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Increased anthropogenic activities over the last decades have led to a gradual increase in cadmium content in the soil, which, due to its high mobility in soil, makes Cd accumulation in plants a serious threat to the health of animals and humans. Plant hormones including melatonin (Mel) and brassinosteroids (BR) are known to provide tolerance against various abiotic stresses. In this work, the role of combined and separate exogenous application of Mel and BR on Cd stress in cherry tomato plants was examined. Cd stress significantly reduced tomato growth by inducing oxidative stress and reduced K + uptake in roots and shoots. Combined application of Mel and BR reduced detrimental effects of Cd in tomato by (i) reducing Cd accumulation in the shoot; (ii) increasing the activities of different antioxidants (SOD, CAT, APX, GR); (iii) triggering higher expression of genes relating to Cd vacuolar sequestration (Na + /H + EXCHANGER, SlNHX1; NATURAL RESISTANCE-ASSOCIATED MACROPHAGE PROTEIN 6, SlNRAMP6), and Cd transport and detoxification (HEAVY-METAL-ASSOCIATED 3, SlHMA3; PLANT CADMIUM RESISTANT 2, SlPCR2); and (iv) improving plant K + homeostasis and contents in root and shoot. The latter trait was associated with the reduced gene expression of K + -permeable outward rectifying channel (SlGORK3), and transcriptional upregulation of high affinity potassium transporter 5 (SIHAK5) under Cd stress. A separate application of Mel and BR showed tissue-specific regulation of tomato growth and Cd tolerance by regulating antioxidant activities, K + uptake, Cd uptake, and translocation from root to shoot and their endogenous contents. Melatonin per se was more effective in improving Cd tolerance in shoot while beneficial BR effects were more pronounced in roots, and their combined application was effective in both tissues. Taken together, reported results show tissue-specific regulation of Cd tolerance by Mel and BR in cherry tomato plants and demonstrate the efficiency of combined Mel + BR treatment as a practical tool to reduce Cd accumulation and mitigate its negative effects on plant growth., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Inc. All rights reserved.)

Mini Chinese cabbage (Brassica rapa L. ssp. Pekinensis) plays an important role in the supply of summer vegetables on the plateau in western China. In recent years, tip-burn has seriously affected the yield, quality and commodity value of mini Chinese cabbage. Calcium (Ca 2+ ) deficiency is a key inducer of tip-burn. As a new type plant hormone, brassinolide (BR) is involved in regulating a variety of biotic and abiotic stresses. To explore the alleviation role of BR in tip-burn caused by Ca 2+ deficiency, a hydroponic experiment was conducted to study the relationship between BR and Ca 2+ absorption and transport. The results showed that foliar spraying with 0.5 µM BR significantly reduced tip-burn incidence rate and disease index of mini Chinese cabbage caused by Ca 2+ deficiency. Moreover, the dynamic monitoring results of tip-burn incidence rate showed that the value reached the highest on the ninth day after treatment. BR promoted the Ca 2+ transport from roots to shoots and from outer leaves to inner leaves by increasing the activities of Ca 2+ -ATPase and H + -ATPase as well as the total ATP content, which provided power for Ca 2+ transport. In addition, exogenous BR upregulated the relative expression levels of BrACA4, BrACA11, BrECA1, BrECA3, BrECA4, BrCAX1, BrCAS and BrCRT2, whereas Ca 2+ deficiency induced down-regulation. In conclusion, exogenous BR can alleviate the Ca 2+ -deficiency induced tip-burn of mini Chinese cabbage by promoting the transport and distribution of Ca 2+ ., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023. Published by Elsevier Inc.)

The application of phytohormones is a viable technique to increase the efficiency of phytoremediation in heavy metal-contaminated soils. The objective of this study was to determine how the application of 24-epibrassinolide (EBR), a brassinosteroid analog, could regulate root growth and tolerance to cadmium (Cd) stress in Kentucky bluegrass. As a result, the number of lateral root primordia and total root length in the Cd-treated seedlings decreased by 33.1 % and 56.5 %, respectively. After the application of EBR, Cd accumulation in roots and leaves, and the negative effect of Cd on root growth were reduced under Cd stress. Additionally, the expression of the brassinosteroid signaling gene PpBRI1 was significantly upregulated by exogenous EBR. Moreover, exogenous EBR upregulated the expression of genes encoding antioxidant enzymes and improved the activity of antioxidant enzymes, thereby reduced oxidative stress in roots. Finally, targeted hormonomics analysis highlighted the utility of the application of EBR to alleviate the effect of Cd on the reduction in auxin (IAA) content and the increase in ethylene (ACC) content. These were known to be associated with the upregulation in the expression of auxin biosynthesis gene PpYUCCA1 and downregulation in the expression of ethylene biosynthesis gene PpACO1 in the roots treated with Cd stress. Overall, the application of EBR alleviated Cd-induced oxidative stress in addition to improving root elongation and lateral root growth crosstalk with auxin and ethylene in Kentucky bluegrass subjected to Cd stress. This study further highlights the potential role of brassinosteroids in improving the efficiency of phytoremediation for Cd-contaminated soils., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 The Authors. Published by Elsevier Inc. All rights reserved.)

Brassinosteroids and hydrogen peroxide (H 2 O 2 ) are extensively used to combat several environmental factors, including heavy metal stress in plants, but their cumulative impact on the maintenance of copper (Cu) homeostasis in plants could not be dissected at elevated level. This study was executed to explore the roles of 24-epibrassinolide (EBL; foliar) and H 2 O 2 (root dipping) in resilience of tomato (Solanum lycopersicum L.) plants to Cu stress. The cumulative effect of EBL and H 2 O 2 in tomato plants grown under Cu stress (10 or 100 mg kg -1 soil) were assessed. Roots of 20 d old plants were submerged in 0.1 mM of H 2 O 2 solution for 4 h and subsequently transplanted in the soil-filled earthen pots and at 30 day after transplantation (DAT), the plants were sprinkled with deionized water (control), and/or 10 -8  M EBL and plant performances were evaluated at 40 DAT. High Cu (100 mg kg -1 soil) concentration considerably reduced photosynthetic efficacy, cell viability, and plant growth, and deformed chloroplast ultrastructure and root morphology with altered stomatal behavior, but boosted the activity of antioxidant enzymes, proline content and electrolyte leakage in the leaves of tomato. Moreover, EBL and H 2 O 2 implemented through distinct modes improved photosynthetic efficiency, modified chloroplast ultrastructure, stomatal behavior, root structure, cell viability and production of antioxidants and proline (osmolyte) that augmented resilience of tomato plants to Cu stress. This study revealed the potential of EBL and H 2 O 2 applied through distinct mode could serve as an effective strategy to reduce Cu-toxicity in tomato crop., (Copyright © 2020 Elsevier Inc. All rights reserved.)

A study was performed to assess if nitrate reductase (NR) participated in brassinosteroid (BR)-induced cadmium (Cd) stress tolerance primarily by accelerating the ascorbate-glutathione (AsA-GSH) cycle. Prior to initiating Cd stress (CdS), the pepper plants were sprayed with 0.5 μM 24-epibrassinolide (EBR) every other day for 10 days. Thereafter the seedlings were subjected to control or CdS (0.1 mM CdCl 2 ) for four weeks. Cadmium stress decreased the plant growth related attributes, water relations as well as the activities of monodehydroascorbate reductase (MDHAR) and dehydroascorbate reductase (DHAR), but enhanced proline content, leaf Cd 2+ content, oxidative stress-related traits, activities of ascorbate peroxidase (APX) and glutathione reductase (GR), and the activities of antioxidant defence system-related enzymes as well as NR activity and endogenous nitric oxide content. EBR reduced leaf Cd 2+ content and oxidative stress-related parameters, enhanced plant growth, regulated water relations, and led to further increases in proline content, AsA-GSH cycle-related enzymes' activities, antioxidant defence system-related enzymes as well as NR activity and endogenous nitric oxide content. The EBR and the inhibitor of NR (tungstate) reversed the positive effects of EBR by reducing NO content, showing that NR could be a potential contributor of EBR-induced generation of NO which plays an effective role in tolerance to CdS in pepper plants by accelerating the AsA-GSH cycle and antioxidant enzymes., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Lead (Pb) is a highly toxic heavy metal to plants, animals, and human beings. The use of growth regulators has reversed the effects of heavy metal stress on germination and early plant development. The aim of this study was to evaluate the effect of brassinosteroids on seed germination and seedling growth of Brassica juncea (L.) Czern. & Coss. under Pb stress conditions. Two forms of application of 24-epibrassinolide (EBL) were evaluated, application on seeds in pre-soaking and on germination paper, using EBL concentrations of 0, 10 -10 , 10 -8 , and 10 -6  M. Germination and seedling growth parameters were evaluated during the germination test. The activity of the enzymes superoxide dismutase, catalase, peroxidase, and ascorbate peroxidase were determined, as well as the lead content in the seeds and seedlings. The EBL applied at the 10 -8  M concentration was the most effective in overcoming Pb stress in both forms of application. The antioxidant enzyme defense system was compromised by Pb exposure. However, 10 -8  M EBL increased the activity of antioxidant enzymes such as catalase and peroxidase to overcome the toxic effects caused by Pb. In addition, EBL at the concentration of 10 -8  M increased Pb content in seedlings without affecting seedling growth., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Lead is one of the most hazardous pollutants to both the environment as well as human beings. As one of the approaches to enhance phytoremediation, brassinosteroids are predicted as a potential candidate phytohormone for assisted phytoremediation. Few studies have focused on the physiological regulations of tall fescue plants (Festuca arundinacea Schreb.), a potential phytoremediation species, for its responses to applications of brassinosteroids under lead stress. Therefore, the objectives of this study were to investigate the effects of foliar application of 24-epibrassinolide, a brassinosteroids analogue, on reactive oxygen species accumulation and antioxidative defense systems of tall fescue when exposed to lead, and ultimately its potential to be used in phytoremediation. When exposed to lead (1000 mg/kg) for 80 d, decreases in shoot and root biomass of tall fescue biomass as well as chlorophyll and carotenoid productions were found. Foliar application of 24-epibrassinolide at three rates and five applications every 7 d improved the biomass of both shoots and roots, and increased the photosynthetic pigments. The improved lead tolerance in tall fescue plants after 24-epibrassinolide applications was associated with reduced H 2 O 2 and O 2 .- accumulations and increased antioxidative enzyme activities including superoxide dismutase, catalase, and guaiacol peroxidase. Additionally, osmoprotectants increased and lipid peroxidation decreased. Ultimately, foliar applications of 24-epibrassinolide enhanced the lead recovery rate of tall fescue plants, proving its potential role in phytoremediation for soil contaminated with heavy metals such as lead., (Copyright © 2019 Elsevier Inc. All rights reserved.)

In the current investigation, we studied role of castasterone (CS), (a bioactive brassinosteroid) in Brassica juncea grown under imidacloprid (IMI) stress. We observed that CS-seed treatment resulted in the recovery of seedling growth under IMI toxicity. Seed treatment with CS, significantly enhanced the contents of pigments like chlorophylls, carotenoids, anthocyanins and xanthophylls under stress. Oxidative stress generated by the production of reactive oxygen species (ROS) like hydrogen peroxide and superoxide anion, was reduced after CS treatment under IMI toxicity. Antioxidative defense system got activated after CS-seed treatment, resulting in the increased activities of enzymes. Moreover, CS-seed treatment under IMI stress also stimulated the biosynthesis of organic acids of Krebs cycle (citrate, succinate, fumarate and malate) and phenolics. We also noticed that CS is also involved in the regulation of the gene expression of some key enzymes involved in pigment metabolism (CHLASE, PSY, CHS), carbon fixation (RUBISCO), Krebs cycle (CS, SUCLG1, SDH, FH), ROS generation (RBO), antioxidative enzymes (SOD, CAT, POD, DHAR, GR, GST), phenolic biosynthesis (PAL) and pesticide detoxification system (CXE, P450, NADH). This modulated gene expression after CS-treatment activated the insecticide detoxification, leading to the reduction of IMI residues. Data analysis using multivariate statistical technique i.e. multiple linear regression, also supported the fact that CS can efficiently reduce IMI induced phytotoxicity in B. juncea., (Copyright © 2019 Elsevier Inc. All rights reserved.)

Salicylic acid (SA), Epi-brassinolide (EBL) and calcium (Ca) play crucial roles in plant development and mediate plant response to biotic and abiotic stress. This study was aimed to investigate the possible mediatory role of SA, EBL, Ca or their combination in protecting tomato plants from cadmium (Cd) toxicity. According to the results, Cd stress resulted in a significant reduction of plant dry mass, photosynthetic pigment content as well as photosynthetic rate. Exogenous application of SA decreased the malondialdehyde (MDA) level by 39.27% and increased catalase (CAT) activity by 81.17%. SA and EBL treatment significantly increased chlorophyll a (Chl a), chlorophyll b (Chl b) content, photosynthetic rate (Pn) as well as water use efficiency (WUE). SA+EBL (1:1)/Ca+SA+EBL (1:1:1) treatment obviously alleviated Cd-induced growth inhibition, the dry mass of different tomato organs were significantly increased (p < 0.05). Especially in Ca+SA+EBL treated plants, the dry mass of roots, stems and leaves increased by 141.18%, 128.57% and 118.52%, respectively. Besides, SA+EBL and Ca+SA+EBL treatments reduced the MDA level, but increased photosynthetic pigment concentration and photosynthetic efficiency. CAT activity was increased by 62.92% in Ca+SA+EBL treated plants, the WUE was increased by 557.76% in SA+EBL pretreated plants. Moreover, exogenous application of SA, SA+EBL and Ca+SA+EBL significantly decreased Cd accumulation in tomato organs (p < 0.05) compared with Cd-stressed plants. Taken together, our results indicated that exogenous application of SA, EBL and Ca individually or in combination could alleviate Cd toxicity in tomato plants, although the extent varies., (Copyright © 2018 Elsevier Inc. All rights reserved.)

The study focuses on potential of combined pre-soaking treatment of 24-Epibrassinolide (EBL) and Salicylic acid (SA) in alleviating Pb phytotoxicity in Brassica juncea L. plants. The seeds after treatment with combination of both the hormones were sown in mixture of soil, sand and manure (3:1:1) and were exposed to Pb concentrations (0.25mM, 0.50mM and 0.75mM). After 30 days of growth, the plants were harvested and processed, for quantification of various metabolites. It was found that pre-sowing of seeds in combination of EBL and SA, mitigated the adverse effects of metal stress by modulating antioxidative defense response and enhanced osmolyte contents. Dry matter content and heavy metal tolerance index were enhanced in response to co-application of EBL and SA. The levels of superoxide anions, hydrogen peroxide and malondialdehyde were lowered by the combined treatment of hormones. Enhancement in activities of guaiacol peroxidase, catalase, glutathione reductase and glutathione-s-transferase were recorded. Contents of glutathione, tocopherol and ascorbic acid were also enhanced in response to co-application of both hormones. Expression of POD, CAT, GR and GST1 genes were up-regulated whereas SOD gene was observed to be down-regulated. Contents of proline, trehalose and glycine betaine were also reported to be elevated as a result of treatment with EBL+SA. The results suggest that co-application of EBL+SA may play an imperative role in improving the antioxidative defense expression of B. juncea plants to combat the oxidative stress generated by Pb toxicity., (Copyright © 2017 Elsevier Inc. All rights reserved.)

Industrialization and urbanization have posed serious threats to the environment. Excessive release of heavy metals from industrial effluents and overuse of pesticides in modern agriculture are limiting crop production by polluting environment and deteriorating food quality. Sustaining food quality under heavy metals and pesticide stress is crucial to meet the increasing demands for food. 24-Epibrassinolide (EBL), a ubiquitously occurring plant growth hormone shows great potential to alleviate heavy metals and pesticide stress in plants. This review sums up the potential role of EBL in ameliorating heavy metals and pesticide toxicity in plants extensively. EBL application increases plant's overall growth, biomass accumulation and photosynthetic efficiency by the modulation of numerous biochemical and physiological processes under heavy metals and pesticide stress. In addition, EBL scavenges reactive oxygen species (ROS) by triggering the production of antioxidant enzymes such as SOD, CAT, POX etc. EBL also induces the production of proline and soluble proteins that helps in maintaining osmotic potential and osmo-protection under both heavy metals and pesticide stress. At the end, future needs of research about the application of 24-epibrassinolide have also been discussed., (Copyright © 2017 Elsevier Inc. All rights reserved.)

Pesticides are widely used to protect crop plants from various insect pests. However, application of pesticides causes phytotoxicity to plants which results in their impaired growth and development. Brassinosteroids are well known to protect plants under abiotic stress conditions. The purpose of the present study was to access the ameliorative role of 24-epibrassinolide (EBR) in Brassica juncea L. under imidacloprid (IMI) toxicity. B. juncea plants were raised from seeds soaked in 0.1, 1 and 100nM of EBR, and grown in soils amended with 250, 300 and 350mgkg(-1) IMI pesticide, and observed for growth, pigments and photosynthetic parameters after 30, 60 and 90 days of seed sowing. The plants grown in soil treated with IMI exhibited a significant reduction in shoot length, number of leaves, chlorophyll contents and photosynthetic parameters like photosynthetic rate, stomatal conductance, inter-cellular CO2 and transpiration rate, when compared with their respective controls. However, pigments which act as antioxidants such as carotenoids, anthocyanins and xanthophylls were increased with IMI stress. Pre-sowing seed treatment with EBR decreased the toxic effects of IMI and increased the growth, pigment biosynthesis and photosynthetic parameters of the plants grown in IMI amended soil. Maximum increase in all the growth and photosynthetic parameters was noticed in plants raised from seeds treated with 100nM EBR and grown in IMI amended soil., (Copyright © 2016 Elsevier Inc. All rights reserved.)

24-Epibrassinolide (EBL) and Selenium (Se) individually confer tolerance to various abiotic stresses, but their interactive effect in the regulation of copper (Cu) homeostasis in plants exposed to toxic levels of Cu is poorly investigated. This study provides an insight into the effects of EBL (foliar) and/or Se (through sand) on Brassica juncea plants exposed to toxic levels of Cu. The combined effect of EBL and Se on compartmentalization of Cu, oxidative stress markers, photosynthetic machinery and biochemical traits in B. juncea were analyzed. Application of EBL and Se through different mode modulated the compartmentalization of Cu in different parts of plants, enhanced the photosynthetic traits, and activities of various antioxidant enzymes and proline accumulation in B. juncea under excess copper levels. These enhanced levels of antioxidant enzymes, proline (osmolyte) accumulation triggered by combination of EBL and Se could have conferred tolerance to the B. juncea plants under toxic level of copper and also maintained Cu homeostasis in various parts of plants. This study indicates that combination of EBL and Se through different mode is an operative approach for Cu detoxification in plants and could be exploited for removal of excess copper from polluted soil., (Copyright © 2016. Published by Elsevier Inc.)

Polycyclic aromatic hydrocarbons (PAHs) are toxic to both plants and animals. The enhancement of plant tolerance and detoxification capacity is important for the plant-based remediation of PAHs. Therefore, we investigated the effects of 24-epibrassinolide (EBR) on the metabolism of a three-ringed PAH (phenanthrene-PHE) and subsequent stress tolerance in tomato (Solanum lycopersicum L.) plants. Exposure to PHE (300 μM) for 21 d significantly decreased biomass and net CO(2) assimilation (P(n)) but induced photoinhibition, malondialdehyde (MDA), H(2)O(2) and antioxidant enzymes. Obvious ultrastructural alterations were observed in the PHE-treated root tip cells. Importantly, the foliar application of EBR (0.1 μM) significantly increased biomass, P(n) and antioxidant enzyme activities but decreased MDA and H(2)O(2) compared with PHE alone and saved the root cells from severe damage. The expression of detoxification genes (CYP90b3, GSH1, GST1), reduced glutathione (GSH) content and glutathione S-transferase activity in the EBR+PHE-treated plants were higher than those of PHE alone. Additionally, lower levels of PHE residues in the roots were observed as a result of EBR+PHE treatment. Taken together, our results strongly suggest an enhanced and coordinated detoxification and degradation of PHE by EBR., (Copyright © 2012 Elsevier Inc. All rights reserved.)