutions Cys(C)→Ser(S) or Tyr(Y) or Trp(W), were also identified in the PCS sequences that might be associated with metal binding. The reported catalytic triad residues from Arabidopsis, Cys56, His162 and Asp180, are all conserved at the respective sites of PCSs. A clear monocot/dicot separation was revealed by phylogenetic analysis and was further corroborated by the exon–intron organisations of the PCS genes. Moreover, gene ontology terms, co-expression network, cis-regulatory motif and miRNA analyses indicated that the complex as well as dynamic regulation of PCSs has significant involvement in different metabolic pathways associated with signalling, defence, stress and phytohormone, in addition to metal detoxification. Moreover, variations in protein structure are suggested to confer the functional divergence in PCS proteins. [ABSTRACT FROM AUTHOR]

It has been suggested that thiol compounds such as glutathione (GSH) and phytochelatin (PC) contribute to arsenic (As) tolerance mechanisms in higher plant. In contrast, the role of thiol compounds in As tolerance and hyperaccumulation in Pteris vittata, an As hyperaccumulator fern, has not been well demonstrated so far. The purpose of this study was to elucidate that role. Gel-filtration chromatography showed that fronds of P. vittata contained high-molecular-weight As compounds containing thiol compounds. Localization of As showed similar trends to that of thiol compounds, increasing with getting higher in position of pinna. Besides, the molar ratio of thiol to As was nearly equal to the ideal molar ratio of As-thiol complexes. These results suggest that As makes complex with thiol compounds in fronds of P. vittata. When comparing P. vittata with Nephrolepis exaltata, an As nonaccumulator fern, As-induced enhancement of thiol synthesis was observed only in P. vittata. The enhancement of thiol synthesis was induced directly by As, not indirectly by As-mediated oxidative stress. γ-glutamylcysteine (γEC) synthetase is a key enzyme for the synthesis of GSH and PC. Two putative genes of γEC synthetase, PvECS1 and PvECS2, were found in P. vittata. Expression of PvECS2 in shoots increased at 48 h after the onset of As exposure, almost corresponding to the increase in concentrations of γEC and GSH. Moreover, As accumulation in shoots began just before these increases. Thus, this study strongly suggests the contribution of thiol compounds to As tolerance in P. vittata.