Drought (D) and chromium (Cr) stress co-occur in agricultural fields due to the accumulation of excessive Cr in soils from industrial pollution and increasing frequency of water scarcity. Carrageenan (Car), a compound extracted from red seaweed, is an emerging biostimulant with multifaceted roles in plants. This study investigated the role of exogenous Car in mediating tolerance to D-, Cr-, and DCr-stress in wheat seedlings, aiming to elucidate the potential of Car in mitigating toxicity and promoting plant resilience. Wheat seedlings exposed to DCr-stress exhibited reduced growth and biomass production, along with elevated levels of reactive oxygen, carbonyl, and nitrogen species. Moreover, D-stress exacerbated Cr-toxicity, as demonstrated by principal component analysis (PCA), which showed a strong positive correlation between DCr-stress and stress marker parameters. This suggests that DCr-stress resulted in higher Cr uptake and increased oxidative damage compared to individual D-or Cr-stress, making DCr-stress more detrimental than either stress applied alone. However, Car priming ameliorated the toxic effects of DCr-stress and promoted the growth performance of DCr-stressed wheat seedlings. In PCA, the positive correlation of D + Car, Cr + Car, and DCr + Car treatments with growth and plant defense-related parameters suggests that Car-mediated improvement in stress tolerance can be attributed to reduced accumulation of toxic Cr, increased levels of total free amino acids and soluble sugars, enhanced antioxidant enzyme activity, elevated non-enzymatic antioxidant levels, higher phenolic and flavonoid content, and improved metal chelation and detoxification. Our results indicated Car is a potential and cost-effective biostimulant for managing D-, Cr-, or DCr-stress in wheat.

Excessive accumulation of cadmium (Cd) impairs crop growth by inducing oxidative damage through the generation of reactive oxygen species (ROS). In this study, a biocompatible ferruginated carbon quantum dots (Fe-CQDs) nanozyme is developed to target ROS, thereby reducing oxidative damage and improving the absorption and transfer of Cd ions in wheat. Notably, Fe-CQDs exhibit multi-enzyme activities mimicking peroxidase (POD), catalase (CAT), and superoxide dismutase (SOD), enabling effective neutralization of active species such as hydroxyl radicals (center dot OH), hydrogen peroxide (H2O2), and superoxide anions (O2 center dot-). Importantly, root application of 10 mg L-1 Fe-CQDs alleviates Cd stress and promotes wheat growth in both hydroponic and soil cultures. Specifically, the levels of O2 center dot-, H2O2, and malondialdehyde (MDA) in leaf tissues decrease, whereas the non-enzyme antioxidant, reduced glutathione (GSH), increases. Cell wall thickness in the Fe-CQDs-treated group is reduced by 42.4% compared with the Cd group. Moreover, Fe-CQDs enhance the expression of genes related to antioxidants, stress resistance, Cd detoxification, and nutrient transport. Transcriptomic and metabolomic analyses show that Fe-CQDs stimulate the production of flavonoids and regulate the activity of metal transporter genes (YSL, ABC, ZIP) to maintain ROS homeostasis. These findings highlight the potential of Fe-CQDs nanozyme platforms in mitigating oxidative damage and enhancing crop growth, offering new insights into the application of nanobiotechnology in agriculture. (c) 2025 Crop Science Society of China and Institute of Crop Science, CAAS. Production and hosting by Elsevier B.V. on behalf of KeAi Communications Co., Ltd. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

Key messageMelatonin increases Pb tolerance in P. ovata seedlings via the regulation of growth and stress-related phytohormones, ROS scavenging and genes responsible for melatonin synthesis, metal chelation, and stress defense.AbstractLead (Pb) is a highly toxic heavy metal that accumulates in plants through soil and air contamination and impairs its plant growth and development. Because of its pharmaceutical importance, improvements in Plantago ovata yield against abiotic stresses are necessary. Melatonin (MEL) is a stress-alleviating biostimulator and our results showed a reduction in Pb induced phytotoxicity by enhancing plant growth attributes and balancing protective osmolytes. Pb-induced reactive oxygen species accumulation, including superoxide and peroxide free radicals and their mitigation through enzymatic antioxidants, was demonstrated in presence of MEL. Cell viability and Pb bioaccumulation were determined to understand the extent of cellular damage. Moreover, MEL increased secondary metabolite (flavonoids and anthocyanins) contents by 2-3-fold at the lowest Pb concentrations. Similar increases in the relative expression of genes (PoPAL and PoPPO), which are responsible for the production of non-enzymatic antioxidants, were observed. Notably, the upregulation of the PoCOMT gene up to 4-fold indicates increased melatonin production, as manifested in the phytomelatonin level. MEL supplementation also increased the auxin (IAA) level by 3-fold in the 100 mu M Pb treatment group, while the abscisic acid (ABA) level decreased (1.4-fold) and the expression of PoMYB (a stress-related transcription factor) increased (up to 2.66-fold). Additionally, we found extreme downregulation (up to 18-fold) in the relative expression of PoMT 2 (a metal binding thiol compound) with melatonin treatment, which is otherwise upregulated (by 6-fold) during Pb stress. In the current study, these effects collectively revealed that MEL contribute to enhanced plant growth and Pb stress tolerance.

The application of 2,4-epibrassinolide (EBR) is considered an effective and environment friendly method to improve plant growth under heavy metal (HM) stress, which is crucial for crop productivity and environmental phytoremediation. This meta-analysis evaluated plant responses to exogenous EBR under HM stress by compiling data from 73 studies, including 2480 observations. Results showed that the most significant effects of exogenous EBR on plant growth and HM uptake parameters were observed on shoot/root length (47.9 %) and HM concentration in plant tissues (-32.9 %). EBR application enhanced photosynthesis and the mitigation of oxidative damage by significantly boosting antioxidant enzyme activity, non-enzymatic antioxidants, and metabolites. Exogenous EBR induced the largest percentage changes in plant growth and HM uptake under nickel stress, with an average increase of 57.5 % and a decline of 38.5 %, respectively. The greatest effects of exogenous EBR on plant growth and HM uptake parameters were observed in plants of the Cruciferae family, while the lowest effects were in the Gramineae family. In terms of EBR application characteristics, seed soaking with lower EBR concentrations (<= 1 nM) is recommended for crop production in HM-contaminated soils. These findings underscore the potential of exogenous EBR in achieving sustainable agriculture and environmental phytoremediation in HM-contaminated soils.

Heavy metal contamination, particularly from cadmium (Cd) and lead (Pb), poses significant risks to soil and water resources and leads to severe damage in plants. This study investigated the physiological and molecular mechanisms of the responses of tomato (Solanum lycopersicum L.) seedlings to Cd and Pb stress by applying 50 mg/L Cd, 100 mg/L Pb, and a combination of 50 mg/L Cd + 100 mg/L Pb. The goal was to understand how these heavy metals impact the growth, antioxidant systems, and secondary metabolic pathways in tomato seedlings. The results showed that compared with the control, Cd + Pb stress significantly increased the content of soluble sugar by 37.40% and 33.46% on days 5 and 15, respectively, and the content of proline by 77.91% to 93.91% during the entire period in tomato seedlings. It also elevated electrical leakage by 110.52% on day 15, maintained the levels of malondialdehyde close to the control, enhanced the activities of superoxide dismutase by 33.32% on day 10 and 11.22% on day 15, peroxidase by 42.15% on day 10, and catalase by 90.78% on day 10. Additionally, it reduced the contents of hydrogen peroxide by 15.47% to 29.64% and the rate of formation of superoxide anions by 26.34% to 53.47% during the entire period of treatment. The transcriptomic analysis revealed a significant differential expression of the genes involved in pathways, such as phenylalanine, glutathione, arginine and proline, and nitrogen metabolism. These genes included PALs, HDCs, GGCT, ODC1, LAPs, SMS, and SAMDC. Notably, transcription factors, such as ERF109, ARF9, GRF3, GRF4, GRF7, and GRF9, were also significantly regulated. The study concluded that Cd and Pb stress enhanced the osmoregulatory and antioxidant defense systems in tomato seedlings, which may contribute to their tolerance to heavy metal stress. Future research could explore the application of these findings to develop strategies to improve the resistance of plants to contamination with heavy metals.

Heavy metal stress can lead to morphological and physiological variations in crops. We aimed to distinguish heavy metal stress levels based on the variations of morphological and physiological parameters from radiative transfer and statistical models. Sentinel-2 satellite images and in situ measured data were collected from heavy metal-contaminated soils of rice growing areas in Zhuzhou City, Hunan Province, China. The chlorophyll content (chlorophyll a + chlorophyll b, Cab) and leaf area index (LAI) were calculated using a PROSAIL radiative transfer model and the multilayer perceptron algorithm. A two-dimensional feature space was established from Cab-LAI. Furthermore, a normalized heavy metal stress index (HMSI) from the established Cab-LAI theoretical triangular model was explored to distinguish heavy metal stress levels in rice. The results indicated that (i) the PROSAIL and artificial neural network algorithm were successful at deriving physiological parameters with high estimation accuracy. Pearson's correlation coefficient between the predicted and measured Cab was 0.85; (ii) the correlation between the measured concentration of cadmium in the soil and the HMSI was 0.84, indicating that it is a good indicator of rice damage caused by heavy metal stress, with the maximum HMSI occurring in rice subjected to high pollution; and (iii) high pollution occurred on both sides of the Xiangjiang River, whereas moderate pollution mainly existed around the heavily polluted areas. Areas with non-pollution and mild pollution were distributed over most of the study area. Combining rice Cab with LAI is a feasible method to determine the distribution of rice heavy metal stress levels over a large area.

Although much interest has been focused on the role of selenium (Se) in plant nutrition over the last 20 years, the influences of organic selenium (selenomethionine; Se-Met) and inorganic selenium (potassium selenite; Se-K) on the growth and physiological characters of cadmium (Cd)-stressed Glycine max L.) seedlings have not yet been studied. In this study, the impacts of Se-Met or Se-K on the growth, water physiological parameters (gaseous exchange and leaf water content), photosynthetic and antioxidant capacities, and hormonal balance of G. max seedlings grown under 1.0 mM Cd stress were studied. The results showed that 30 mu M Se-K up-regulates water physiological parameters, photosynthetic indices, antioxidant systems, enzymatic gene expression, total antioxidant activity (TAA), and hormonal balance. In addition, it down-regulates levels of reactive oxygen species (ROS; superoxide free radicals and hydrogen peroxide), oxidative damage (malondialdehyde content as an indicator of lipid peroxidation and electrolyte leakage), Cd translocation factor, and Cd content of Cd-stressed G. max seedlings. These positive findings were in favor of seedling growth and development under Cd stress. However, 50 mu M Se-Met was more efficient than 30 mu M Se-K in promoting the above-mentioned parameters of Cd-stressed G. max seedlings. From the current results, we conclude Se-Met could represent a promising strategy to contribute to the development and sustainability of crop production on soils contaminated with Cd at a concentration of up to 1.0 mM. However, further work is warranted to better understand the precise mechanisms of Se-Met action under Cd stress conditions.

Simple Summary: Zinc is a vital nutrient required by all living organisms; however, its impact varies based on Zn concentration and chemical form. This study examined the effect of zinc chloride (ZnCl2) and zinc sulfate (ZnSO4) on the life history performance and hemolymph metabolism of the common moth, Spodoptera litura, which is known to damage many crops. We found that, while low levels of ZnCl2 benefit the reproduction of Spodoptera litura, higher levels of ZnCl2 prolong the preadult developmental period and decrease the preadult survival rate. Additionally, dietary ZnSO4 exerts a devastating effect on the survival of S. litura larvae, even at the lowest concentration. This helps us better understand the effect of the chemical forms and concentrations of zinc on the biological processes and toxicological impacts on insects. Zinc is an essential micronutrient crucial in various biological processes of an organism. However, the effects of zinc vary depending on its chemical form. Therefore, the aim of this study was to conduct a comparative analysis of the life history performances and hemolymph metabolism of Spodoptera litura exposed to different concentrations of dietary zinc chloride (ZnCl2) and zinc sulfate (ZnSO4), utilizing two-sex life tables and untargeted metabolomics. The preadult survival rate of S. litura significantly decreased, while the preadult developmental period of S. litura was prolonged as the dietary ZnCl2 concentration increased. However, the fecundity of S. litura at 50 mg/kg dietary ZnCl2 was significantly increased. The intrinsic rate of increase (r) and the finite rate of increase (lambda) in S. litura in the control group (CK, no exogenous ZnCl2 or ZnSO4 added) and with 50 mg/kg dietary ZnCl2 were significantly higher than those at 100 mg/kg, 200 mg/kg, and 300 mg/kg. Dietary ZnSO4 exerts a devastating effect on the survival of S. litura. Even at the lowest concentration of 50 mg/kg dietary ZnSO4, only 1% of S. litura could complete the entire life cycle. Furthermore, as the dietary ZnSO4 concentration increased, the developmental stage achievable by the S. litura larvae declined. High-throughput untargeted metabolomics demonstrated that both 100 mg/kg dietary ZnCl2 and ZnSO4 decreased the hemolymph vitamins levels and increased the vitamin C content, thereby helping S. litura larvae to counteract the stress induced by ZnCl2 and ZnSO4. Simultaneously, dietary ZnCl2 obstructed the chitin synthesis pathway in the hemolymph of S. litura, thus extending the developmental period of S. litura larvae. These results indicate that low concentrations of Zn2+ positively impact populations of S. litura, but the effectiveness and toxicity of Zn depend on its chemical form and concentration.

With the growth of the population and the development of modern industry and the economy, the problem of heavy metal pollution in cultivated soil has become increasingly prominent. Moreover, heavy metal poses a serious threat to plant growth due to its characteristics of difficult degradation, high mobility, easy enrichment, and potential toxicity and has become a social topic. Melatonin is a new type of plant hormone widely present in animals, plants, fungi, and bacteria, and its biological role has begun investigated in the last dozen years. Facing heavy metal stress, melatonin can play a pleiotropic role in the physiological processes of plants, such as stress resistance and growth regulation, mitigate the damage caused by stress on plants, and provide a new research idea for alleviating heavy metal stress in plants. From the aspects of the plant phenotype, physiology, element absorption, and molecular structure, this paper, therefore, mainly reviews the effects of melatonin on plants subjected to heavy metal stress and the mechanism of melatonin alleviating heavy metal stress and then puts forward future research directions. This information may be of great significance to the normal growth of crops under heavy metal stress and will provide an important theoretical basis for the genetic improvement of crop resistance in the future.

In this study, wild barley (Hordeum brevisubulatum) infected (E+) and uninfected (E-) by Epichlo & euml; bromicola were used for hydroponic experiments during the seedling stage. Various attributes, such as the effect of fungal endophyte on the growth and development of wild barley, the absorption of cadmium (Cd) and mineral elements (Ca, Mg, Fe, Mn, Cu, Zn), subcellular distribution, and chemical forms were investigated under CdCl2 stress. The results showed that the fungal endophy significantly reduced the Ca content and percentage of plant roots under Cd stress. The Fe and Mn content of roots, the mineral element content of soluble fractions, and the stems in the pectin acid or protein-chelated state increased significantly in response to fungal endophy. Epichlo & euml; endophyte helped Cd2+ to enter into plants; and reduced the positive correlation of Ca-Fe and Ca-Mn in roots. In addition, it also decreased the correlation of soluble components Cd-Cu, Cd-Ca, Cd-Mg in roots, and the negative correlation between pectin acid or protein-chelated Cd in stems and mineral elements, to increase the absorbance of host for mineral elements. In conclusion, fungal endophy regulated the concentration and distribution of mineral elements, while storing more Cd2+ to resist the damage caused by Cd stress. The study could provide a ground for revealing the Cd tolerance mechanism of endophytic fungal symbionts. NOVELTY STATEMENT The present study is the first to study the effect of fungal endophy on essential mineral elements of plants under heavy metal stress, filling a gap in the existing research. The study could be helpful to reveal the mechanism of endophytic fungi to improve the host's tolerance to heavy metals and provide a foundation for the grass-endophyte symbionts to improve heavy metal-contaminated soils as ecological grasses.