Cd-sensitive Cultivar Research Articles

Cadmium tolerance and accumulation from the perspective of metal ion absorption and root exudates in broomcorn millet

Cadmium (Cd) is a persistent heavy metal that poses environmental and public health concerns. This study aimed to identify the potential biomarkers responsible for Cd tolerance and accumulation by investigating the response of the content of essential metal elements, transporter gene expression, and root exudates to Cd stress in broomcorn millet (Panicum miliaceum). A hydroponics experiment was conducted using two broomcorn millet cultivars with distinct Cd tolerance levels and accumulation phenotypes (Cd-tolerant and Cd-sensitive cultivars). Cd stress inhibited lateral root growth, especially in the Cd-sensitive cultivar. Furthermore, Cd accumulation was significantly greater in the Cd-tolerant cultivar than in the Cd-sensitive cultivar. Cd stress significantly inhibited the absorption of essential metal elements and significantly increased the calcium concentration. Differentially expressed genes involved in metal ion transport were identified via transcriptome analysis. Cd stress altered the composition of root exudates, thus increasing lipid species and decreasing alkaloid, lignan, sugar, and alcohol species. Moreover, Cd stress significantly reduced most alkaloid, organic acid, and phenolic acid exudates in the Cd-tolerant cultivar, while it increased most lipid and phenolic acid exudates in the Cd-sensitive cultivar. Some significantly changed root exudates (ferulic acid, O-coumaric acid, and spermine) are involved in the phenylalanine biosynthesis, and arginine and proline metabolic pathways, thus, may be potential biomarkers of Cd stress response. Overall, metal ion absorption and root exudates are critical for Cd tolerance and accumulation in broomcorn millet. These findings provide valuable insights into improving Cd phytoremediation by applying mineral elements or metabolites.

Multiomics reveals an essential role of long-distance translocation in regulating plant cadmium resistance and grain accumulation in allohexaploid wheat (Triticum aestivum).

Cadmium (Cd) is a highly toxic heavy metal that readily enters cereals, such as wheat, via the roots and is translocated to the shoots and grains, thereby posing high risks to human health. However, the vast and complex genome of allohexaploid wheat makes it challenging to understand Cd resistance and accumulation. In this study, a Cd-resistant cultivar of wheat, 'ZM1860', and a Cd-sensitive cultivar, 'ZM32', selected from a panel of 442 accessions, exhibited significantly different plant resistance and grain accumulation. We performed an integrated comparative analysis of the morpho-physiological traits, ionomic and phytohormone profiles, genomic variations, transcriptomic landscapes, and gene functionality in order to identify the mechanisms underlying these differences. Under Cd toxicity, 'ZM1860' outperformed 'ZM32', which showed more severe leaf chlorosis, poorer root architecture, higher accumulation of reactive oxygen species, and disordered phytohormone homeostasis. Ionomics showed that 'ZM32' had a higher root-to-shoot translocation coefficient of Cd and accumulated more Cd in the grains than 'ZM1860'. Whole-genome re-sequencing (WGS) and transcriptome sequencing identified numerous DNA variants and differentially expressed genes involved in abiotic stress responses and ion transport between the two genotypes. Combined ionomics, transcriptomics, and functional gene analysis identified the plasma membrane-localized heavy metal ATPase TaHMA2b-7A as a crucial Cd exporter regulating long-distance Cd translocation in wheat. WGS- and PCR-based analysis of sequence polymorphisms revealed a 25-bp InDel site in the promoter region of TaHMA2b-7A, and this was probably responsible for the differential expression. Our multiomics approach thus enabled the identification of a core transporter involved in long-distance Cd translocation in wheat, and it may provide an elite genetic resource for improving plant Cd resistance and reducing grain Cd accumulation in wheat and other cereal crops.

Deciphering distinct root exudation, ionomics, and physio-biochemical attributes of Serratia marcescens CP-13 inoculated differentially Cd tolerant Zea mays cultivars.

Cadmium (Cd) being a non-essential, mobile, and toxic heavy metal, negatively affects the plant growth and physiology. Current work investigated the impact of Serratia marcescens CP-13 inoculation on root organic acids and nutrient exudates of two maize cultivars varying in Cd tolerance under induced Cd toxicity. Seedlings of Cd-sensitive (Sahiwal-2002) and Cd-tolerant (MMRI-Yellow) cultivars were grown either inoculated or non-inoculated with CP-13 in Petri plates having various Cd stress levels (0, 6, 12, 18, 24, 30μM). Seedlings were transferred to rhizoboxes for the collection of root exudates and analysis of physio-biochemical traits. Both maize cultivars exuded higher organic acids and nutrient exudates under non-inoculated conditions as compared to inoculated ones. Non-inoculated tolerant cultivar exhibited higher nutrient accumulation, biomass, antioxidants, total chlorophyll, Cd release meanwhile reduced Cd uptake, lipid peroxidation, exudation of organic acids, and nutrients than the sensitive one. However, under CP-13 inoculation, Cd sensitive cultivar exhibited less exudation of organic acids (citric acid, acetic acid, malic acid, glutamic acid, formic acid, succinic acid, and oxalic acid), nutrients mobilization (K, Na, Zn, Ca, and Mg), total chlorophyll, antioxidants (APX, SOD, POD), total soluble sugar, diminished MDA, and Cd uptake. The significant reduction in release of root exudates by both cultivars was likely due to the plant growth promoting traits of CP-13 which confer Cd tolerance. The maximum release of rhizospheric root exudates were documented at 30μM applied Cd stress. Therefore, the Serratia sp. CP-13 was proposed as a potential inoculant for bioremediation of Cd together with maize cultivars.

Changes in plant growth, Cd partitioning and xylem sap composition in two sunflower cultivars exposed to low Cd concentrations in hydroponics

This study characterizes sunflower response to the levels of Cd encountered in moderately Cd-polluted soils. Two sunflower cultivars differing in their ability to sequestrate Cd in roots were exposed to low concentrations of Cd (0.5 nM or 100 nM) in hydroponics and sampled after 18 days (258 degree-days) when ten leaves were fully expanded. Plant growth, Cd uptake and partitioning among organs were monitored along with the ionomic (ICP-MS) and the metabolic (1H-NMR) composition of the xylem sap. Sunflower tolerance to Cd differed between the two cultivars. The cultivar with the highest ability to sequestrate Cd in roots (Kapllan) was more tolerant to Cd than the one with the lowest ability (ES RICA). The 23% penalization of plant growth observed at 100 nM in cultivar ES RICA was associated with reduced xylem loading fluxes of soluble sugars, perhaps pointing to disruption of carbohydrate metabolism. Retention of Cd in the stem was higher at 100 nM than at 0.5 nM in the Cd-sensitive cultivar ES RICA, which can be seen as a sunflower strategy to restrict the amount of Cd delivered to the leaves under Cd stress. No direct connection was found between the speciation of Cd in the xylem sap and the Cd translocation efficiency, although significant changes in the free ionic fraction of Cd were observed between the two cultivars at 0.5 nM. The relevance of these results in promoting the use of sunflower in phytomanagement of Cd-polluted soils is discussed.

Silicon nutrition lowers cadmium content of wheat cultivars by regulating transpiration rate and activity of antioxidant enzymes

Given that cadmium (Cd) uptake by plants is linked to transpiration rate and activity of antioxidant enzymes and further that silicon (Si) can regulate them, it was hypothesized that improved Si nutrition could reduce Cd concentration in plants. Thus, present study was carried out to elucidate the positive effect of Si nutrition on the growth, activities of antioxidant enzymes and tissue cadmium (Cd) concentration in Cd-tolerant (Iqbal-2000) and Cd-sensitive wheat (Triticum aestivum L.) cultivars. Fifteen days after seedling transplantation, 15 μM Cd stress alone and in combination with 0.6 mM Si was applied. Silicon application improved root and shoot dry matter of Cd-sensitive cultivar Sehar-2006 while the effect was non-significant in Cd-tolerant cultivar Iqbal-2000. Silicon-treated Cd-sensitive cultivar showed marked improvements in chlorophyll content and photosynthesis, while stomatal conductance and transpiration rate decreased by Si application. Silicon treatment enhanced the activities of enzymatic antioxidants including catalase, ascorbate peroxidase, guaiacol peroxidase and superoxide dismutase and the increase was higher for Cd-tolerant cultivar Iqbal-2000. Although Si nutrition depressed malondialdehyde (MDA) content in both Cd-stressed cultivars, the response was more evident in Cd-sensitive Sehar-2006. Lower lipid peroxidation was related to Si-induced increase in antioxidant activities only in Cd-sensitive cultivar. Silicon application decreased Cd accumulation in the roots and shoots of both the cultivars. The decrease in shoot Cd was associated with a decrease in Cd uptake by roots and Cd translocation from roots to shoots. Overall, it is concluded that Si suppressed Cd contents by decreasing transpiration rate in Cd-sensitive cultivar and by increasing antioxidant activity in Cd-tolerant cultivar.

Sulfur Protects Pakchoi (Brassica chinensis L.) Seedlings against Cadmium Stress by Regulating Ascorbate-Glutathione Metabolism.

Cadmium (Cd) pollution in food chains pose a potential health risk for humans. Sulfur (S) is a significant macronutrient that plays a significant role in the regulation of plant responses to diverse biotic and abiotic stresses. However, no information is currently available about the impact of S application on ascorbate-glutathione metabolism (ASA-GSH cycle) of Pakchoi plants under Cd stress. The two previously identified genotypes, namely, Aikangqing (a Cd-tolerant cultivar) and Qibaoqing (a Cd-sensitive cultivar), were utilized to investigate the role of S to mitigate Cd toxicity in Pakchoi plants under different Cd regimes. Results showed that Cd stress inhibited plant growth and induced oxidative stress. Exogenous application of S significantly increased the tolerance of Pakchoi seedlings suffering from Cd stress. This effect was demonstrated by increased growth parameters; stimulated activities of the antioxidant enzymes and upregulated genes involved in the ASA-GSH cycle and S assimilation; and by the enhanced ASA, GSH, phytochelatins, and nonprotein thiol production. This study shows that applying S nutrition can mitigate Cd toxicity in Pakchoi plants which has the potential in assisting the development of breeding strategies aimed at limiting Cd phytoaccumulation and decreasing Cd hazards in the food chain.

Antioxidant enzyme systems and the ascorbate–glutathione cycle as contributing factors to cadmium accumulation and tolerance in two oilseed rape cultivars (Brassica napus L.) under moderate cadmium stress

Oilseed rape (Brassica napus L.) with high tolerance to cadmium (Cd) may be used in the phytoremediation of Cd-contaminated fields. However, the mechanisms responsible for Cd accumulation and tolerance in oilseed rape are still poorly understood. Here, we investigated the physiological and molecular processes involved in Cd tolerance of two oilseed rape cultivars with different Cd accumulation abilities. The total Cd accumulation in cultivar L351 was higher than cultivar L338, particularly with increasing concentrations of Cd exposure. L338 was a more pronounced Cd-sensitive cultivar than L351, while higher activities of antioxidant enzymes (CAT, APX, GR, DHAR) as well as higher contents of GSH and AsA were all observed in L351 under Cd treatments, especially at high levels. No differences were found in SOD activities between the two cultivars under the same Cd treatments, suggesting that SOD was not the key factor in relation to the differences of Cd tolerance and accumulation between them. Gene expression levels of BnFe-SOD, BnCAT, BnAPX, BcGR and BoDHAR in roots of L351 were relatively higher than that in L338 under Cd exposure as well as BnCAT and BcGR in leaves. It is concluded that antioxidant enzymes and the ascorbate–glutathione cycle play important roles in oilseed rape Cd accumulation and tolerance.

The decline in potassium concentration is associated with cadmium toxicity of rice seedlings

Cadmium (Cd) is one of the most dangerous environmental pollutants, among other things, affecting plant mineral composition. Thus, in this study, we investigated the changes in potassium (K) concentration in Cd-treated rice (Oryza sativa L.) seedlings of two cultivars. On treatment with 5 μM CdCl2, the Cd concentration increased in the shoot and roots of Cd-sensitive cultivar (cv. Taichung Native 1, TN1) but not or slightly in the Cd-tolerant cultivar (cv. Tainung 67, TNG67). The decrease in K concentration in the shoot and roots of TN1 caused by Cd was more pronounced than that of TNG67. Exogenous addition of KCl decreased Cd concentration and reduced Cd toxicity of TN1 seedlings. Evidence presented in this study suggests that the improvement of K status is able to reduce toxicity of rice seedlings to CdCl2.

Silicon alleviates cadmium toxicity in peanut plants in relation to cadmium distribution and stimulation of antioxidative enzymes

Silicon (Si) is generally considered a beneficial element for the growth of higher plants, especially for those grown under stressed environments. Recently, the mitigating role of Si in cadmium (Cd) stress has received some attention. However, its mechanisms involved remain poorly understood. We studied the effects of Si on tissue and subcellular distribution of Cd, as well as the activities of major antioxidant enzymes (SOD, POD and CAT) with two contrasting peanut (Arachis hypogaea L.) cultivars (Luhua 11 and Luzi 101) differing in their Cd tolerance. The results showed that Cd exposure alone depressed plant growth and caused oxidative stress for both cultivars, and this toxicity was more obvious in Cd-sensitive cultivar (Luhua 11) than in Cd-tolerant cultivar (Luzi 101). Si supply significantly alleviated the toxicity of Cd in peanut seedlings; this was correlated with a reduction of shoot Cd accumulation, an alteration of Cd subcellular distribution in leaves, and a stimulation of antioxidative enzymes. The mechanisms of Si amelioration of Cd stress were cultivar and tissue dependent. For Luhua 11, Si-mediated inhibition of Cd transport from roots to shoots, reduction of Cd content in cell organelle fractions of leaves, and enhancement of the SOD, POD and CAT activities in roots, might responsible for the role of Si in alleviating Cd toxicity. For Luzi 101, Si alleviation of Cd toxicity is mainly attributed to the decrease in Cd concentration in shoot and stimulation of antioxidants systems.

Silicon-enhanced resistance to cadmium toxicity in Brassica chinensis L. is attributed to Si-suppressed cadmium uptake and transport and Si-enhanced antioxidant defense capacity

A series of hydroponics experiments were performed to investigate roles of silicon (Si) in enhancing cadmium (Cd) tolerance in two pakchoi ( Brassica chinensis L.) cultivars: i.e. cv. Shanghaiqing, a Cd-sensitive cultivar, and cv. Hangyoudong, a Cd-tolerant cultivar. Plants were grown under 0.5 and 5 mg Cd L −1 Cd stress without or with 1.5 mM Si. Plant growth of the Cd-tolerant cultivar was stimulated at the lower Cd level, but was decreased at the higher Cd level when plants were treated with Cd for one week. However, Plant growth was severely inhibited at both Cd levels as stress duration lasted for up to three weeks. Plant growth of the Cd-sensitive cultivar was severely inhibited at both Cd levels irrespective of Cd stress duration. Addition of Si increased shoot and root biomass of both cultivars at both Cd levels and decreased Cd uptake and root-to-shoot transport. Superoxide dismutase, catalase and ascorbate peroxidase activities decreased, but malondialdehyde and H 2O 2 concentrations increased at the higher Cd level, which were counteracted by Si added. Ascorbic acid, glutathione and non-protein thiols concentrations increased at the higher Cd level, which were further intensified by addition of Si. The effects of Si and Cd on the antioxidant enzyme activity were further verified by isoenzyme analysis. Silicon was more effective in enhancing Cd tolerance in the Cd-tolerant cultivar than in the Cd-sensitive cultivar. It can be concluded that Si-enhanced Cd tolerance in B. chinensis is attributed mainly to Si-suppressed Cd uptake and root-to-shoot Cd transport and Si-enhanced antioxidant defense activity.

Contributions of apoplasmic cadmium accumulation, antioxidative enzymes and induction of phytochelatins in cadmium tolerance of the cadmium-accumulating cultivar of black oat (Avena strigosa Schreb.)

The contributions of cadmium (Cd) accumulation in cell walls, antioxidative enzymes and induction of phytochelatins (PCs) to Cd tolerance were investigated in two distinctive genotypes of black oat (Avena strigosa Schreb.). One cultivar of black oat 'New oat' accumulated Cd in the leaves at the highest concentration compared to another black oat cultivar 'Soil saver' and other major graminaceous crops. The shoot:root Cd ratio also demonstrated that 'New oat' was the high Cd-accumulating cultivar, whereas 'Soil saver' was the low Cd-accumulating cultivar. Varied levels of Cd exposure demonstrated the strong Cd tolerance of 'New oat'. By contrast, low Cd-accumulating cultivar 'Soil saver' suffered Cd toxicity such as growth defects and increased lipid peroxidation, even though it accumulated less Cd in shoots than 'New oat'. Higher activities of ascorbate peroxidase (EC 1.11.1.11) and superoxide dismutase (EC 1. 15. 1. 1) were observed in the leaves of 'New oat' than in 'Soil saver'. No advantage of 'New oat' in PCs induction was observed in comparison to Cd-sensitive cultivar 'Soil saver', although Cd exposure increased the concentration of total PCs in both cultivars. Higher and increased Cd accumulation in cell wall fraction was observed in shoots of 'New oat'. On the other hand, in 'Soil saver', apoplasmic Cd accumulation showed saturation under higher Cd exposure. Overall, the present results suggest that cell wall Cd accumulation and antioxidative activities function in the tolerance against Cd stress possibly in combination with vacuolar Cd compartmentation.

Differences in Mn uptake and subcellular distribution in different barley genotypes as a response to Cd toxicity

A hydroponics experiment was carried out in greenhouse to study the genotypic differences in Mn uptake and subcellular distribution in response to Cd toxicity. Increased Cd level in medium caused a significant reduction in plant height and fresh weight, and ZAU3 and Wumaoliuling being the least and the most affected genotypes, respectively. There was a marked difference in proportion of Mn accumulation in different fractions relative to the total Mn amount in tissues among the 4 fractions, with the soluble fraction FIV showing the largest proportion in shoots, followed by organelle containing fraction (FIII), while cell wall (FI) and chloroplasts FII being the smallest. Meanwhile, Cd significantly increased FIII Mn accumulation proportion, but decreased FIV proportion, with significant genotypic difference of Wumaoliuling being the least increase in FIII and the greatest decrease in FIV among the 4 genotypes. In roots, the major pool of Mn content was FI, FIV, and FIII, and Cd induced no significant changes. Furthermore, Cd caused a significant reduction in subcellular Mn concentration of FI and FIV fractions in shoots and the 4 fractions in roots, with more pronounced in Cd-sensitive cultivar Wumaoliuling in root FII, FIII and FIV, and shoot FI, FII, and FIII, while little difference in both Mn concentrations of root FI, and shoot FIV.

Antioxidant enzyme activities are upregulated in response to cadmium in sensitive, but not in tolerant, rice (Oryza sativa L.) seedlings

Changes in H 2 O 2 and malondialdehyde (MDA) contents and antioxidant enzyme activities in Cd-treated rice (Oryza sativa L.) seedlings of two cultivars were investigated. On treatment with CdCl 2 , increases in H 2 O 2 and MDA contents and antioxidant enzyme activities (speroxide dismutase (SOD), ascorbate peroxidase, glutathione reductase, catalase, and peroxidase (POX)) were observed in the leaves of Cd-sensitive cultivar (cv. Taichung Native 1, TN1) but not in Cd-tolerant cultivar (cv. Tainung 67, TNG67). The increased content of MDA and activities of SOD and POX preceded the occurrence of toxicity in CdCl 2 -treated TN1 leaves. Pretreatment with abscisic acid (ABA) enhanced Cd tolerance and reduced Cd-induced increase in the content of MDA and increase in the activities of SOD and POX in TN1 leaves. Exogenous application of ABA biosynthesis inhibitor, fluridone, decreased Cd tolerance, increased the content of MDA, and increased the activities of SOD and POX in Cd-treated TNG67 leaves. Furthermore, fluridone's effects on toxicity, the content of MDA, and the activities of SOD and POX in Cd-treated TNG67 leaves were reversed by the application of ABA. In conclusion, the oxidative stress is differently expressed in TN1 and TNG67 rice seedlings in response to CdCl 2 . Results also suggest that CdCl 2 causes an oxidative stress and CdCl 2 -induced toxicity is mediated through oxidative stress in TN1 leaves.

Role of abscisic acid in cadmium tolerance of rice (Oryza sativa L.) seedlings.

Changes in abscisic acid (ABA) contents in Cd-treated rice (Oryza sativa L.) seedlings of two cultivars were investigated. On treatment with CdCl2, the ABA content rapidly increased in the leaves and roots of Cd-tolerant cultivar (cv. Tainung 67, TNG67) but not in the Cd-sensitive cultivar (cv. Taichung Native 1, TN1). The reduction of transpiration rate of TN1 caused by Cd was less than that of TNG67. Exogenous application of ABA reduced transpiration rate, decreased Cd content, and enhanced Cd tolerance of TN1 seedlings. Exogenous application of the ABA biosynthesis inhibitor, fluridone, reduced ABA accumulation, increased transpiration rate and Cd content, and decreased Cd tolerance of TNG67 seedlings. Fluridone effect on Cd toxicity of TNG67 seedlings was reversed by the application of ABA. The roles of endogenous ABA in Cd tolerance of rice seedlings are discussed and suggested.

Accumulation of ammonium ion in cadmium tolerant and sensitive cultivars of Oryza sativa

Cd-tolerant and Cd-sensitive rice cultivars were used to study the role of NH4+ accumulation in Cd-induced toxicity. NH4+ accumulation seems to be involved in regulating the toxicity of rice seedlings caused by CdCl2. This conclusion was based on the observations that (a) on treatment with CdCl2, NH4+ content increased rapidly in the leaves of the Cd-sensitive cultivar (cv. Taichung Native 1, TN1) but not in the Cd-tolerant cultivar (cv. Tainumg 67, TNG67), (b) pretreatment with abscisic acid (ABA) enhanced Cd tolerance and reduced Cd-induced NH4+ accumulation in TN1 seedlings, (c) exogenous application of the ABA biosynthesis inhibitor, fluridone, decreased Cd tolerance and increased NH4+ content in leaves of TNG67, (d) exogenous application of phosphinothricin, an inhibitor of glutamine synthetase (GS), which resulted in NH4+ accumulation in the leaves, also induced toxicity similar to Cd in TN1 seedlings. Evidence is presented to show that Cd-induced NH4+ accumulation in TN1 leaves is attributable to a decrease in GS activity. Since Cd-treated TN1 leaves had higher glutamine and glutamate contents than control leaves, it is unlikely that glutamine (or glutamate) depletion is the mechanism which regulates Cd-induced toxicity.