Lead and cadmium tolerance and accumulation of proanthocyanidin- deficient mutants of the fern Athyrium yokoscense

The fern Athyrium yokoscense often flourishes around mine sites in Japan and can tolerate and accumulate heavy metals such as lead (Pb) and cadmium (Cd). In this work, we examined whether proanthocyanidins, also called condensed tannins, were involved in the mechanisms of Pb and Cd tolerance and accumulation of A. yokoscense because proanthocyanidins are known to alleviate metal stress in several plant species and are present at high levels in A. yokoscense. For this purpose, we used mutant gametophytes deficient in proanthocyanidins, in which the relative proanthocyanidin contents were 20% of those of the wild-type gametophytes. Although the proanthocyanidin contents of the mutant were quite low, the growth of the mutant was very similar to that of the wild-type gametophytes even in the presence of 80 mg/kg Pb or 48 mg/kg Cd. Under the same conditions, the mutant gametophytes also accumulated Pb and Cd as much as the wild-type gametophytes did. These results indicate that the proanthocyanidins in A. yokoscense are not important for the Pb and Cd tolerance and accumulation properties.

Fungal Seed Endophyte FZT214 Improves Dysphania ambrosioides Cd Tolerance Throughout Different Developmental Stages

Phytoremediation is a promising remediation method of heavy metal (HM)–contaminated soils. However, lower HM tolerance of metal accumulator inhibits its practical application and effects. The current study was aimed to illustrate the role of fungal seed endophyte (FZT214) in improving Dysphania ambrosioides Cd tolerance during different developmental stages under various Cd stresses (5, 15, 30 mg kg–1) by pot experiments. The results showed that FZT214 significantly (p < 0.05) improved the host plant’s growth at the flowering and fruiting stage in most of the treatment, while at the growing stage the increase was less (p > 0.05). The seed yield was also improved (p < 0.05) in the FZT214-inoculated plants (E+) and induced early flowering was observed. Moreover, the inoculation also positively affected total chlorophyll content, antioxidant process, and lipid peroxidation in most of the treatments throughout three developmental stages. Not all but in most cases, IAA and GA were more in E+ plants while JA was more in the E− plants (non-inoculated plants) during three developmental stages. The results suggested that the colonization of FZT214 to the D. ambrosioides might trigger multiple and comprehensive protective strategies against Cd stress, which mainly include activation of the dilution effects, induced biochemical changes to overcome damage from Cd toxicity, and alteration of the endogenous phytohormones. FZT214 can find competent application in the future to improve the growth of other crop plants.

Effects of the Dark Septate Endophyte (DSE) Exophiala pisciphila on the Growth of Root Cell Wall Polysaccharides and the Cadmium Content of Zea mays L. under Cadmium Stress

This paper aims to investigate the mechanism by which dark septate endophytes (DSEs) enhance cadmium (Cd) tolerance in there host plants. Maize (Zea mays L.) was inoculated with a DSE, Exophiala pisciphila, under Cd stress at different concentrations (0, 5, 10, and 20 mg·kg−1). The results show that, under 20 mg/kg Cd stress, DSE significantly increased maize biomass and plant height, indicating that DSE colonization can be utilized to increase the Cd tolerance of host plants. More Cd was retained in DSE-inoculated roots, especially that fixed in the root cell wall (RCW). The capability of DSE to induce a higher Cd holding capacity in the RCW is caused by modulation of the total sugar and uronic acid of DSE-colonized RCW, mainly the pectin and hemicellulose fractions. The fourier-transform spectroscopy analysis results show that carboxyl, hydroxyl, and acidic groups are involved in Cd retention in the DSE-inoculated RCW. The promotion of the growth of maize and improvement in its tolerance to Cd due to DSEs are related to restriction of the translocation of Cd from roots to shoots; resistance of Cd uptake Cd inside cells; and the increase in RCW-integrated Cd through modulating RCW polysaccharide components.

Differences in Cadmium Accumulation, Detoxification and Antioxidant Defenses between Contrasting Maize Cultivars Implicate a Role of Superoxide Dismutase in Cd Tolerance

Cadmium (Cd), a readily absorbed and translocated toxic heavy metal, inhibits plant growth, interrupts metabolic homeostasis and induces oxidative damage. Responses towards Cd-stress differ among plant cultivars, and the complex integrated relationships between Cd accumulation, detoxification mechanisms and antioxidant defenses still need to be unraveled. To this end, 12 Egyptian maize cultivars were grown under Cd-stress to test their Cd-stress tolerance. Out of these cultivars, tolerant (TWC360 and TWC321), moderately sensitive (TWC324) and sensitive (SC128) cultivars were selected, and we determined their response to Cd in terms of biomass, Cd accumulation and antioxidant defense system. The reduction in biomass was highly obvious in sensitive cultivars, while TWC360 and TWC321 showed high Cd-tolerance. The cultivar TWC321 showed lower Cd uptake concurrently with an enhanced antioxidant defense system. Interestingly, TWC360 accumulated more Cd in the shoot, accompanied with increased Cd detoxification and sequestration. A principal component analysis revealed a clear separation between the sensitive and tolerant cultivars with significance of the antioxidant defenses, including superoxide dismutase (SOD). To confirm the involvement of SOD in Cd-tolerance, we studied the effect of Cd-stress on a transgenic maize line (TG) constitutively overexpressing AtFeSOD gene in comparison to its wild type (WT). Compared to their WT, the TG plants showed less Cd accumulation and improved growth, physiology, antioxidant and detoxification systems. These results demonstrate the role of SOD in determining Cd-tolerance.

Praseodymium enhanced the tolerance of maize seedlings subjected to cadmium stress by up-regulating the enzymes in the regeneration and biosynthetic pathways of ascorbate and glutathione

To test whether praseodymium (Pr) regulates cadmium (Cd) tolerance, we explored the effects of Pr on enzymatic activities in the regeneration and biosynthetic pathways of ascorbate and glutathione in maize seedlings under Cd stress. The findings demonstrated that Cd stress increased enzymatic activities in the regeneration pathway (ascorbate peroxidase (APX), dehydroascorbate reductase (DHAR), glutathione reductase (GR), monodehydroascorbate reductase (MDHAR)) and in the biosynthetic pathway of ascorbate and glutathione (γ-ECS and GalLDH), as well as ascorbate (AsA) and glutathione (GSH) contents. However, Cd stress significantly decreased AsA/dehydroascorbic acid (DHA) ratio and GSH/oxidised glutathione (GSSG) ratio, net photosynthetic rate (P_n), chlorophylls (Chl) and carotenoids (Car) contents, maximum photochemical efficiency of PSII (F_v/F_m), photochemical quenching (qP) and quantum efficiency of PSII photochemistry (Φ_PSII), as well as plant height and biomass. Application of Pr to Cd-stressed seedlings enhanced above enzymatic activities, AsA and GSH contents, AsA/DHA and GSH/GSSG ratios, P_n, Chl and Car contents, F_v/F_m, qP and Φ_PSII, as well as plant height and biomass. Meanwhile, the application of Pr to Cd-stressed seedlings reduced malondialdehyde (MDA) content and electrolyte leakage. The above results indicated that Pr enhanced Cd tolerance of maize by up-regulating enzymatic activities in regeneration and biosynthetic pathways of ascorbate and glutathione.

Melatonin Confers Plant Cadmium Tolerance: An Update

Cadmium (Cd) is one of the most injurious heavy metals, affecting plant growth and development. Melatonin (N-acetyl-5-methoxytryptamine) was discovered in plants in 1995, and it is since known to act as a multifunctional molecule to alleviate abiotic and biotic stresses, especially Cd stress. Endogenously triggered or exogenously applied melatonin re-establishes the redox homeostasis by the improvement of the antioxidant defense system. It can also affect the Cd transportation and sequestration by regulating the transcripts of genes related to the major metal transport system, as well as the increase in glutathione (GSH) and phytochelatins (PCs). Melatonin activates several downstream signals, such as nitric oxide (NO), hydrogen peroxide (H2O2), and salicylic acid (SA), which are required for plant Cd tolerance. Similar to the physiological functions of NO, hydrogen sulfide (H2S) is also involved in the abiotic stress-related processes in plants. Moreover, exogenous melatonin induces H2S generation in plants under salinity or heat stress. However, the involvement of H2S action in melatonin-induced Cd tolerance is still largely unknown. In this review, we summarize the progresses in various physiological and molecular mechanisms regulated by melatonin in plants under Cd stress. The complex interactions between melatonin and H2S in acquisition of Cd stress tolerance are also discussed.

Comparative root transcriptome analysis of two soybean cultivars with different cadmium sensitivities reveals the underlying tolerant mechanisms

The soybean can provide rich protein and fat and has great economic value worldwide. Cadmium (Cd) is a toxic heavy metal to organisms. It can accumulate in plants and be transmitted to the human body via food chain. Cd is a serious threat to soybean development, especially to root growth. Some soybean cultivars present tolerant symptoms under Cd stress; however, the potential mechanisms are not fully understood. Here, we optimized RNA-seq to identify the differentially expressed genes (DEGs) in Cd-sensitive (KUAI) and Cd-tolerant (KAIYU) soybean roots and compared the DEGs between KAIYU and KUAI. A total of 1,506 and 1,870 DEGs were identified in the roots of KUAI and KAIYU, respectively. Through Gene Ontology (GO), Kyoto Encyclopedia of Genes and Genomes (KEGG) pathway and gene function analyses, we found that genes related to antioxidants and sequestration were responsible for Cd tolerance in KAIYU. In addition, overexpression of Glyma11g02661, which encodes a heavy metal transporting ATPase, significantly improved Cd tolerance in transgenic hairy roots. These results provide a preliminary understanding of the tolerance mechanisms in response to Cd stress in soybean root development and are of great importance in developing Cd-resistant soybean cultivars by using the identified DEGs through genetic modification.

The Endophytic Fungus Piriformospora Indica-Assisted Alleviation of Cadmium in Tobacco

Increasing evidence suggests that the endophytic fungus Piriformospora indica helps plants overcome various abiotic stresses, especially heavy metals. However, the mechanism of heavy metal tolerance has not yet been elucidated. Here, the role of P. indica in alleviating cadmium (Cd) toxicities in tobacco was investigated. It was found that P. indica improved Cd tolerance to tobacco, increasing Cd accumulation in roots but decreasing Cd accumulation in leaves. The colonization of P. indica altered the subcellular repartition of Cd, increasing the Cd proportion in cell walls while reducing the Cd proportion in membrane/organelle and soluble fractions. During Cd stress, P. indica significantly enhanced the peroxidase (POD) activity and glutathione (GSH) content in tobacco. The spatial distribution of GSH was further visualized by Raman spectroscopy, showing that GSH was distributed in the cortex of P. indica-inoculated roots while in the epidermis of the control roots. A LC-MS/MS-based label-free quantitative technique evaluated the differential proteomics of P. indica treatment vs. control plants under Cd stress. The expressions of peroxidase, glutathione synthase, and photosynthesis-related proteins were significantly upregulated. This study provided extensive evidence for how P. indica enhances Cd tolerance in tobacco at physiological, cytological, and protein levels.