Differences in yield components and kernel Cd accumulation in response to Cd toxicity in four barley genotypes

A greenhouse hydroponics experiment was carried out to investigate genotypic difference in yield components in response to Cd toxicity, and kernel Cd concentrations and its relationship with Cd levels in roots and shoots during ontogenesis and as affected by shading and awn-removal. Root, shoot biomass and yield components of the four barley genotypes were impaired by increasing external Cd levels, with cv Wumaoliuling being most affected. Cadmium accumulation in roots and shoots increased with external Cd levels and differed significantly among genotypes. Meanwhile, 1 and 5 microM Cd treatments induced significant genotypic difference in kernel Cd concentrations, and Mimai 114, with the lowest Cd levels in roots and shoots, being the highest, whereas ZAU 3 being the lowest genotype. Shading had no significant effect on kernel Cd concentration, whereas awn-removal caused a significant decrease. Significantly negative correlation was discovered between Zn, Cu or Mn and Cd concentration in kernels, and there were positive relationships between Zn and Cu, Fe or Mn concentrations. Grain Cd concentrations were strongly correlated with both shoot and root levels. Regression equations between kernel- and shoot/root-Cd concentrations at different days of Cd exposure were established, allowing prediction of kernel Cd levels at harvest by measuring root- and shoot-Cd levels at early growth stage.     

Subcellular distribution and chemical form of Cd and Cd-Zn interaction in different barley genotypes

A hydroponic experiment was carried out to study the genotypic difference in subcellular distribution and chemical form of Cd and Zn uptake and their interaction in four barley genotypes. Increased Cd level in the medium caused a significant increase of Cd concentration in all fractions of roots/shoots, with most accumulation in FI (cell wall) and FIV (soluble). In root, the greatest amount of Cd was found in extraction solution of 2% HAC or 0.6M HCl, followed by 1M NaCl (20%), and lowest in extraction of 80% ethanol or d-H2O. While in shoot, the highest value of Cd accumulation was recorded in the fractions extracted by 1M NaCl and 2% HAC, followed by 0.6M HCl or d-H2O, and least in 80% ethanol extraction solution. There was a distinct difference among genotypes in Cd concentration in subcellular and chemical forms and it was found that the Cd-sensitive genotype Wumaoliuling, in comparison with the other three Cd-resistant genotypes, had higher Cd concentration in chloroplast-shoot/trophoplast-root (FII), membrane and organelle (FIII) and in inorganic and water-soluble Cd of roots, while lower in FI, FIV and pectates/protein integrated Cd. After 48 h of Cd treatment, the plants were replaced into Cd-free nutrient solution and grown for 72 h, a significant decrease in Cd concentration of root FI was observed, with less Cd reduction in Wumaoliuling. In comparison with control (no Cd), addition of Cd significantly increased Zn accumulation in chloroplast (FII), with least increase in Wumaoliuling. The Zn content in the other 3 fractions decreased significantly with addition of Cd, especially in roots, while Wumaoliuling was the most severely affected genotype. Moreover, it could be suggested Zn supplement could significantly reduce Cd concentration in root trophoplast and in shoot soluble fraction of Cd-treated plants.     

Role of salicylic acid in alleviating oxidative damage in rice roots (Oryza sativa) subjected to cadmium stress

Time-dependent changes in enzymatic and non-enzymatic antioxidants, and lipid peroxidation were investigated in roots of rice (Oryza sativa) grown hydroponically with Cd, with or without pretreatment of salicylic acid (SA). Exposure to 50 microM Cd significantly decreased root growth, and activities of superoxide dismutase (SOD), catalase (CAT) and peroxidase (POD), but increased the concentrations of H(2)O(2), malondialdehyde (MDA), ascorbic acid (AsA), glutathione (GSH) and non-protein thiols (NPT). However, pretreatment with 10 microM SA enhanced the activities of antioxidant enzymes and the concentrations of non-enzymatic antioxidants, but lowered the concentrations of H(2)O(2) and MDA in the Cd-stressed rice compared with the Cd treatment alone. Pretreatment with SA alleviated the Cd-induced inhibition of root growth. The results showed that pretreatment with SA enhanced the antioxidant defense activities in Cd-stressed rice, thus alleviating Cd-induced oxidative damage and enhancing Cd tolerance. The possible mechanism of SA-induced H(2)O(2) signaling in mediating Cd tolerance was discussed.     

Involvement of abscisic acid in regulating antioxidative defense systems and IAA-oxidase activity and improving adventitious rooting in mung bean [Vigna radiata (L.) Wilczek] seedlings under cadmium stress

In vitro experiments were conducted to investigate the effects of abscisic acid (ABA) and Cd on antioxidative defense systems and indole-3-acetic acid (IAA) oxidase during adventitious rooting in mung bean [Vigna radiata (L.) Wilczek] seedlings. The exogenous ABA significantly enhanced the number and fresh weight of the adventitious roots. CdCl₂ strongly inhibited adventitious rooting. Pretreatment with 10 μM ABA clearly alleviated the inhibitory effect of Cd on rooting. ABA significantly reduced superoxide dismutase (SOD), ascorbate peroxidase (APX), peroxidase (POD), and catalase (CAT) activities, as well as the levels of glutathione (GSH) and ascorbic acid (ASA) during adventitious rooting. ABA strongly increased IAA-oxidase activity during the induction (0–12 h) and expression (after 48 h) phases and increased the phenols levels. Cd treatment significantly reduced the activities of SOD, APX, POD, and IAA oxidase, as well as GSH level. Cd strongly increased ASA levels. ABA pretreatment counteracted Cd-induced alterations of certain antioxidants and antioxidative enzymes, e.g., remarkably rescued APX and POD activities, reduced the elevated SOD and CAT activities and ASA levels, and recovered the reduced GSH levels, caused by Cd stress. Thus, the physiological effects of the combination of ABA and Cd treatments were opposite of those obtained with Cd treatment alone, suggesting that ABA involved in the regulation of antioxidative defense systems and the alleviation of wounding- and Cd-induced oxidative stress.     

Contrasting behaviour of cadmium and zinc in a soil-plant-arthropod system

This study investigates the transfer of Cd and Zn from a soil amended with sewage sludge at rates up to 100 t ha(-1) through a multi-trophic system consisting of barley, the aphid Sitobion avenae and the larvae of the lacewing Chrysoperla carnae. Results show marked differences in the transfer of the two metals. Cadmium was freely accumulated in barley roots, but accumulation in the shoot was restricted to a concentration of around 0.22 mg kg(-1) (dry weight). This limited the transfer of Cd to higher trophic levels and resulted in no significant accumulation of Cd in S. avenae or in C. carnae. Zinc transfer in the system was largely unrestricted, resulting in significant accumulation in roots and shoots, in S. avenae and in C. carnae. Cadmium biomagnification occurred in lacewing pupae, with concentrations up to 3.6 times greater than in aphids. S. avenae biomagnified Zn by a factor of ca. 2.5 at low sludge amendment rates, but biomagnification decreased to a factor of 1.4 at the highest amendment rate. Biomagnification of Zn did not occur in C. carnae, but concentrations were up to 3.5 time higher than in soil. Results are discussed in light of the mechanisms regulating transfer of the two metals in the system.     

Effects of cadmium on glutathione synthesis in hepatopancreas of freshwater crab, Sinopotamon yangtsekiense

Cadmium (Cd) is one of the most deleterious heavy metals in aquatic systems that could promote oxidative damage. To explore the effects of Cd exposure of a freshwater crab (Sinopotamon yangtsekiense) on hepatopancreatic glutathione (GSH) synthesis, crabs were exposed to the reagent with a dose range of 7.25-116.00 mg L(-1) for 48 h. The concentrations of GSH, oxidized glutathione (GSSG), NADPH and NADP(+), as well as the activities of enzymes involved in GSH synthesis, i.e. glutathione reductase (GR), glucose-6-phosphate dehydrogenase (G6PD), gamma-glutamylcysteine synthetase (gamma-GCS) were determined. Progressive depletion of cellular GSH content was observed with the increasing of Cd concentrations, while the level of GSSG remained constant. In response to Cd exposure, crabs showed significant induction of G6PD and NADPH, however, only up to moderate exposures. GR activity remained at a steady level at all exposure concentrations. The activity of gamma-GCS was significantly positively correlated with the Cd concentration. These results suggested that GSH synthesis could be activated against reactive oxygen species induced by lower Cd exposure; under the higher Cd exposure conditions, an inhibition of NADPH-dependant redox cycling and de novo GSH synthesis led to significant decrease in GSH content.     

Antioxidative response to Cd in a newly discovered cadmium hyperaccumulator, Arabis paniculata F

A hydroponic experiment was carried out to study the effect of cadmium (Cd) on growth, Cd accumulation, lipid peroxidation, reactive oxygen species (ROS) content and antioxidative enzymes in leaves and roots of Arabis paniculata F., a new Cd hyperaccumuator found in China. The results showed that 22-89 microM Cd in solution enhanced the growth of A. paniculata after three weeks, with 21-27% biomass increase compared to the control. Cd concentrations in shoots and roots increased with increasing Cd supply levels, and reached a maximum of 1662 and 8670 mg kg(-1) Cd dry weight at 178 microM Cd treatment, respectively. In roots, 22-89 microM Cd reduced the content of malondialdehyde (MDA), superoxide (O(2)(-1)) and H(2)O(2) as well as the activities of superoxide dismutase (SOD), guaiacol peroxidase (GPX), ascorbate peroxidase (APX) and glutathione reductase (GR). In leaves, the contents of MDA, O(2)(-1) and H(2)O(2) remained unaffected by 22-89 microM Cd, while 178 microM Cd treatment significantly increased the MDA content, 69.5% higher than that of the control; generally, the activities of SOD, catalase (CAT), GPX and APX showed an increasing pattern with increasing Cd supply levels. Our present work concluded that A. paniculata has a great capability of Cd tolerance and accumulation. Moderate Cd treatment (22-89 microM Cd) alleviated the oxidative stress in roots, while higher level of Cd addition (178 microM) could cause an increasing generation of ROS, which was effectively scavenged by the antioxidative system.     

The effects of cadmium and zinc interactions on the accumulation and tissue distribution of zinc and cadmium in lettuce and spinach

The interactions between Zn and Cd on the concentration and tissue distribution of these metals in lettuce and spinach were studied at levels corresponding to background and Zn-Cd contaminated sites. Plants were grown in nutrient solutions containing 0.398-8.91 microM Zn and 0.010-0.316 microM Cd. Cadmium accumulated more in old than in young leaves of both crops at any solution Cd level, whereas Zn followed that pattern only at Zn levels > or = 3.16 microM. Increasing solution Cd increased Zn concentrations in young leaves of lettuce but not of spinach, regardless of Zn levels. Cadmium concentrations in young leaves of both crops decreased exponentially with increasing solution Zn at low (0.0316 microM) but not at high (0.316 microM) solution Cd. The Zn: Cd concentration ratios in young leaves of lettuce and spinach grown at 0.316 microM Cd became greater as the solution Zn increased. Cadmium and Zn concentrations in young leaves were related more closely to the relative concentrations of Zn and Cd in solution than were the concentrations in old leaves, especially in lettuce. Studies of Zn-Cd interactions and Cd bioavailability should differentiate between basal and upper leaves of lettuce and spinach. Compared to Cd-only pollution, Zn-Cd combined pollution may not decrease Cd concentrations in lettuce and spinach edible tissues, but because it increases their Zn concentrations it lowers plant Cd bioavailability.     

Calcium protects roots of Sedum alfredii H. against cadmium-induced oxidative stress

Sedum alfredii is a well-known Cd (cadmium) hyperaccumulator native to China. The impacts of exogenous Ca on Cd-induced oxidative stress and antioxidant systems in roots of S. alfredii were investigated by using cellular and biochemical approaches. Supplementation of the medium with higher Ca levels resulted in alleviated growth inhibition and decreased Cd concentration, as well as increased Ca concentration in roots. Cadmium induced lipid peroxidation and loss of plasma membrane integrity, reactive oxygen species overproduction, as well as ultrastructural changes of root cells were largely reversed by Ca supplementation in the medium. Calcium application significantly altered the Cd effects on antioxidant enzymes and non-enzyme antioxidants (non-protein thiols), and significantly increased glutathione (GSH) biosynthesis. The results suggest that Ca is able to protect the roots of S. alfredii against Cd toxicity by restoration of Cd-displaced Ca, alleviation of the metal induced oxidative stress, as well as promotion of GSH biosynthesis.     

Abolishing cadmium toxicity in Chlorella vulgaris by ascorbic acid, calcium, glucose and reduced glutathione

Addition of ascorbic acid, calcium (Ca2+), glucose and reduced glutathione (GSH) to the cadmium (Cd2+)-amended Kuhl medium significantly accelerated the growth and chlorophyll content of Chlorella vulgaris. This enhancement was found to be dose dependent. Their relative effectiveness against Cd2+ on growth and chlorophyll content has been arranged as GSH > Ca2+ > glucose > ascorbic acid. However, Ca2+ offered more protection against Cd2+ toxicity on protein contents, followed by ascorbic acid, GSH and glucose. The different concentrations of all amelioratives completely alleviated the toxicity of Cd2+ on total carbohydrates to exceed the control level. Photosynthetic oxygen evolution showed more response to Ca2+ and glucose in alleviating Cd2+ toxicity. On the other hand, at certain concentrations, GSH, Ca2+, ascorbic acid and glucose stimulated respiratory oxygen uptake over the control level. The uptake of Cd2+ dropped significantly in cultures supplemented with 20 microM ascorbic acid, glucose and GSH, while Ca2+ (30 microM) significantly lowered the Cd2+ uptake by 58% indicating that Ca2+ had a more antagonistic effect against Cd2+.     

The role of citric acid in cadmium and nickel uptake and translocation, in Halimione portulacoides

Some plants have high ability to absorb heavy metals in high concentrations. In this study, Halimione portulacoides was tested in conjunction with citric acid, in order to evaluate the possible use of this plant in phytoremediation processes in salt marshes. Two different concentrations of chelator were used combined with two heavy metal concentrations. When 25microM of citric acid was applied, Cd uptake and translocation was enhanced while for Ni these processes were almost inhibited. Increasing citric acid concentration to 50microM, Ni absorption decreased by the roots while for Cd there was still an increase in root uptake. Analysing translocation with this concentration of chelator, a decreased metal content in the upper organs for both metals was observed. While for Cd an optimal concentration of 25microM of citric acid was observed for phytoremediative processes, for nickel neither concentrations of chelator showed advantages for application in this remediative method.     

Silicon-mediated enhancement of cadmium tolerance in maize (Zea mays L.) grown in cadmium contaminated soil

Pot experiments were performed to study the alleviative effects of exogenous silicon (Si) on cadmium (Cd) phytotoxicity in maize grown in an acid soil experimentally contaminated with Cd. Five treatments were investigated in the first trial consisting of a control (neither Cd nor Si added), Cd added at 20 or 40 mg kg(-1) Cd without or with Si added at 400 mg kg(-1) Si. A following-up trial was conducted with almost the same treatments as in the first trial except that Si was incorporated at 50 mg kg(-1) Si. The results showed that Cd treatment significantly decreased shoot and root dry weight, while addition of Si at both levels significantly enhanced biomass. Addition of Si at 400 mg kg(-1) Si significantly increased soil pH but decreased soil Cd availability, thus reducing Cd concentration in the shoots and roots and total Cd in the shoots. Moreover, more Cd was found to be in the form of specific adsorbed or Fe-Mn oxides-bound fraction in the Si-amended soil. In contrast, soil pH, available Cd and Cd forms were unaffected by addition of Si at 50 mg kg(-1) Si, but shoot Cd concentration in the Si-amended Cd treatments significantly decreased at both Cd levels used compared to the non-Si-amended Cd treatments. Total Cd in the shoots and roots was considerably and significantly higher in the Si-amended Cd treatments than in the non-Si-amended Cd treatments. The xylem sap significantly increased but Cd concentration in the xylem sap significantly decreased in the Si-amended Cd treatments compared with the non-Si-amended Cd treatments irrespective of Cd and Si levels used. The results suggest that Si-enhanced tolerance to Cd can be attributed not only to Cd immobilization caused by silicate-induced pH rise in the soils but also to Si-mediated detoxification of Cd in the plants.     

The redox state and activity of superoxide dismutase classes in Arabidopsis thaliana under cadmium or copper stress

The redox state of glutathione and ascorbate as well as the activity of superoxide dismutase classes were determined in leaves of Arabidopsis thaliana grown for seven days in the nutrient solution containing 0, 5 and 50 microM Cd or Cu excess. A decrease in GSH/GSSG ratio was found in plants under Cd and Cu stress. In the plants exposed to Cu stress the activity of all SOD classes increased. However, in the plants treated with Cd the activity of FeSOD and MnSOD was elevated, but CuZnSOD activity was diminished in comparison with control. In these plants the activity of SOD classes was dependent on both the GSH/GSSG and AA/DHA ratios, while in those exposed to Cu excess - on the GSH/GSSG ratio. Differences were shown in the changes both in redox state and activity of SOD classes caused by the metals differing in physiochemical properties. Moreover, relationships between changes in SOD class activities and ROS levels were discussed.     

Overexpressing GSH1 and AsPCS1 simultaneously increases the tolerance and accumulation of cadmium and arsenic in Arabidopsis thaliana

The goal of this study was to develop transgenic plants with increased tolerance for and accumulation of heavy metals and metalloids from soil by simultaneous overexpression of AsPCS1 and GSH1 (derived from garlic and baker's yeast) in Arabidopsis thaliana. Phytochelatins (PCs) and glutathione (GSH) are the main binding peptides involved in chelating heavy metal ions in plants and other living organisms. Single-gene transgenic lines had higher tolerance to and accumulated more Cd and As than wild-type. Compared to single-gene transgenic lines, dual-gene transformants exhibited significantly higher tolerance to and accumulated more Cd and As. One of the dual-gene transgenic lines, PG1, accumulated twice the amount of Cd as single-gene transgenic lines. Simultaneous overexpression of AsPCS1 and GSH1 led to elevated total PC production in transgenic Arabidopsis. These results indicate that such a stacking of modified genes is capable of increasing Cd and As tolerance and accumulation in transgenic lines, and represents a highly promising new tool for use in phytoremediation efforts.     

Assessment of cadmium accumulation,toxicity, and tolerance in Brassicaceae and Fabaceae plants—implications for phytoremediation

This study, based on a greenhouse pot culture experiment conducted with 15-day-old rapeseed (Brassica campestris L. cv. Pusa Gold; family Brassicaceae) and moong bean (Vigna radiata L. Wilczek cv. Pusa Ratna; family Fabaceae) plants treated with cadmium (Cd) concentrations (0, 50, and 100 mg kg^?1 soil), investigates their potential for Cd accumulation and tolerance, and dissects the underlying basic physiological/biochemical mechanisms. In both species, plant dry mass decreased, while Cd concentration of both root and shoot increased with increase in soil Cd. Roots harbored a higher amount of Cd (vs. shoot) in B. campestris, while the reverse applied to V. radiata. By comparison, root Cd concentration was higher in B. campestris than in V. radiata. The high Cd concentrations in B. campestris roots and V. radiata shoots led to significant elevation in oxidative indices, as measured in terms of electrolyte leakage, H₂O₂ content, and lipid peroxidation. Both plants displayed differential adaptation strategies to counteract the Cd burden-caused anomalies in their roots and shoots. In B. campestris, increasing Cd burden led to a significantly decreased reduced glutathione (GSH) content but a significant increase in activities of GSH reductase (GR), GSH peroxidase (GPX), and GSH sulfotransferase (GST). However, in V. radiata, increasing Cd burden caused significant increase in GSH content and GR activity, but a significant decline in activities of GPX and GST. Cross talks on Cd burden of tissues and the adapted Cd tolerance strategies against Cd burden-accrued toxicity indicated that B. campestris and V. radiata are good Cd stabilizer and Cd extractor, respectively, wherein a fine tuning among the major components (GR, GPX, GST, GSH) of the GSH redox system helped the plants to counteract differentially the Cd load-induced anomalies in tissues. On the whole, the physiological/biochemical characterization of the B. campestris and V. radiata responses to varying Cd concentrations can be of great help in elaborating the innovative plant-based remediation technologies for metal/metalloid-contaminated sites.     

Characteristics of cadmium tolerance in 'Hermes' flax seedlings: contribution of cell walls

Most flax (Linum usitatissimum) varieties are described as tolerant to high concentrations of Cd. The aim of the present paper was to better characterize this tolerance, by studying the responses of flax plantlets, cv Hermes, to 18d growth on 0.5mM Cd. In Cd-treated seedlings, the majority of Cd was compartmentalized in the roots. Analysis of other elements showed that only Fe concentration was reduced, while Mn increased. Growth parameters of Cd treated flax were only moderately altered, with similar mass tolerance-indices for roots and shoots. Tissue anatomy was unaffected by treatment. The effect on lipid peroxidation, protein carbonylation and antioxidative activities appeared low but slightly higher in roots. The most important impacts of Cd were, in all organs, cell expansion, cell-wall thickening, pectin cross-linking and increase of cell-wall enzymatic activities (pectin methylesterase and peroxidase). Thus, the role of the cell wall in Cd tolerance might be important at two levels: (i) in the reinforcement of the tissue cohesion and (ii) in the sequestration of Cd.     

Effect of dietary protein on the distribution of cadmium,zinc and copper in rats following the oral administration of cadmium

Cadmium, 100 μg/rat/day, was administered orally 100 times (total amount of Cd administered, 10 mg), under conditions of low protein intake. The total amount of Cd in the liver and kidneys was as follows: females fed 20% protein diets > females fed 5% protein diets > males fed 20% protein diets ≒ males fed 5% protein diets. A decreased Zn concentration was found in the liver of males and the pancreas of males and females fed with the 5% protein diet. The Cu concentration in the kidneys was markedly influenced by low protein intake.     

Influence of cadmium on antioxidant capacity and four microelement concentrations in tomato seedlings (Lycopersicon esculentum)

Tomato (Lycopersicon esculentum) seedlings were grown in four cadmium (Cd) levels of 0-10 microM in a hydroponic system to analyze the antioxidative enzymes, Cd concentration in the plants, and the interaction between Cd and four microelements. The results showed that there was a significant increase in malondialdehyde (MDA) concentration, and superoxide dismutase (SOD) and peroxidase (POD) activities in the plants subjected to 1-10 microM Cd. This indicates that Cd stress induces an oxidative stress response in tomato plants, characterized by an accumulation of MDA and increase in activities of SOD and POD. Root, stem and leaf Cd concentrations increased with its exposure Cd level, and the highest Cd concentration occurred in roots, followed by leaves and stems. A concentration- and tissue-dependent response was found in the four microelement concentrations to Cd stress in the tomato leaves, stems and roots. Regression analysis showed that there was a significantly negative correlation between Cd and Mn, implying the antagonistic effect of Cd on Mn absorption and translocation. The correlation between Cd and Zn, Cu and Fe were inconsistent among leaves, stems and roots.     

Improved phytoaccumulation of cadmium by genetically modified tobacco plants (Nicotiana tabacum L.). Physiological and biochemical response of the transformants to cadmium toxicity

The response of tobacco plants (Nicotiana tabacum L.)--non-transformed and transformed with a metallothionein gene MThis from Silene vulgaris L.--to increase cadmium supply in the nutrient solution was compared. The transgenic plants accumulated significantly more Cd both in the roots and the leaves. Visual toxicity symptoms and disturbance in water balance were correlated with Cd tissue content. Treatment with 300 microM CdCl(2) resulted in inhibition of photosynthesis and mobilization of the ascorbate-glutathione cycle. Treatment with 500 microM CdCl(2) led to irreversible damage of photosynthesis and oxidative stress. An appearance of a new peroxidase isoform and changes in the leaf polypeptide pattern were observed at the highest Cd concentration. The level of non-protein thiols gradually increased following the Cd treatment both in transgenic and non-transformed plants.