Photosynthetic variation and detoxification strategies based on cadmium uptake, non-protein thiols, and secondary metabolites in Miscanthus sacchariflorus under cadmium exposure.

Miscanthus sacchariflorus is previously demonstrated to be a potential candidate for remediation of cadmium (Cd) pollution. To explore its resistance strategy to Cd, a hydroponic experiment was conducted to determine the variations of photosynthetic activity in leaves and physiological response in roots of this plant. Results showed that the root of M. sacchariflorus was the primary location for Cd accumulation. The bioconcentration factor in the roots and rhizomes was >1, and the translocation factor from underground to aboveground was <1. Throughout the experimental period, treatment with 0.06 mM Cd2+ did not significantly alter the contents of chlorophyll a, chlorophyll b, or carotenoid. By contrast, treatment with 0.15 and 0.30 mM Cd2+ decreased the contents of chlorophyll a, chlorophyll b, and carotenoid; caused the deformation of the chlorophyll fluorescence transient curve; reduced the photochemical efficiency of photosystem II; and increased the contents of non-protein thiols, total flavone, and total phenol. These results indicate that M. sacchariflorus has good adaptability to 0.06 mM Cd2+. Moreover, the accumulation of the non-protein thiols, total flavone, and total phenol in roots may promote the chelation of Cd2+, thus alleviating Cd toxicity. This study provides theoretical support for using M. sacchariflorus to remediate Cd-polluted wetlands.

Role of methylglyoxal and glyoxalase in the regulation of plant response to heavy metal stress.

KEY MESSAGE: Methylglyoxal and glyoxalase function a significant role in plant response to heavy metal stress. We update and discuss the most recent developments of methylglyoxal and glyoxalase in regulating plant response to heavy metal stress. Methylglyoxal (MG), a by-product of several metabolic processes, is created by both enzymatic and non-enzymatic mechanisms. It plays an important role in plant growth and development, signal transduction, and response to heavy metal stress (HMS). Changes in MG content and glyoxalase (GLY) activity under HMS imply that they may be potential biomarkers of plant stress resistance. In this review, we summarize recent advances in research on the mechanisms of MG and GLY in the regulation of plant responses to HMS. It has been discovered that appropriate concentrations of MG assist plants in maintaining a balance between growth and development and survival defense, therefore shielding them from heavy metal harm. MG and GLY regulate plant physiological processes by remodeling cellular redox homeostasis, regulating stomatal movement, and crosstalking with other signaling molecules (including abscisic acid, gibberellic acid, jasmonic acid, cytokinin, salicylic acid, melatonin, ethylene, hydrogen sulfide, and nitric oxide). We also discuss the involvement of MG and GLY in the regulation of plant responses to HMS at the transcriptional, translational, and metabolic levels. Lastly, considering the current state of research, we present a perspective on the future direction of MG research to elucidate the MG anti-stress mechanism and offer a theoretical foundation and useful advice for the remediation of heavy metal-contaminated environments in the future.

An integrated transcriptome, metabolomic, and physiological investigation uncovered the underlying tolerance mechanisms of Monochoria korsakowii in response to acute/chronic cadmium exposure.

Identifying the physiological response and tolerance mechanism of wetland plants to heavy metal exposure can provide theoretical guidance for an early warning for acute metal pollution and metal-contaminated water phytoremediation. A hydroponic experiment was employed to investigate variations in the antioxidant enzyme activity, chlorophyll content, and photosynthesis in leaves of Monochoria korsakowii under 0.12 mM cadmium ion (Cd2+) acute (4 d) and chronic (21 d) exposure. Transcriptome and metabolome were analyzed to elucidate the underlying defensive strategies. The acute/chronic Cd2+ exposure decreased chlorophyll a and b contents, and disturbed photosynthesis in the leaves. The acute Cd2+ exposure increased catalase activity by 36.42%, while the chronic Cd2+ exposure markedly increased ascorbate peroxidase, superoxide dismutase, and glutathione peroxidase activities in the leaves. A total of 2 685 differentially expressed genes (DEGs) in the leaves were identified with the plants exposed to the acute/chronic Cd2+ contamination. In the acute Cd2+ exposure treatment, DEGs were preferentially enriched in the plant hormone transduction pathway, followed by phenylrpopanoid biosynthesis. However, the chronic Cd2+ exposure induced DEGs enriched in the biosynthesis of secondary metabolites pathway as priority. With acute/chronic Cd2+ exposure, a total of 157 and 227 differentially expressed metabolites were identified in the leaves. Conjoint transcriptome and metabolome analysis indicated the plant hormone signal transduction pathway and biosynthesis of secondary metabolites was preferentially activated by the acute and chronic Cd2+ exposure, respectively. The phenylpropanoid pathway functioned as a chemical defense, and the positive role of deoxyxylulose phosphate pathway in leaves against acute/chronic Cd2+ exposure was impaired.

[Phenylpropanoid pathway in plants and its role in response to heavy metal stress: a review].

Phenylpropanoid metabolic pathway is one of the most important secondary metabolic pathways in plants. It directly or indirectly plays an antioxidant role in plant resistance to heavy metal stress, and can improve the absorption and stress tolerance of plants to heavy metal ions. In this paper, the core reactions and key enzymes of the phenylpropanoid metabolic pathway were summarized, and the biosynthetic processes of key metabolites such as lignin, flavonoids and proanthocyanidins and relevant mechanisms were analyzed. Based on this, the mechanisms of key products of phenylpropanoid metabolic pathway in response to heavy metal stress were discussed. The perspectives on the involvement of phenylpropanoid metabolism in plant defense against heavy metal stress provides a theoretical basis for improving the phytoremediation efficiency of heavy metal polluted environment.

Photosynthetic response, antioxidase activity, and cadmium uptake and translocation in Monochoria korsakowii with cadmium exposure.

To identify the tolerance mechanisms of wetland plants exposed to heavy metal, a hydroponic experiment was used to investigate variations in photosynthetically physiological parameters and antioxidant enzyme activities in leaves of Monochoria korsakowii exposed to 0.05, 0.15, 0.30, and 0.45 mM Cd2+ for 7 d. The Cd2+ concentrations in the plant roots, stems, and leaves were also investigated. Cd2+ exposure significantly decreased the total chlorophyll content, net photosynthetic rate, intercellular carbon dioxide concentration, and stomatal conductance, while stomatal limitation value had the opposite trend (P < 0.05). During Cd2+ stress, ascorbate peroxidase activity significantly increased (P < 0.05). The translocation factor for Cd2+ was significantly lower than that of the control, and both were less than 1 (P < 0.05). Cd2+ stress damaged the photosynthetic apparatus in the leaves. During Cd2+ stress, M. korsakowii alleviated oxidative stress by increasing the activities of antioxidant enzymes, such as APX. Under 0.45 mM Cd2+ stress, increased heat dissipation was responsible for alleviating the photooxidative damage to photosynthetic organs in the leaves. Meanwhile, the majority of Cd2+ was immobilized in the roots, thus alleviating excessive Cd2+ phytotoxicity in the aboveground parts. Generally, M. korsakowii has potential application in the phytoremediation of low-cadmium-polluted water.

Colonization and phytoremediation potential for Miscanthus sacchariflorus in copper tailings.

A pot experiment was conducted to explore the effects of copper (Cu) tailings with various proportions in the substrate on seed germination and morphological traits of the plant. Concurrently, to identify the adaptive and tolerance strategies of the plant to Cu tailings, the uptake and accumulation of the plant to heavy metals, variations in soil enzymatic activities, and metal speciation in the blank, rhizospheric, and non-rhizospheric soils were estimated. Cu tailings at 25% proportion in the substrate exerted no significant negative effects on seed germination and seedling growth. However, Cu tailings at higher proportions (≧50%) inhibited seed germination and disturbed the plant physiological metabolism and growth. More biomass allocated to the plant roots could contribute to more heavy metals being immobilized, arresting their translocation from roots to shoots. This was accepted as a crucial defense mechanism for the plant against heavy metal co-contamination. The plant can improve the biological properties of Cu tailings in terms of enhanced invertase and phosphatase activities. And in turn, this can effectively alleviate heavy metal phytotoxicity. Simultaneously, it markedly decreased exchangeable Cu and Zn content in the rhizosphere in 25% Cu tailings treatments. In 50% Cu tailings treatment, no differences were observed in Zn speciation between rhizosphere and non-rhizosphere soils. In 75% Cu tailings, compared to the non-rhizosphere, an obvious reduction in exchangeable Cu in rhizosphere occurred, while an opposite tendency was demonstrated in carbonate-bound Zn. The plant could successfully colonize in Cu tailings, and represent a phytoremediation potential in Cu tailings.

Physiological defense and metabolic strategy of Pistia stratiotes in response to zinc-cadmium co-pollution.

Pistia stratiotes is a cadmium (Cd) hyperaccumulating plant with strong bioaccumulation and translocation capacity for Cd. A hydroponic experiment was used to evaluate the combined effect of Zinc (Zn) and Cd at different concentrations on leaf growth and metabolism of P. stratiotes. This study revealed the physiological defense and metabolic strategy of responses to Zn-Cd co-pollution. With the Zn50Cd1, Zn50Cd10, Zn100Cd1, and Zn100Cd10 treatments for 9 d, the relative crown diameter, relative leave number, and ramet number of the plant had no significant difference with the control. Under the compound treatments containing Zn50Cd50 and Zn100Cd50, the activity of the glyoxalase system and amino acid metabolism in the leaves were inhibited. The leaf photosynthetic apparatus increased heat dissipation to reduce the damage to the photosystem II (PS II) reaction center caused by excess excitation energy under Zn-Cd stress. This safeguarded the balance between the absorption and utilization of light energy. Compared to the control, the Zn and Cd co-pollution for 9 d had no effect on the reduced glutathione (GSH) and oxidized glutathione (GSSG) contents. There was no effect on the dehydroascorbate reductase (DHAR) and glutathione reductase (GR) activities, but there was increased ascorbate peroxidase (APX) activity and oxidized ascorbic acid (DHA) content. These increased the antioxidant capacity of the ascorbate-glutathione (AsA-GSH) cycle. The treated plants also had increased levels of carnosol and substances related to lipid metabolism including 9, 10-Dihydroxystearate, Prostaglandin G2, Sphingosine, and 13-L-Hydroperoxylinoleic acid, maintaining the cell stability and resistance to the Zn-Cd stress.

Transcriptional, secondary metabolic, and antioxidative investigations elucidate the rapid response mechanism of Pontederia cordata to cadmium.

Pontederia cordata is previously demonstrated a cadmium (Cd) tolerant plant, and also a candidate for the phytoremediation of heavy-metal-contaminated wetlands. A hydroponic experiment was used to investigate variations in photosynthetic gas exchange parameters, antioxidative activities, chlorophyll and secondary metabolite contents, and transcriptome in leaves of the plant exposed to 0.44 mM Cd2+ for 0 h, 24 h, and 48 h. Under Cd2+ exposure for 24 h, the plant presented a favorable photosynthesis by maintaining relatively higher antioxidant activity. Cd2+ exposure for 48 h accelerated membrane peroxidation, declined photosynthetic pigment content, and increased polyphenol oxidase activity, thus interfering with photosynthesis. The phenylpropane pathway served as a chemical rather than physical defense against Cd2+ in the plant leaves. A total of 20,998, 4743, and 4413 differentially expressed genes (DEGs) were identified in the groups of 0 h vs 24 h, 0 h vs 48 h, and 24 h vs 48 h, respectively. The primary metabolic pathways of the DEGs were mainly enriched in nitrogen metabolism, starch and sucrose metabolism, fructose and mannose metabolism, as well as pentose-phosphate pathway, contributing to a stable cell structure and function. Flavonoid biosynthesis directly or indirectly played an antioxidative role against Cd2+ in the leaves. Forty-nine transcription factor (TF) families were identified, and 8 TF families were shared among the three groups. The present study provides a theoretical foundation for investigating tolerance mechanisms of wetland plants to Cd stress in terms of secondary metabolism and transcriptional regulation.

Tolerance mechanism and phytoremediation potential of Pistia stratiotes to zinc and cadmium co-contamination.

Pistia stratiotes can not only effectively remediate eutrophic water, but also displays strong absorption and bioaccumulation abilities for heavy metals. However, it has not been well-understood how the plant resists the combined stress of heavy metals. In these experiments, the morphophysiological traits, the ascorbate-glutathione (AsA-GSH) cycle, the glyoxalase system, and the contents of zinc (Zn) and cadmium (Cd) were investigated under Zn and Cd co-pollution. The AsA-GSH cycle and glyoxalase system could coordinately alleviate the oxidative and carbonyl stress, which was identified as an important tolerance mechanism. With Zn50Cd1, Zn50Cd10, Zn100Cd1, and Zn100Cd10 treatments for 18 days, 90.75-93.69% of Zn and 88.13-96.96% Cd accumulated in the roots. Treatments with Zn50Cd50, and Zn100Cd50 for 18 days resulted in a decrease of stress tolerance and chlorophyll content in leaves, an increase in plasma membrane permeability, a massive accumulation of methylglyoxal (MG), and visible toxic symptoms. Additionally, the bioaccumulation factor (BCF) for roots and shoots and the translocation factor (TF) were >1, and the content of Cd in shoots was no <100 mg·kg-1. This indicated P. stratiotes was a Cd hyperaccumulator and have great potential for the phytoremediation of heavy metal contaminated water.Novelty statement Pistia stratiotes, a cadmium hyperaccumulator, has great application potential for the phytoremediation of zinc and cadmium co-polluted water.

Succinic acid inhibits photosynthesis of Microcystis aeruginosa via damaging PSII oxygen-evolving complex and reaction center.

To elucidate the mechanism of succinic acid (SA) inhibition of Microcystis aeruginosa, the chlorophyll fluorescence transients, photosynthesis, photosynthetic electron transport activity, and gene expression of M. aeruginosa were evaluated under various doses of SA. The results demonstrated that, after treatment with 60 mg L-1 SA for 1 h, the chlorophyll fluorescence transients and related parameters changed significantly, indicating that the function and structure of photosynthetic apparatuses of Microcystis were seriously damaged. The initial quantum efficiency α, maximum net photosynthetic rate Pnmax, dark respiration rate Rd, and gross photosynthetic rate decreased to 57%, 49%, 49%, and 46%, respectively, relative to the control. Furthermore, photosystem II (PSII) activity (H2O→p-BQ) and the electron transport activity of H2O→MV and DPC→MV significantly decreased. Real-time PCR analysis revealed that, following incubation with 60 mg L-1 SA for 24 h, the expression level of core protein genes (psbA, psaB, psbD, and psbO) of the photosynthesis centers photosystem I (PSI) and PSII decreased significantly. However, the transcription of gene nblA encoding phycobilisome degradation protein was elevated. The downregulation of the rbcL gene, which encodes the large subunit of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), resulted in the suppression of CO2 fixation and assimilation. High concentration (60 mg L-1) of SA resulted in damage to oxygen-evolving complex (OEC) and reaction center of PSII, blocking photosynthetic electron transport, thereby lowering the rate photosynthesis and inhibiting the growth of Microcystis. We concluded that inhibition of photosynthesis is an important mechanism of SA inhibition in M. aeruginosa.

Pontederia cordata, an ornamental aquatic macrophyte with great potential in phytoremediation of heavy-metal-contaminated wetlands.

Pontederia cordata can tolerate heavy metal toxicity and possesses great potential for phytoremediation of heavy-metal-contaminated wetlands, yet how it copes with heavy metal stress has still not been determined. Hydroponic experiments were used to assess the effects of various levels of Cd2+ on the photosynthesis and activity of redox-regulatory systems in the plant leaves, and we also sought to elucidate the tolerance mechanism of the plant to Cd2+ by investigating Cd2+ enrichment characteristics and chemical forms. The plant can manage a low cadmium concentration (≤0.04 mM) with relatively stable biomass and photosynthetic performance. Cd2+ at the highest concentration (0.44 mM) decreased superoxide dismutase and peroxidase activities by 37.17% and 93.29%, respectively. Similar trends were demonstrated in the contents of ascorbic acid, carotenoids, lutein, glutathione, and non-protein thiol, as well as phytochelation in the leaves, exacerbating membrane peroxidation despite the significantly increased catalase activity observed. Moreover, the highest Cd2+ concentration disturbed the biosynthesis of chlorophyll precursors in the leaves, reduced chlorophyll a and b, as well as total chlorophyll contents by 60.47%, 67.47%, and 68.12%, respectively, which inhibited photosynthesis, leading to a decline in biomass. Compared with maximum quantum efficiency (Fv/Fm) and the potential activity (Fv/Fo) of photosystem II, the performance index for energy conservation from photons absorbed by PSII to the reduction of intersystem electron acceptors (PIabs), and of PSI end acceptors (PItotal), can indicate Cd2+ toxicity to the photosynthetic apparatus in the leaves. 49.95%-76.90% of the Cd2+ was sequestered in the plant roots, restraining translocation from roots to shoots, which is considered a tolerance mechanism, probably resulting from disturbed transpiration in leaves and increased Cd2+ content with low activity. Pontederia cordata is a candidate plant for phytoremediation of heavy-metal -contaminated wetlands.

Cadmium phytotoxicity, related physiological changes in Pontederia cordata: antioxidative, osmoregulatory substances, phytochelatins, photosynthesis, and chlorophyll fluorescence.

Pontederia cordata is a heavy metal accumulator, while the heavy metal tolerance mechanisms of this plant are not well understood. Hydroponic experiments were used to assess the effects of Cd2+ on antioxidative activities, osmoregulatory substances and photosynthesis in leaves. Exposure of 5 mg L-1 Cd2+ for 7 days, the photosynthetic apparatus functioned normally and sustained a relatively high photosynthetic rate, and good growth was observed. Under 50 and 75 mg L-1 Cd2+, accelerated lipid peroxidation and increased peroxidase activity (POD; E.C.1.11.1.7) were detected, while no significant differences were observed in superoxide dismutase (SOD; E.C.1.15.1.1) and catalase (CAT; E.C.1.11.1.6) activities, as well as in lutein, ascorbic acid, and glutathione contains of leaves. Proline content increased, while soluble sugar and soluble protein contents decreased under 75 mg L-1 Cd2+. Cd2+ at different concentrations induced a reduction in carotenoid, total carotenoid, and ascorbic acid-dehydroascorbate contents. A significant increase in phytochelatin content was induced by 75 mg L-1. Chlorophyll content decreased under Cd stress and disturbed photosynthesis, causing dramatic reductions in photosynthetic parameters. Stomatal closure was responsible for a reduced photosynthetic rate under Cd2+ exposure. Cd2+ concentrations of no less than 25 mg L-1 disorganized the photosynthetic apparatus, induced the partial closure, and decreased activity of the photosystem II (PS II) reaction center, thus disturbing light conversion and utilization, thereby decreasing the photosynthetic efficiency in PS II.

Pre-aeration of the rhizosphere offers potential for phytoremediation of heavy metal-contaminated wetlands.

Two solution cultures with different oxygen pretreatments were used to investigate (ⅰ) the variation in the radial oxygen loss in the roots and root morphology of Triarrhena sacchariflora seedlings and (ii) their tolerance to Cu2+ and Cd2+, as well as both the metal uptake and accumulation by pretreated seedlings. Developed aerenchyma in the roots was induced by the hypoxia pretreatment (HP) and aeration pretreatment (AP), for which root porosity, respectively, increased by 45.76%-53.39% and 84.07%-88.66%. AP altered the natural radial oxygen loss coupled to an enhanced secretion of oxygen in the root tips. AP was found to effectively improve the seedlings' tolerance to Cu2+ and Cd2+, facilitating their growth, thereby increasing their root diameter, dry weight, and number of root tips, as well as promoting shoot growth. AP was capable of promoting the uptake and bioaccumulation in seedlings of Cu2+ and Cd2+; it also induced more Cu2+ and Cd2+ immobilized in roots so that less of either metal was transported from roots to shoots, which may well be a key mechanism for strengthening seedlings' tolerance to metal ions. Our experimental results suggest that AP offers great potential for the remediation of heavy metal-contaminated wetlands.

Tolerance mechanism of Triarrhena sacchariflora (Maxim.) Nakai. seedlings to lead and cadmium: Translocation, subcellular distribution, chemical forms and variations in leaf ultrastructure.

Hydroponic experiments were conducted to assess the accumulation, translocation, and chemical forms of lead (Pb) and cadmium (Cd) in the roots, stems, and leaves of Triarrhena sacchariflora seedlings and the associated variation in leaf ultrastructure. The leaves and leaf ultrastructure showed no significant symptoms of toxicity with 0.05 mM Pb or 0.01 mM Cd exposure for 10d. Chlorosis and wilting were observed in leaves when the Pb and Cd concentration was higher than 0.1 and 0.05 mM in the medium, respectively, as demonstrated by severe ultrastructural modifications at higher concentration in the leaves, such as plasmolysis, cell wall detachment, chloroplast swelling, nuclear condensation, and even nuclear fragmentation. The Pb and Cd concentrations in the roots was significantly higher than those in the stems and leaves. This indicated low Pb and Cd translocation from the roots to the aboveground parts. Subcellular distribution analysis showed that the majority of Pb and Cd was bound to the cell wall, especially in the roots, indicating that the cell wall likely constitutes a crucial storage site for Pb and Cd. This mechanism decreases the translocation of Pb and Cd across membranes and is more effective than vacuolar compartmentation. The majority of Pb and Cd exited in form of insoluble Pb/Cd-pectate or -oxalate complexes in the plant. In conclusion, higher concentrations of Pb or Cd induced premature senescence. High Pb and Cd enrichment was observed in the roots, which decreased the translocation of Pb and Cd from the roots to the aboveground tissues. The immobilization of Pb or Cd by the cell wall is important for plant detoxification and can protect protoplasts from Pb or Cd toxicity. Pb and Cd mainly existed in insoluble Pb/Cd-phosphate or -oxalate complexes, exhibiting low activity and thereby limiting symplastic transport and suppressing toxicity.