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1.  Brassinosteroids Improve Quality of Summer Tea (Camellia sinensis L.) by Balancing Biosynthesis of Polyphenols and Amino Acids 
Summer grown green tea is less popular due to bitterness and high astringency, which are attributed to high levels of tea polyphenols (TP) and low levels of amino acids (AA) in tea leaves (Camellia sinensis L.). Brassinosteroids (BRs), a group of steroidal plant hormones can regulate primary and secondary metabolism in a range of plant species under both normal and stress conditions. However, specific effects of BRs on the photosynthesis of tea plants and the quality of summer green tea are largely unknown. Here we show that 24-epibrassinolide (EBR), a bioactive BR, promoted photosynthesis in tea plants in a concentration-dependent manner. Stimulation in photosynthesis by EBR resulted in an increased summer tea yield. Although all tested concentrations (0.01, 0.05, 0.1, 0.5, and 1.0 ppm) of EBR increased concentrations of TP and AA, a moderate concentration (0.5 ppm) caused the highest decrease in TP to AA ratio, an important feature of quality tea. Time-course analysis using 0.5 ppm EBR as foliar spray revealed that TP or AA concentration increased as early as 3 h after EBR application, reaching the highest peak at 24 h and that remained more or less stable. Importantly, such changes in TP and AA concentration by EBR resulted in a remarkably decreased but stable TP to AA ratio at 24 h and onward. Furthermore, concentrations of catechins and theanine increased, while that of caffeine remained unaltered following treatment with EBR. EBR improved activity of phenylalanine ammonia-lyase (PAL) and glutamine: 2-oxoglutarate aminotransferase (GOGAT) enzymes involved in catechins and theanine biosynthesis, respectively. Transcript analysis revealed that transcript levels of CsPAL and CsGS peaked as early as 6 h, while that of CsGOGAT peaked at 12 h following application of EBR, implying that EBR increased the concentration of TP and AA by inducing their biosynthesis. These results suggest a positive role of BR in enhancing green tea quality, which might have potential implication in improving quality of summer tea.
PMCID: PMC5003824  PMID: 27625668
24-epibrassinolide; amino acids; photosynthesis; polyphenol; secondary metabolism; summer tea; tea quality
2.  Transcriptome Analysis of Pepper (Capsicum annuum) Revealed a Role of 24-Epibrassinolide in Response to Chilling 
Brassinosteroids (BRs) have positive effects on many processes during plant growth, development, and various abiotic stress responses. However, little information is available regarding the global gene expression of BRs in response to chilling stress in pepper. In this study, we used RNA sequencing to determine the molecular roles of 24-epibrassinolide (EBR) during a chilling stress response. There were 39,829 transcripts, and, among them, 656 were differently-expressed genes (DEGs) following EBR treatment (Chill+EBR) compared with the control (Chill only), including 335 up-regulated and 321 down-regulated DEGs. We selected 20 genes out of the 656 DEGs for RT-qPCR analysis to confirm the RNA-Seq. Based on GO enrich and KEGG pathway analysis, we found that photosynthesis was significantly up-enriched in biological processes, accompanied by significant increases in the net photosynthetic rate (Pn), Fv/Fm, and chlorophyll content. Furthermore, the results indicate that EBR enhanced endogenous levels of salicylic acid (SA) and jasmonic acid (JA) while suppressing the ethylene (ETH) biosynthesis pathway, suggesting that BRs function via a synergistic cross-talk with SA, JA, and ETH signaling pathways in response to chilling stress. In addition, EBR induced cellulose synthase-like protein and UDP-glycosyltransferase, suggesting a contribution to the formation of cell wall and hormone metabolism. EBR also triggered the calcium signaling transduction in cytoplasm, and activated the expression of cellular redox homeostasis related genes, such as GSTX1, PER72, and CAT2. This work, therefor, identified the specific genes showed different expression patterns in EBR-treated pepper and associated with the processes of hormone metabolism, redox, signaling, transcription, and defense. Our study provides the first evidence of the potent roles of BRs, at the transcription level, to induce the tolerance to chilling stress in pepper as a function of the combination of the transcriptional activities, signaling transduction, and metabolic homeostasis.
PMCID: PMC5002408  PMID: 27621739
pepper; Brassinosteroid; chill-stress; transcriptome; RNA sequencing
3.  Brassinosteroids are involved in response of cucumber (Cucumis sativus) to iron deficiency 
Annals of Botany  2012;110(3):681-688.
Background and Aims
Brassinosteroids (BR) are a class of plant polyhydroxysteroids with diverse functions in plant growth and development. However, there is little information on the role of BRs played in the response to nutrient deficiency.
To evaluate the role of BR in the response of plants to iron (Fe) deficiency, the effect of 24-epibrassinolide (EBR) on ferric reductase (FRO) activity, acidification of the rhizosphere and Fe content in cucumber (Cucumis sativus) seedlings under Fe-deficient (1 µm FeEDTA) and Fe-sufficient (50 µm FeEDTA) conditions were investigated.
Key Results
There was a significant increase in FRO activity upon exposure of cucumber seedlings to an Fe-deficient medium, and the Fe deficiency-induced increase in FRO activity was substantially suppressed by EBR. In contrast, application of EBR to Fe-sufficient seedlings stimulated FRO activity. Ethylene production evoked by Fe deficiency was suppressed by EBR, while EBR induced ethylene production from Fe-sufficient seedlings. Fe contents in shoots were reduced by treatment with EBR, while Fe contents in roots were markedly increased under both Fe-deficient and Fe-sufficient conditions. The reductions in Fe contents of shoots were independent of chlorophyll (CHL) contents under Fe-sufficient conditions, but they were positively correlated with CHL contents under Fe-deficient conditions. At the transcriptional level, transcripts encoding FRO (CsFRO1) and Fe transporter (CsIRT1) were increased upon exposure to the Fe-deficient medium, and the increases in transcripts were reversed by EBR.
The results demonstrate that BRs are likely to play a negative role in regulating Fe-deficiency-induced FRO, expressions of CsFRO1 and CsIRT1, as well as Fe translocation from roots to shoots.
PMCID: PMC3400454  PMID: 22684685
Brassinosteroids; iron deficiency; cucumber; Cucumis sativus; ferric reductase activity; Fe translocation
4.  Brassinosteroids alleviate high-temperature injury in Ficus concinna seedlings via maintaining higher antioxidant defence and glyoxalase systems 
AoB Plants  2015;7:plv009.
Brassinosteroids play a significant role in the amelioration of various biotic and abiotic stresses. To investigate the effects of exogenously applied brassinosteroids on the thermotolerance of plants, reactive oxygen species (ROS), antioxidant defense and methylglyoxal (MG) detoxification systems were examined in Ficus concinna seedlings with or without 24-epibrassinolide (EBR) application. Our results showed that EBR treatment reduced high temperature-induced increases in the levels of ROS, MG and lipid peroxidation. We also demonstrate that EBR attenuates high temperature-induced oxidative stress by simultaneously increasing non-enzymatic and enzymatic antioxidant responses, as well as MG detoxification systems.
Although brassinosteroids (BRs) play crucial roles in plant development and stress tolerance, the mechanisms by which they have these effects are poorly understood. Here, we investigated the possible mechanism of exogenously applied BRs on reactive oxygen species (ROS), antioxidant defence and methylglyoxal (MG) detoxification systems in Ficus concinna seedlings grown under high-temperature (HT) stress for 48 h. Our results showed that the activities of ascorbate peroxidase (APX), superoxide dismutase (SOD), glutathione S-transferase (GST), glutathione peroxidase (GPX) and glyoxalase II (Gly II) were increased under two levels of HT stress. Compared with control the activities of catalase (CAT) and dehydroascorbate reductase (DHAR) were not changed due to HT stress. The activities of glutathione reductase (GR), monodehydroascorbate reductase (MDHAR) and glyoxalase I (Gly I) were increased only at moderate HT stress. Despite these protective mechanisms, HT stress induced oxidative stress in F. concinna seedlings, as indicated by the increased levels of ROS, malondialdehyde (MDA) and MG, and the reductions in chlorophyll levels and relative water content. The contents of reduced glutathione (GSH) and ascorbate (AsA) were not changed under moderate HT stress. Spraying with 24-epibrassinolide (EBR) alone had little influence on the non-enzymatic antioxidants and the activities of antioxidant enzymes. However, EBR pretreatment under HT stress resulted in an increase in GSH and AsA content, maintenance of high redox state of GSH and AsA, and enhanced ROS and MG detoxification by further elevating the activities of SOD, GST, GPX, APX, MDHAR, GR, DHAR, Gly I and Gly II, as evident by lower level of ROS, MDA and MG. It may be concluded that EBR could alleviate the HT-induced oxidative stress by increasing the enzymatic and non-enzymatic antioxidant defence, and glyoxalase systems in F. concinna seedlings.
PMCID: PMC4344480  PMID: 25609563
Antioxidants; brassinosteroids; glyoxalase system; high-temperature stress; methylglyoxal
5.  Role of brassinosteroids in alleviation of phenanthrene–cadmium co-contamination-induced photosynthetic inhibition and oxidative stress in tomato 
Journal of Experimental Botany  2012;64(1):199-213.
Heavy metal pollution often occurs together with organic contaminants. Brassinosteroids (BRs) induce plant tolerance to several abiotic stresses, including phenanthrene (PHE) and cadmium (Cd) stress. However, the role of BRs in PHE+Cd co-contamination-induced stress amelioration is unknown. Here, the interactive effects of PHE, Cd, and 24-epibrassinolide (EBR; a biologically active BR) were investigated in tomato plants. The application of Cd (100 µM) alone was more phytotoxic than PHE applied alone (100 µM); however, their combined application resulted in slightly improved photosynthetic activity and pigment content compared with Cd alone after a 40 d exposure. Accumulation of reactive oxygen species and membrane lipid peroxidation were induced by PHE and/or Cd; however, the differences in effect were insignificant between Cd and PHE+Cd. The foliar application of EBR (0.1 µM) to PHE- and/or Cd-stressed plants alleviated photosynthetic inhibition and oxidative stress by causing enhancement of the activity of the enzymes and related transcript levels of the antioxidant system, secondary metabolism, and the xenobiotic detoxification system. Additionally, PHE and/or Cd residues were significantly decreased in both the leaves and roots after application of EBR, more specifically in PHE+Cd-stressed plants when treated with EBR, indicating a possible improvement in detoxification of these pollutants. The findings thus suggest a potential interaction of EBR and PHE for Cd stress alleviation. These results advocate a positive role for EBR in reducing pollutant residues for food safety and also strengthening phytoremediation.
PMCID: PMC3528031  PMID: 23201830
Brassinosteroids; food safety; heavy metal; photosynthesis; phytoremediation; polycyclic aromatic hydrocarbons (PAHs)
6.  SILAC-Based Mass Spectrometry Analysis Reveals That Epibrassinolide Induces Apoptosis via Activating Endoplasmic Reticulum Stress in Prostate Cancer Cells 
PLoS ONE  2015;10(9):e0135788.
Epibrassinolide (EBR) is a polyhydroxylated sterol derivative and biologically active compound of the brassinosteroids. In addition to well-described roles in plant growth, EBR induces apoptosis in the LNCaP prostate cancer cells expressing functional androgen receptor (AR). Therefore, it is suggested that EBR might have an inhibitory potential on androgen receptor signaling pathway. However, the mechanism by which EBR exerts its effects on LNCaP is poorly understood. To address this gap in knowledge, we used an unbiased global proteomics approach, i.e., stable-isotope labeling by amino acids in cell culture (SILAC). In total, 964 unique proteins were identified, 160 of which were differentially expressed after 12 h of EBR treatment. The quantification of the differentially expressed proteins revealed that the expression of the unfolded protein response (UPR) chaperone protein, calreticulin (CALR), was dramatically downregulated. The decrease in CALR expression was also validated by immunoblotting. Because our data revealed the involvement of the UPR in response to EBR exposure, we evaluated the expression of the other UPR proteins. We demonstrated that EBR treatment downregulated calnexin and upregulated BiP and IRE1α expression levels and induced CHOP translocation from the cytoplasm to nucleus. The translocation of CHOP was associated with caspase-9 and caspase-3 activation after a 12 h EBR treatment. Co-treatment of EBR with rapamycin, an upstream mTOR pathway inhibitor, prevented EBR-induced cell viability loss and PARP cleavage in LNCaP prostate cancer cells, suggesting that EBR could induce ER stress in these cells. In addition, we observed similar results in DU145 cells with nonfunctional androgen receptor. When proteasomal degradation of proteins was blocked by MG132 co-treatment, EBR treatment further induced PARP cleavage relative to drug treatment alone. EBR also induced Ca2+ sequestration, which confirmed the alteration of the ER pathway due to drug treatment. Therefore, we suggest that EBR promotes ER stress and induces apoptosis.
PMCID: PMC4564160  PMID: 26353013
7.  Physiological and biochemical effect of 24-epibrassinoslide on cold tolerance in maize seedlings 
Germination and early seedling growth are important for establishment of maize because maize is chilling sensitive crop and low temperature during early period of growth can be detrimental to subsequent crop growth and productivity. Therefore, it is important to protect maize seedling from cold stress. A study was conducted on induced cold tolerance by 24-epibrassinoslide (EBR) at the Indian Agricultural Research Institute, New Delhi, India. Maize seedlings were raised in green house condition (25/18 °C day-night temperatures). Ten days old seedlings were treated with EBR (0.0, 0.01, 0.1, 1.0 and 10 μM) and then divided into two sets, one set was kept in greenhouse (25/18 °C day-night temperatures) and another was transferred to net house (cold stress). Data on various morpho-physiological traits was recorded after 7, 14 and 21 days of treatment. Exogenous application of 1.0 μM EBR had significant effect on growth and morpho-physiological traits under both conditions. The maize seedlings treated with EBR were more tolerant to cold stress than the untreated one. Significant increase in plant height, dry matter accumulation, chlorophyll content, total soluble proteins and starch contents was observed under both conditions, however, the results were more pronounced under cold stress. 1.0 μ M concentration being the most effective under both conditions. Maintenance of high tissue water content, reduced membrane injury index, increased total chlorophyll, soluble sugar and protein content were taken as the possible indicators of EBR induced chilling tolerance.
PMCID: PMC3550514  PMID: 23814437
24-epibrassinoslide; Cold tolerance; Maize; Glycine betaine; Total chlorophyll; Membrane injury index
8.  A role of brassinosteroids in early fruit development in cucumber 
Journal of Experimental Botany  2008;59(9):2299-2308.
Brassinosteroids (BRs) are essential for many biological processes in plants, however, little is known about their roles in early fruit development. To address this, BR levels were manipulated through the application of exogenous BRs (24-epibrassinolide, EBR) or a BR biosynthesis inhibitor (brassinazole, Brz) and their effects on early fruit development, cell division, and expression of cyclin and cyclin-dependent kinases (CDKs) genes were examined in two cucumber cultivars that differ in parthenocarpic capacity. The application of EBR induced parthenocarpic growth accompanied by active cell division in Jinchun No. 4, a cultivar without parthenocarpic capacity, whereas Brz treatment inhibited fruit set and, subsequently, fruit growth in Jinchun No. 2, a cultivar with natural parthenocarpic capacity, and this inhibitory effect could be rescued by the application of EBR. RT-PCR analysis showed both pollination and EBR induced expression of cell cycle-related genes (CycA, CycB, CycD3;1, CycD3;2, and CDKB) after anthesis. cDNA sequences for CsCycD3;1 and CsCycD3;2 were isolated through PCR amplification. Both CsCycD3;1 and CsCycD3;2 transcripts were up-regulated by EBR treatment and pollination but strongly repressed by Brz treatment. Meanwhile, BR6ox1 and SMT transcripts, two genes involved in BR synthesis, exhibited feedback regulation. These results strongly suggest that BRs play an important role during early fruit development in cucumber.
PMCID: PMC2423651  PMID: 18515830
Brassinosteroids; cell division; Cucumis sativus; cyclin; flow cytometry; parthenocarpy
9.  Brassinosteroids are involved in Fe homeostasis in rice (Oryza sativa L.) 
Journal of Experimental Botany  2015;66(9):2749-2761.
Brassinosteroids negatively regulate Fe transport and translocation in a strategy II rice plants.
Brassinosteroids (BRs) are steroid hormones that modulate numerous physiological processes in plants. However, few studies have focused on the involvement of BRs in sensing and responding to the stress of mineral nutrient deficiency. In the present study, we evaluated the roles of BRs in the response of rice (Oryza sativa) to iron (Fe) deficiency during Fe uptake, transport, and translocation. Exogenous application of 24-epibrassinolide (EBR) to wild-type (WT) plants exaggerated leaf symptoms of Fe deficiency and suppressed growth. EBR increased and decreased Fe concentrations in roots and shoots, respectively, under both Fe-deficient and Fe-sufficient conditions. Transcripts involved in Fe homeostasis, including OsIRT1, OsYSL15, OsYSL2, OsNAS1, and OsNAS2, were enhanced by EBR under Fe-deficient conditions. EBR depressed expression of OsNAS1, OsNAS2, and OsYSL2 in shoots, and inhibited Fe transport and translocation via the phloem. Rice mutant d2-1, which is defective in BR biosynthesis, was more tolerant to Fe deficiency than the WT, and accumulated greater amounts of Fe in roots than the WT under Fe-sufficient conditions. A greater upregulation of OsIRT1, OsYSL15, OsYSL2, OsNAS1, and OsNAS2 in the d2-1 mutant compared to the WT was found under Fe-sufficient conditions, while expression of these genes in the d2-1 mutant was lower than in the WT under Fe-deficient conditions. The greater tolerance of the d2-1 mutant could be partly mitigated by exogenous application of EBR. These novel findings highlight the important role of BR in mediating the response of strategy II plants to Fe deficiency by regulating Fe uptake and translocation in rice.
PMCID: PMC4986876  PMID: 25770588
Brassinosteroids (BRs); d2-1 mutant; Fe deficiency; Fe translocation; Fe uptake; rice (Oryza sativa); strategy II plant.
10.  Identification of MicroRNA 395a in 24-Epibrassinolide-Regulated Root Growth of Arabidopsis thaliana Using MicroRNA Arrays 
Brassinosteroids (BRs) are endogenous plant hormones and are essential for normal plant growth and development. MicroRNAs (miRNAs) of Arabidopsis thaliana are involved in mediating cell proliferation in leaves, stress tolerance, and root development. The specifics of BR mechanisms involving miRNAs are unknown. Using customized miRNA array analysis, we identified miRNAs from A. thaliana ecotype Columbia (Col-0) regulated by 24-epibrassinolide (EBR, a highly active BR). We found that miR395a was significantly up-regulated by EBR treatment and validated its expression under these conditions. miR395a was over expressed in leaf veins and root tissues in EBR-treated miR395a promoter::GUS plants. We integrated bioinformatics methods and publicly available DNA microarray data to predict potential targets of miR395a. GUN5—a multifunctional protein involved in plant metabolic functions such as chlorophyll synthesis and the abscisic acid (ABA) pathway—was identified as a possible target. ABI4 and ABI5, both genes positively regulated by ABA, were down-regulated by EBR treatment. In summary, our results suggest that EBR regulates seedling development and root growth of A. thaliana through miR395a by suppressing GUN5 expression and its downstream signal transduction.
PMCID: PMC3742243  PMID: 23839095
brassinosteroids; miR395a; root growth; Arabidopsis thaliana; microRNA array
11.  Pre-sowing Seed Treatment with 24-Epibrassinolide Ameliorates Pesticide Stress in Brassica juncea L. through the Modulation of Stress Markers 
The present experiment was designed to assess the effects of seed soaking with 24-epibrassinolide (EBR) on the physiology of Brassica juncea L. seedlings grown under imidacloprid (IMI) toxicity. Application of EBR increased the length of seedlings, dry weight, and pigment contents, polyphenols, total phenols, and organic acids under IMI toxicity. The expression of genes coding key enzymes of pigment, phenols, polyphenols, and organic acid biosynthetic pathways was also studied including CHLASE (chlorophyllase), PSY (phytoene synthase), CHS (chalcone synthase) and PAL (phenylalanine ammonialyase), CS (citrate synthase), SUCLG1 (succinyl Co-A ligase,), SDH (succinate dehydrogenase), FH (fumarate hydratase), MS (malate synthase). Multiple linear regression (MLR) analysis revealed that IMI application regressed negatively on seedling length, dry weight and total chlorophyll content. However, EBR seed treatment regressed positively on all the parameters studied. Moreover, interaction between IMI and EBR showed positive regression for growth parameters, content of pigments, total polyphenol, total phenol and malate, and expression of PSY and PAL. Negative interactions were noticed for the contents of fumarate, succinate and citrate, and expression of CHS and all genes studied related to organic acid metabolism. In conclusion, EBR enhanced the growth and contents of all studied metabolites by regulating the gene expression of B. juncea seedlings under IMI stress.
PMCID: PMC5089990  PMID: 27853460
mustard; brassinosteroids; pigments; total phenols; phenylalanine ammonialyase; GC-MS; HPLC
12.  Endothelial bioreactor system ameliorates multiple organ dysfunction in septic rats 
The endothelium is a potentially valuable target for sepsis therapy. We have previously studied an extracorporeal endothelial cell therapy system, called the endothelial bioreactor (EBR), which prolonged the survival time of endotoxemia sepsis in swine. To further study of the therapeutic effects and possible mechanisms, we established a miniature EBR system for septic rats induced by cecal ligation and puncture (CLP).
In the miniature EBR system, the extracorporeal circulation first passed through a mini-hemofilter, and the ultrafiltrate (UF) was separated, then the UF passed through an EBR (a 1-mL cartridge containing approximately 2 × 106 endothelial cells grown on microcarriers) and interact with endothelial cells. Eighteen hours after CLP, the rats were treated for 4 h with this extracorporeal system containing either endothelial cells (EBR group) or no cells (sham EBR group). Physiologic and biochemical parameters, cytokines, endothelial functions, and 7-day survival time were monitored. In vitro, the pulmonary endothelial cells of the septic rats were treated with the EBR system and the resulting changes in their functions were monitored.
The EBR system ameliorated CLP-induced sepsis compared with the sham EBR system. After CLP, the 7-day survival rate of sham-treated rats was only 25.0 %, while in the EBR-treated group, it increased to 57.1 % (p = 0.04). The EBR system protected the liver and renal function and ameliorated the kidney and lung injury. Meanwhile, this therapy reduced pulmonary vascular leakage and alleviated the infiltration of inflammatory cells in the lungs, especially neutrophils. Furthermore, after the EBR treatment both in vivo and in vitro, the expression of intercellular adhesion molecule-1 and the secretion of CXCL1 and CXCL2 of pulmonary endothelium decreased, which helped to alleviate the adhesion and chemotaxis of neutrophils. In addition, the EBR system decreased CD11b expression and intracellular free calcium level of peripheral blood neutrophils, modulated the activation of these neutrophils.
The EBR system significantly ameliorated CLP-induced sepsis and improved survival and organ functions. Compared with the sham EBR system, this extracorporeal endothelial therapy may be involved in modulating the function of pulmonary endothelial cells, reducing the adhesion and chemotaxis of neutrophil, and modulating the activation of peripheral blood neutrophils.
Electronic supplementary material
The online version of this article (doi:10.1186/s40635-016-0097-y) contains supplementary material, which is available to authorized users.
PMCID: PMC4958089  PMID: 27447715
Sepsis; Multiple organ dysfunction; Endothelial cells; Bioreactor; Extracorporeal circulation; Neutrophils; Inflammation
13.  Brassinosteroid-mediated stress tolerance in Arabidopsis shows interactions with abscisic acid, ethylene and salicylic acid pathways 
BMC Plant Biology  2010;10:151.
Brassinosteroids (BRs) play crucial roles in plant development and also promote tolerance to a range of abiotic stresses. Although much has been learned about their roles in plant development, the mechanisms by which BRs control plant stress responses and regulate stress-responsive gene expression are not fully known. Since BR interacts with other plant hormones, it is likely that the stress tolerance conferring ability of BR lies in part in its interactions with other stress hormones.
Using a collection of Arabidopsis mutants that are either deficient in or insensitive to abscisic acid (ABA), ethylene (ET), jasmonic acid (JA) and salicylic acid (SA), we studied the effects of 24-epibrassinloide (EBR) on basic thermotolerance and salt tolerance of these mutants. The positive impact of EBR on thermotolerance in proportion to wild type was evident in all mutants studied, with the exception of the SA-insensitive npr1-1 mutant. EBR could rescue the ET-insensitive ein2 mutant from its hypersensitivity to salt stress-induced inhibition of seed germination, but remained ineffective in increasing the survival of eto1-1 (ET-overproducer) and npr1-1 seedlings on salt. The positive effect of EBR was significantly greater in the ABA-deficient aba1-1 mutant as compared to wild type, indicating that ABA masks BR effects in plant stress responses. Treatment with EBR increased expression of various hormone marker genes in both wild type and mutant seedlings, although to different levels.
These results together indicate that the redox-sensitive protein NPR1 (NONEXPRESSOR OF PATHOGENESIS-RELATED GENES1), a master regulator of SA-mediated defense genes, is likely a critical component of EBR-mediated increase in thermotolerance and salt tolerance, but it is not required for EBR-mediated induction of PR-1 (PATHOGENESIS-RELATED1) gene expression; that BR exerts anti-stress effects independently as well as through interactions with other hormones; that ABA inhibits BR effects during stress; and that BR shares transcriptional targets with other hormones.
PMCID: PMC3095295  PMID: 20642851
14.  Effect of 24-epibrassinolide treatment on the metabolism of eggplant fruits in relation to development of pulp browning under chilling stress 
This study aims to investigate the effect of 24-epibrassinolide (EBR) on the metabolism in relation to development of chilling injury-induced pulp browning of eggplant fruit. The fruits were dipped for 10 min in solutions containing 10 μmM EBR and then stored at 1 °C for 15 days. Chilling injury index, weight loss, electrolyte leakage and malondialdehyde (MDA) content of control fruit increased during storage. Chilling injury improved phenylalanine ammonia-lyase (PAL), polyphenol oxidase (PPO), and peroxidase (POD) activities, which are correlated with the increase of total phenolic content and pulp browning of eggplant fruit. The inhibition of pulp browning by EBR treatment was possibly attributed to preserving the cell membrane integrity, reducing total phenolic content, and decreasing PAL, PPO, and POD activities. These results suggest that EBR may inhibit chilling injury and pulp browning in eggplant fruit during cold storage.
PMCID: PMC4444923  PMID: 26028720
Eggplant fruit; Brassinosteroids; Chilling injury; Pulp browning; Phenolic metabolism
15.  Interaction of Brassinosteroids and Polyamines Enhances Copper Stress Tolerance in Raphanus Sativus  
Journal of Experimental Botany  2012;63(15):5659-5675.
Brassinosteroids (BRs) and polyamines (PAs) regulate various responses to abiotic stress, but their involvement in the regulation of copper (Cu) homeostasis in plants exposed to toxic levels of Cu is poorly understood. This study provides an analysis of the effects of exogenously applied BRs and PAs on radish (Raphanus sativus) plants exposed to toxic concentrations of Cu. The interaction of 24-epibrassinolide (EBR, an active BR) and spermidine (Spd, an active PA) on gene expression and the physiology of radish plants resulted in enhanced tolerance to Cu stress. Results indicated that the combined application of EBR and Spd modulated the expression of genes encoding PA enzymes and genes that impact the metabolism of indole-3-acetic acid (IAA) and abscisic acid (ABA) resulting in enhanced Cu stress tolerance. Altered expression of genes implicated in Cu homeostasis appeared to be the main effect of EBR and Spd leading to Cu stress alleviation in radish. Ion leakage, in vivo imaging of H2O2, comet assay, and improved tolerance of Cu-sensitive yeast strains provided further evidence for the ability of EBR and Spd to improve Cu tolerance significantly. The study indicates that co-application of EBR and Spd is an effective approach for Cu detoxification and the maintenance of Cu homeostasis in plants. Therefore, the use of these compounds in agricultural production systems should be explored.
PMCID: PMC3444278  PMID: 22915739
Abscisic acid; brassinosteroids; comet assay; copper transporters; Cu homeostasis; Cu-sensitive yeast; indole-3-acetic acid; oxidative stress; polyamines
16.  Changes in Photosynthetic Rates and Gene Expression of Leaves during a Source–Sink Perturbation in Sugarcane 
Annals of Botany  2007;101(1):89-102.
Background and Aims
In crops other than sugarcane there is good evidence that the size and activity of carbon sinks influence source activity via sugar-related regulation of the enzymes of photosynthesis, an effect that is partly mediated through coarse regulation of gene expression.
In the current study, leaf shading treatments were used to perturb the source–sink balance in 12-month-old Saccharum spp. hybrid ‘N19’ (N19) by restricting source activity to a single mature leaf. Changes in leaf photosynthetic gas exchange variables and leaf and culm sugar concentrations were subsequently measured over a 14 d period. In addition, the changes in leaf gene response to the source–sink perturbation were measured by reverse northern hybridization analysis of an array of 128 expressed sequence tags (ESTs) related to photosynthetic and carbohydrate metabolism.
Key Results
Sucrose concentrations in immature culm tissue declined significantly over the duration of the shading treatment, while a 57 and 88% increase in the assimilation rate (A) and electron transport rate (ETR), respectively, was observed in the source leaf. Several genes (27) in the leaf displayed a >2-fold change in expression level, including the upregulation of several genes associated with C4 photosynthesis, mitochondrial metabolism and sugar transport. Changes in gene expression levels of several genes, including Rubisco (EC 4·1·1·39) and hexokinase (HXK; EC 2·7·1·1), correlated with changes in photosynthesis and tissue sugar concentrations that occurred subsequent to the source–sink perturbation.
These results are consistent with the notion that sink demand may limit source activity through a kinase-mediated sugar signalling mechanism that correlates to a decrease in source hexose concentrations, which, in turn, correlate with increased expression of genes involved in photosynthesis and metabolite transport. The signal feedback system reporting sink sufficiency and regulating source activity may be a potentially valuable target for future genetic manipulation to increase sugarcane sucrose yield.
PMCID: PMC2701831  PMID: 17942591
C4 metabolism; array; hexose; photosynthesis; sucrose; Saccharum spp.; source–sink; gene expression
17.  Brassinosteroids-Induced Systemic Stress Tolerance was Associated with Increased Transcripts of Several Defence-Related Genes in the Phloem in Cucumis sativus 
PLoS ONE  2013;8(6):e66582.
Brassinosteroids (BRs), a group of naturally occurring plant steroidal compounds, are essential for plant growth, development and stress tolerance. Recent studies showed that BRs could induce systemic tolerance to biotic and abiotic stresses; however, the molecular mechanisms by which BRs signals lead to responses in the whole plant are largely unknown. In this study, 24-epibrassinosteroid (EBR)-induced systemic tolerance in Cucumis sativus L. cv. Jinyan No. 4 was analyzed through the assessment of symptoms of photooxidative stress by chlorophyll fluorescence imaging pulse amplitude modulation. Expression of defense/stress related genes were induced in both treated local leaves and untreated systemic leaves by local EBR application. With the suppressive subtractive hybridization (SSH) library using cDNA from the phloem sap of EBR-treated plants as the tester and distilled water (DW)-treated plants as the driver, 14 transcripts out of 260 clones were identified. Quantitative Real Time-Polymerase Chain Reaction (RT-qPCR) validated the specific up-regulation of these transcripts. Of the differentially expressed transcripts with known functions, transcripts for the selected four cDNAs, which encode an auxin-responsive protein (IAA14), a putative ankyrin-repeat protein, an F-box protein (PP2), and a major latex, pathogenesis-related (MLP)-like protein, were induced in local leaves, systemic leaves and roots after foliar application of EBR onto mature leaves. Our results demonstrated that EBR-induced systemic tolerance is accompanied with increased transcript of genes in the defense response in other organs. The potential role of phloem mRNAs as signaling components in mediating BR-regulated systemic resistance is discussed.
PMCID: PMC3686678  PMID: 23840504
18.  Source/sink interactions underpin crop yield: the case for trehalose 6-phosphate/SnRK1 in improvement of wheat 
Considerable interest has been evoked by the analysis of the regulatory pathway in carbohydrate metabolism and cell growth involving the non-reducing disaccharide trehalose (TRE). TRE is at small concentrations in mesophytes such as Arabidopsis thaliana and Triticum aestivum, excluding a role in osmoregulation once suggested for it. Studies of TRE metabolism, and genetic modification of it, have shown a very wide and more important role of the pathway in regulation of many processes in development, growth, and photosynthesis. It has now been established that rather than TRE, it is trehalose 6-phosphate (T6P) which has such profound effects. T6P is the intermediary in TRE synthesis formed from glucose-6-phosphate and UDP-glucose, derived from sucrose, by the action of trehalose phosphate synthase. The concentration of T6P is determined both by the rate of synthesis, which depends on the sucrose concentration, and also by the rate of breakdown by trehalose-6-phosphate phosphatase which produces TRE. Changing T6P concentrations by genetically modifying the enzymes of synthesis and breakdown has altered photosynthesis, sugar metabolism, growth, and development which affect responses to, and recovery from, environmental factors. Many of the effects of T6P on metabolism and growth occur via the interaction of T6P with the SnRK1 protein kinase system. T6P inhibits the activity of SnRK1, which de-represses genes encoding proteins involved in anabolism. Consequently, a large concentration of sucrose increases T6P and thereby inhibits SnRK1, so stimulating growth of cells and their metabolic activity. The T6P/SnRK1 mechanism offers an important new view of how the distribution of assimilates to organs, such as developing grains in cereal plants, is achieved. This review briefly summarizes the factors determining, and limiting, yield of wheat (particularly mass/grain which is highly conserved) and considers how T6P/SnRK1 might function to determine grain yield and might be altered to increase them. Increasing the potential rate of filling and mass/grain are ways in which total crop yield could be increased with good husbandry which maintains crop assimilation Cereal yields globally are not increasing, despite the greater production required to meet human demand. Careful targeting of T6P is showing much promise for optimization of source/sink for yield improvement and offers yet further possibilities for increasing sink demand and grain size in wheat.
PMCID: PMC4142875  PMID: 25202319
source/sink; wheat; grain yield; yield components; food security; sucrose; trehalose 6-phosphate; SnRK1
19.  Metabolic and diffusional limitations of photosynthesis in fluctuating irradiance in Arabidopsis thaliana 
Scientific Reports  2016;6:31252.
A better understanding of the metabolic and diffusional limitations of photosynthesis in fluctuating irradiance can help identify targets for improving crop yields. We used different genotypes of Arabidopsis thaliana to characterise the importance of Rubisco activase (Rca), stomatal conductance (gs), non-photochemical quenching of chlorophyll fluorescence (NPQ) and sucrose phosphate synthase (SPS) on photosynthesis in fluctuating irradiance. Leaf gas exchange and chlorophyll fluorescence were measured in leaves exposed to stepwise increases and decreases in irradiance. rwt43, which has a constitutively active Rubisco enzyme in different irradiance intensities (except in darkness), showed faster increases than the wildtype, Colombia-0, in photosynthesis rates after step increases in irradiance. rca-2, having decreased Rca concentration, showed lower rates of increase. In aba2-1, high gs increased the rate of change after stepwise irradiance increases, while in C24, low gs tended to decrease it. Differences in rates of change between Colombia-0 and plants with low levels of NPQ (npq1-2, npq4-1) or SPS (spsa1) were negligible. In Colombia-0, the regulation of Rubisco activation and of gs were therefore limiting for photosynthesis in fluctuating irradiance, while levels of NPQ or SPS were not. This suggests Rca and gs as targets for improvement of photosynthesis of plants in fluctuating irradiance.
PMCID: PMC4977489  PMID: 27502328
20.  Role of Evidence-Based Reviews in Surgery in teaching critical appraisal skills and in journal clubs 
Canadian Journal of Surgery  2013;56(4):E98-E102.
Evidence-Based Reviews in Surgery (EBRS) is a program developed to teach critical appraisal skills to general surgeons and residents. The purpose of this study was to assess the use of EBRS by general surgery residents across Canada and to assess residents’ opinions regarding EBRS and journal clubs.
We surveyed postgraduate year 2–5 residents from 15 general surgery programs. Data are presented as percentages and means.
A total of 231 residents (58%, mean 56% per program, range 0%–100%) responded: 172 (75%) residents indicated that they know about EBRS and that it is used in their programs. More than 75% of residents who use EBRS agreed or strongly agreed that the EBRS clinical and methodological articles and reviews are relevant. Only 55 residents (24%) indicated that they used EBRS online. Most residents (198 [86%]) attend journal clubs. The most common format is a mandatory meeting held at a special time every month with faculty members with epidemiological and clinical expertise. Residents stated that EBRS articles were used exclusively (13%) or in conjunction with other articles (57%) in their journal clubs. Most respondents (176 of 193 [91%]) stated that journal clubs are very or somewhat valuable to their education.
The EBRS program is widely used among general surgery residents across Canada. Although most residents who use EBRS rate it highly, a large proportion are unaware of EBRS online features. Thus, future efforts to increase awareness of EBRS online features and increase its accessibility are required.
PMCID: PMC3728260  PMID: 23883511
21.  The Enterococcus faecium Enterococcal Biofilm Regulator, EbrB, Regulates the esp Operon and Is Implicated in Biofilm Formation and Intestinal Colonization 
PLoS ONE  2013;8(5):e65224.
Nowadays, Enterococcus faecium is one of the leading nosocomial pathogens worldwide. Strains causing clinical infections or hospital outbreaks are enriched in the enterococcal surface protein (Esp) encoding ICEEfm1 mobile genetic element. Previous studies showed that Esp is involved in biofilm formation, endocarditis and urinary tract infections. In this study, we characterized the role of the putative AraC type of regulator (locus tag EfmE1162_2351), which we renamed ebrB and which is, based on the currently available whole genome sequences, always located upstream of the esp gene, and studied its role in Esp surface exposure during growth. A markerless deletion mutant of ebrB resulted in reduced esp expression and complete abolishment of Esp surface exposure, while Esp cell-surface exposure was restored when this mutant was complemented with an intact copy of ebrB. This demonstrates a role for EbrB in esp expression. However, during growth, ebrB expression levels did not change over time, while an increase in esp expression at both RNA and protein level was observed during mid-log and late-log phase. These results indicate the existence of a secondary regulation system for esp, which might be an unknown quorum sensing system as the enhanced esp expression seems to be cell density dependent. Furthermore, we determined that esp is part of an operon of at least 3 genes putatively involved in biofilm formation. A semi-static biofilm model revealed reduced biofilm formation for the EbrB deficient mutant, while dynamics of biofilm formation using a flow cell system revealed delayed biofilm formation in the ebrB mutant. In a mouse intestinal colonization model the ebrB mutant was less able to colonize the gut compared to wild-type strain, especially in the small intestine. These data indicate that EbrB positively regulates the esp operon and is implicated in biofilm formation and intestinal colonization.
PMCID: PMC3669262  PMID: 23741484
22.  Effects of manganese-excess on CO2 assimilation, ribulose-1,5-bisphosphate carboxylase/oxygenase, carbohydrates and photosynthetic electron transport of leaves, and antioxidant systems of leaves and roots in Citrus grandis seedlings 
BMC Plant Biology  2010;10:42.
Very little is known about the effects of manganese (Mn)-excess on citrus photosynthesis and antioxidant systems. Seedlings of sour pummelo (Citrus grandis) were irrigated for 17 weeks with nutrient solution containing 2 μM (control) or 500 μM (excess) MnSO4. The objective of this study were to understand the mechanisms by which Mn-excess leads to a decrease in CO2 assimilation and to test the hypothesis that Mn-induced changes in antioxidant systems differ between roots and leaves.
Mn-excess decreased CO2 assimilation and stomatal conductance, increased intercellular CO2 concentration, but did not affect chlorophyll (Chl) level. Both initial and total ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) activity in Mn-excess leaves decreased to a lesser extent than CO2 assimilation. Contents of glucose, fructose, starch and total nonstructural carbohydrates did not differ between Mn-excess leaves and controls, while sucrose content was higher in the former. Chl a fluorescence (OJIP) transients from Mn-excess leaves showed increased O-step and decreased P-step, accompanied by positive L- and K-bands. Mn-excess decreased maximum quantum yield of primary photochemistry (Fv/Fm) and total performance index (PItot,abs), but increased relative variable fluorescence at I-steps (VI) and energy dissipation. On a protein basis, Mn-excess leaves displayed higher activities of monodehydroascorbate reductase (MDAR), glutathione reductase (GR), superoxide dismutase (SOD), catalase (CAT) and guaiacol peroxidase (GPX) and contents of antioxidants, similar ascorbate peroxidase (APX) activities and lower dehydroascorbate reductase (DHAR) activities; while Mn-excess roots had similar or lower activities of antioxidant enzymes and contents of antioxidants. Mn-excess did not affect malondialdehyde (MDA) content of roots and leaves.
Mn-excess impaired the whole photosynthetic electron transport chain from the donor side of photosystem II (PSII) up to the reduction of end acceptors of photosystem I (PSI), thus limiting the production of reducing equivalents, and hence the rate of CO2 assimilation. Both the energy dissipation and the antioxidant systems were enhanced in Mn-excess leaves, while the antioxidant systems in Mn-excess roots were not up-regulated, but still remained high activity. The antioxidant systems in Mn-excess roots and leaves provided sufficient protection to them against oxidative damage.
PMCID: PMC2848762  PMID: 20205939
23.  Harvest index, a parameter conditioning responsiveness of wheat plants to elevated CO2  
Journal of Experimental Botany  2013;64(7):1879-1892.
The expansion of the world’s population requires the development of high production agriculture. For this purpose, it is essential to identify target points conditioning crop responsiveness to predicted [CO2]. The aim of this study was to determine the relevance of ear sink strength in leaf protein and metabolomic profiles and its implications in photosynthetic activity and yield of durum wheat plants exposed to elevated [CO2]. For this purpose, a genotype with high harvest index (HI) (Triticum durum var. Sula) and another with low HI (Triticum durum var. Blanqueta) were exposed to elevated [CO2] (700 µmol mol–1 versus 400 µmol mol–1 CO2) in CO2 greenhouses. The obtained data highlighted that elevated [CO2] only increased plant growth in the genotype with the largest HI; Sula. Gas exchange analyses revealed that although exposure to 700 µmol mol–1 depleted Rubisco content, Sula was capable of increasing the light-saturated rate of CO2 assimilation (Asat) whereas, in Blanqueta, the carbohydrate imbalance induced the down-regulation of Asat. The specific depletion of Rubisco in both genotypes under elevated [CO2], together with the enhancement of other proteins in the Calvin cycle, revealed that there was a redistribution of N from Rubisco towards RuBP regeneration. Moreover, the down-regulation of N, NO3 –, amino acid, and organic acid content, together with the depletion of proteins involved in amino acid synthesis that was detected in Blanqueta grown at 700 µmol mol–1 CO2, revealed that inhibition of N assimilation was involved in the carbohydrate imbalance and consequently with the down-regulation of photosynthesis and growth in these plants.
PMCID: PMC3638836  PMID: 23564953
2D proteomic; Acclimation; C/N; CO2; Harvest index; Wheat.
24.  Phenotypic landscape inference reveals multiple evolutionary paths to C4 photosynthesis 
eLife  2013;2:e00961.
C4 photosynthesis has independently evolved from the ancestral C3 pathway in at least 60 plant lineages, but, as with other complex traits, how it evolved is unclear. Here we show that the polyphyletic appearance of C4 photosynthesis is associated with diverse and flexible evolutionary paths that group into four major trajectories. We conducted a meta-analysis of 18 lineages containing species that use C3, C4, or intermediate C3–C4 forms of photosynthesis to parameterise a 16-dimensional phenotypic landscape. We then developed and experimentally verified a novel Bayesian approach based on a hidden Markov model that predicts how the C4 phenotype evolved. The alternative evolutionary histories underlying the appearance of C4 photosynthesis were determined by ancestral lineage and initial phenotypic alterations unrelated to photosynthesis. We conclude that the order of C4 trait acquisition is flexible and driven by non-photosynthetic drivers. This flexibility will have facilitated the convergent evolution of this complex trait.
eLife digest
Plants rely on carbon for their growth and survival: in a process called photosynthesis, they use energy from sunlight to convert carbon dioxide and water into carbohydrates and oxygen gas. The chemical reactions that make up photosynthesis are powered by a chain of enzymes, and plants must ensure that these enzymes—which are in the leaves of the plant—are supplied with enough carbon dioxide and water. Carbon dioxide from the atmosphere enters plants through pores in their leaves, but water must be carried up the plant from the roots.
The type of photosynthesis used by about 90% of flowering plant species—including tomatoes and rice—is called C3 photosynthesis. The first step in this process begins with an enzyme called RuBisCO, which reacts with carbon dioxide and a substance called RuBP to form molecules that contain three carbon atoms (hence the name C3 photosynthesis).
In a hot climate, however, a plant can lose a lot of water through the pores in its leaves: closing these pores allows the plant to retain water, but this also reduces the supply of carbon dioxide. Under these circumstances this causes problems because RuBisCO uses oxygen to break down RuBP, instead of creating sugars, when carbon dioxide is not readily available. To prevent this process, which wastes a lot of energy and resources, some plants—including maize, sugar cane and many other agricultural staples—have evolved an alternative process called C4 photosynthesis. Although it is more complex than C3 photosynthesis, and required many changes to be made to the structure of leaves, C4 photosynthesis has evolved on more than 60 different occasions.
In C4 plants, the mesophyll—the region that is associated with the capture of carbon dioxide by RuBisCO in C3 plants—contains high levels of an alternative enzyme called PEPC that converts carbon dioxide molecules into an acid that contains four carbon atoms. To avoid carbon dioxide being captured by both enzymes, C4 plants evolved to relocate RuBisCO from the mesophyll to a second set of cells in an airtight structure known as the bundle sheath. The four-carbon acids produced by PEPC diffuse to the cells in the bundle sheath, where they are broken down into carbon dioxide molecules, and photosynthesis then proceeds as normal. This process allows photosynthesis to continue when the level of carbon dioxide in the leave is low because the plant has closed its pores to retain water.
Since C4 plants grow faster than C3 plants, and also require less water, plant biologists would like to introduce certain C4 traits into C3 crop plants. To help with this process, Williams, Johnston et al. have used computational methods to explore how C4 photosynthesis evolved from ancestral C3 plants. This involved investigating the prevalence of 16 traits that are common to C4 plants in a total of 73 species that undergo C3 or C4 photosynthesis (including 37 species that possess characteristics of both C3 and C4).
Williams, Johnston et al. then went on to produce a new mathematical model that represents evolutionary processes as pathways across a multi-dimensional “landscape”. The model shows that traits can be acquired in various orders, and that C4 photosynthesis evolved through a number of independent pathways. Some traits that evolved early in the transitions to C4 photosynthesis influenced how evolution proceeded, providing “foundations” upon which further changes evolved.
Interestingly, the structure of the leaf itself appeared to change before any of the photosynthetic enzymes changed. This led Williams, Johnston et al. to conclude that climate change—in particular, the declines in carbon dioxide levels that occurred in prehistoric times—was probably not responsible for the original evolution of C4 photosynthesis. Nevertheless, these results could help with efforts to adapt important C3 crop plants to on-going changes in our climate.
PMCID: PMC3786385  PMID: 24082995
convergent evolution; C4 photosynthesis; Bayesian model; Other
25.  Chloroplastic thioredoxin-f and thioredoxin-m1/4 play important roles in brassinosteroids-induced changes in CO2 assimilation and cellular redox homeostasis in tomato 
Journal of Experimental Botany  2014;65(15):4335-4347.
Virus-induced gene silencing (VIGS) was used in this study to characterize the role of thioredoxin-f and thioredoxin-m1/4 in brassinosteroid-induced changes in CO2 assimilation and cellular redox homeostasis in tomato.
Chloroplast thioredoxins (TRXs) and glutathione function as redox messengers in the regulation of photosynthesis. In this work, the roles of chloroplast TRXs in brassinosteroids (BRs)-induced changes in cellular redox homeostasis and CO2 assimilation were studied in the leaves of tomato plants. BRs-deficient d ^im plants showed decreased transcripts of TRX-f, TRX-m2, TRX-m1/4, and TRX-x, while exogenous BRs significantly induced CO2 assimilation and the expression of TRX-f, TRX-m2, TRX-m1/4, and TRX-x. Virus-induced gene silencing (VIGS) of the chloroplast TRX-f, TRX-m2, TRX-m1/4, and TRX-y genes individually increased membrane lipid peroxidation and accumulation of 2-Cys peroxiredoxin dimers, and decreased the activities of the ascorbate–glutathione cycle enzymes and the ratio of reduced glutathione to oxidized glutathione (GSH/GSSG) in the leaves. Furthermore, partial silencing of TRX-f, TRX-m2, TRX-m1/4, and TRX-y resulted in decreased expression of genes involved in the Benson–Calvin cycle and decreased activity of the associated enzymes. Importantly, the BRs-induced increase in CO2 assimilation and the increased expression and activities of antioxidant- and photosynthesis-related genes and enzymes were compromised in the partially TRX-f- and TRX-m1/4-silenced plants. All of these results suggest that TRX-f and TRX-m1/4 are involved in the BRs-induced changes in CO2 assimilation and cellular redox homeostasis in tomato.
PMCID: PMC4112637  PMID: 24847092
Antioxidant; Benson–Calvin cycle; chloroplast; 2-Cys peroxiredoxin; glutathione; photosynthesis.

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