Jasmonate Synthesis Research Articles

Chloroplast Galactolipids: The Link Between Photosynthesis, Chloroplast Shape, Jasmonates, Phosphate Starvation and Freezing Tolerance.

Monogalactosyldiacylglycerol (MGDG) and digalactosyldiacylglycerol (DGDG) together constitute approximately 80% of chloroplast lipids. Apart from facilitating the photosynthesis light reaction in the thylakoid membrane, these two lipids are important for maintaining chloroplast morphology and for plant survival under abiotic stresses such as phosphate starvation and freezing. Recently it was shown that severe growth retardation phenotypes of the DGDG-deficient mutant dgd1 were due to jasmonate overproduction, linking MGDG and DGDG homeostasis with phytohormone production and suggesting MGDG as a major substrate for jasmonate biosynthesis. Induction of jasmonate synthesis and jasmonic acid (JA) signaling was also observed under conditions of phosphate starvation. We hypothesize that when DGDG is recruited to substitute for phospholipids in extraplastidic membranes during phosphate deficiency, the altered MGDG to DGDG ratio in the chloroplast envelope triggers the conversion of galactolipids into jasmonates. The conversion may contribute to rebalancing the MGDG to DGDG ratio rapidly to maintain chloroplast shape, and jasmonate production can reduce the growth rate and enhance predator deterrence. We also hypothesize that other conditions, such as suppression of dgd1 phenotypes by trigalactosyldiacylglycerol (tgd) mutations, may all be linked to altered jasmonate production, indicating that caution should be exercised when interpreting phenotypes caused by conditions that may alter the MGDG to DGDG ratio at the chloroplast envelope.

Carlos L. Ballaré.

Carlos L. Ballaré.

Verticillium dahliae-Arabidopsis Interaction Causes Changes in Gene Expression Profiles and Jasmonate Levels on Different Time Scales.

Verticillium dahliae is a soil-borne vascular pathogen that causes severe wilt symptoms in a wide range of plants. Co-culture of the fungus with Arabidopsis roots for 24 h induces many changes in the gene expression profiles of both partners, even before defense-related phytohormone levels are induced in the plant. Both partners reprogram sugar and amino acid metabolism, activate genes for signal perception and transduction, and induce defense- and stress-responsive genes. Furthermore, analysis of Arabidopsis expression profiles suggests a redirection from growth to defense. After 3 weeks, severe disease symptoms can be detected for wild-type plants while mutants impaired in jasmonate synthesis and perception perform much better. Thus, plant jasmonates have an important influence on the interaction, which is already visible at the mRNA level before hormone changes occur. The plant and fungal genes that rapidly respond to the presence of the partner might be crucial for early recognition steps and the future development of the interaction. Thus they are potential targets for the control of V. dahliae-induced wilt diseases.

A role for 9-lipoxygenases in maize defense against insect herbivory

ABSTRACTFeeding by Spodoptera exigua (beet armyworm) larvae on Zea mays (maize) induces expression of 9-lipoxygenases to a greater extent than 13-lipoxygenases. Whereas 13-lipoxygenases have an established role in the synthesis of jasmonates that serve as defense signaling molecules in many plant species, relatively little is known about the role of 9-lipoxygenases in herbivore defense. Phylogenetic analysis of lipoxygenases from maize inbred lines B73 and W22 shows that, although most Lox genes are present in both lines, Lox12, a 9-lipoxygenase that has been implicated in fungal defense, is truncated and unlikely to encode a functional protein in W22. Two independent Mutator transposon insertions in another 9-lipoxygenase, Lox4, caused improved S. exigua growth on the mutant lines relative to wildtype W22. This observation suggests a function in herbivore defense for metabolic products downstream of maize Lox4, either through direct toxicity or a perhaps an as yet unknown signaling function.

Molecular mechanisms underlying Grateloupia imbricata (Rhodophyta) carposporogenesis induced by methyl jasmonate.

When applied invitro, methyl jasmonate is sensed by the red seaweed Grateloupia imbricate, substantially and visually affecting its carposporogenesis. However, although there is some understanding of the morphological changes induced by methyl jasmonate invitro, little is known about the genes that are involved in red seaweed carposporogenesis and how their protein products act. For the work reported herein, the expression of genes in red seaweed that encode enzymes involved in the synthesis of methyl jasmonate (jasmonic acid carboxyl methyl transferase and a putative methyl transferase) was monitored. Additionally the genes involved in oxidation (cytochrome P450 and WD40), jasmonate synthesis, signal transduction, and regulation of reactive oxygen species (MYB), and reproduction (ornithine decarboxylase) were monitored. To determine when or if the aforementioned genes were expressed during cystocarp development, fertilized and fertile thalli were exposed to methyl jasmonate and gene expression was measured after 24 and 48h. The results showed that methyl jasmonate promoted differential gene expression in fertilized thalli by 24h and upregulated expression of the ornithine decarboxylase gene only by 48h in fertile thalli (0.75±003 copies·μL-1 at 24h vs. 1.11±0.04 copies·μL-1 at 48h). We conclude that Ornithine decarboxylase expression involves methyl jasmonate signaling as well as development and maturation of cystocarps.

GhJAZ2 attenuates cotton resistance to biotic stresses via the inhibition of the transcriptional activity of GhbHLH171.

Plants have evolved effective mechanisms to protect themselves against multiple stresses, and employ jasmonates (JAs) as vital defence signals to defend against pathogen infection. The accumulation of JA, induced by signals from biotic and abiotic stresses, results in the degradation of Jasmonate-ZIM-domain (JAZ) proteins, followed by the de-repression of JAZ-repressed transcription factors (such as MYC2) to activate defence responses and developmental processes. Here, we characterized a JAZ family protein, GhJAZ2, from cotton (Gossypium hirsutum) which was induced by methyl jasmonate (MeJA) and inoculation of Verticillium dahliae. The overexpression of GhJAZ2 in cotton impairs the sensitivity to JA, decreases the expression level of JA-response genes (GhPDF1.2 and GhVSP) and enhances the susceptibility to V.dahliae and insect herbivory. Yeast two-hybrid and bimolecular fluorescence complementation assays showed that GhJAZ2 may be involved in the regulation of cotton disease resistance by interaction with further disease-response proteins, such as pathogenesis-related protein GhPR10, dirigent-like protein GhD2, nucleotide-binding site leucine-rich repeat (NBS-LRR) disease-resistant protein GhR1 and a basic helix-loop-helix transcription factor GhbHLH171. Unlike MYC2, overexpression of GhbHLH171 in cotton activates the JA synthesis and signalling pathway, and improves plant tolerance to the fungus V.dahliae. Molecular and genetic evidence shows that GhJAZ2 can interact with GhbHLH171 and inhibit its transcriptional activity and, as a result, can restrain the JA-mediated defence response. This study provides new insights into the molecular mechanisms of GhJAZ2 in the regulation of the cotton defence response.

Transcriptome and phytohormone analysis reveals a comprehensive phytohormone and pathogen defence response in pear self-/cross-pollination

Key messageCandidate genes were identified and the role of phytohormones such as JA-Me and ABA in the synthesis of S-RNase was emphasized in pear self-incompatibility.Self-incompatibility (SI) occurs widely in flowering plants as an intraspecific reproductive barrier. This phenomenon promotes variation within species, but for some species such as Pyrus, SI is a nuisance rather than a benefit in agricultural production. Although many studies have been conducted on SI in pears, its mechanism remains unclear. In this study, high-throughput Illumina RNA sequencing (RNA-seq) was used to identify SI-related genes in pear styles. Using transcriptome comparisons, differentially expressed genes of unpollinated (UP), cross-pollinated (CP), and self-pollinated (SP) styles were identified after 48 h. A total of 1796 and 1890 genes were identified in DSC (UP vs. CP) and DSI (UP vs. SP), respectively. KEGG analysis revealed that genes involved in the “plant hormone signal transduction pathway” and “plant–pathogen interaction pathway” were significantly enriched in DSI (UP vs. SP) compared to those in DSC (UP vs. CP). The expression level of S-glycoprotein ribonuclease (S-RNase) was dramatically reduced in cross-pollinated (CP) styles. To better understand the relationship between the expression patterns of S-RNase and two major KEGG pathways, the concentrations of phytohormones were measured, and the expression pattern of S-RNase was analysed using qRT-PCR. Our results demonstrate that methyl jasmonate and abscisic acid may enhance the expression level of S-RNase, and pollination can affect the synthesis of methyl jasmonate and abscisic acid in pear styles. Overall, this study is a global transcriptome analysis of SI in pear. A relationship between self-rejection, plant hormones, and pathogen defence was shown in pear.

Regulation of growth-defense balance by the JASMONATE ZIM-DOMAIN (JAZ)-MYC transcriptional module.

Summary The plant hormone jasmonate (JA) promotes the degradation of JASMONATE ZIM‐DOMAIN (JAZ) proteins to relieve repression on diverse transcription factors (TFs) that execute JA responses. However, little is known about how combinatorial complexity among JAZ–TF interactions maintains control over myriad aspects of growth, development, reproduction, and immunity.We used loss‐of‐function mutations to define epistatic interactions within the core JA signaling pathway and to investigate the contribution of MYC TFs to JA responses in Arabidopsis thaliana.Constitutive JA signaling in a jaz quintuple mutant (jazQ) was largely eliminated by mutations that block JA synthesis or perception. Comparison of jazQ and a jazQ myc2 myc3 myc4 octuple mutant validated known functions of MYC2/3/4 in root growth, chlorophyll degradation, and susceptibility to the pathogen Pseudomonas syringae. We found that MYC TFs also control both the enhanced resistance of jazQ leaves to insect herbivory and restricted leaf growth of jazQ. Epistatic transcriptional profiles mirrored these phenotypes and further showed that triterpenoid biosynthetic and glucosinolate catabolic genes are up‐regulated in jazQ independently of MYC TFs.Our study highlights the utility of genetic epistasis to unravel the complexities of JAZ–TF interactions and demonstrates that MYC TFs exert master control over a JAZ‐repressible transcriptional hierarchy that governs growth–defense balance.

Acetate-mediated novel survival strategy against drought in plants.

Water deficit caused by global climate changes seriously endangers the survival of organisms and crop productivity, and increases environmental deterioration1,2. Plants' resistance to drought involves global reprogramming of transcription, cellular metabolism, hormone signalling and chromatin modification3-8. However, how these regulatory responses are coordinated via the various pathways, and the underlying mechanisms, are largely unknown. Herein, we report an essential drought-responsive network in which plants trigger a dynamic metabolic flux conversion from glycolysis into acetate synthesis to stimulate the jasmonate (JA) signalling pathway to confer drought tolerance. In Arabidopsis, the ON/OFF switching of this whole network is directly dependent on histone deacetylase HDA6. In addition, exogenous acetic acid promotes de novo JA synthesis and enrichment of histone H4 acetylation, which influences the priming of the JA signalling pathway for plant drought tolerance. This novel acetate function is evolutionarily conserved as a survival strategy against environmental changes in plants. Furthermore, the external application of acetic acid successfully enhanced the drought tolerance in Arabidopsis, rapeseed, maize, rice and wheat plants. Our findings highlight a radically new survival strategy that exploits an epigenetic switch of metabolic flux conversion and hormone signalling by which plants adapt to drought.

Key metabolic traits of Pisum sativum maintain cell vitality during Didymella pinodes infection: cultivar resistance and the microsymbionts' influence

Ascochyta blight causes severe losses in field pea production and the search for resistance traits towards the causal agent Didymella pinodes is of particular importance for farmers. Various microsymbionts have been reported to shape the plants' immune response. However, regardless their contribution to resistance, they are hardly included in experimental designs. We delineate the effect of symbionts (rhizobia, mycorrhiza) on the leaf proteome and metabolome of two field pea cultivars with varying resistance levels against D. pinodes and, furthermore, show cultivar specific symbiont colonisation efficiency. The pathogen infection showed a stronger influence on the interaction with the microsymbionts in the susceptible cultivar, which was reflected in decreased nodule weight and root mycorrhiza colonisation. Vice versa, symbionts induced variation of the host's infection response which, however, was overruled by genotypic resistance associated traits of the tolerant cultivar such as maintenance of photosynthesis and provision of sugars and carbon back bones to fuel secondary metabolism. Moreover, resistance appears to be linked to sulphur metabolism, a functional glutathione-ascorbate hub and fine adjustment of jasmonate and ethylene synthesis to suppress induced cell death. We conclude that these metabolic traits are essential for sustainment of cell vitality and thus, a more efficient infection response. SignificanceThe infection response of two Pisum sativum cultivars with varying resistance levels towards Didymella pinodes was analysed most comprehensively at proteomic and metabolomic levels. Enhanced tolerance was linked to newly discovered cultivar specific metabolic traits such as hormone synthesis and presumably suppression of cell death.

Exogenous application of methyl jasmonate induces defence against Meloidogyne hapla in soybean

Phytohormones play important roles in plant defence against plant-parasitic nematodes, although the role of jasmonate (JA) in defence against root-knot nematodes (RKN, Meloidogyne spp.) in soybean (Glycine max) was unknown. In this study, two commercial soybean cultivars, cvs DongSheng1 (DS1) and SuiNong14 (SN14), were identified as susceptible and resistant, respectively, to M. hapla. Quantitative reverse transcription (qRT)-PCR analysis showed that the expression of genes involved in JA synthesis or signalling was significantly induced in both susceptible and resistant roots at 24 and 48 h after inoculation. Exogenous application of methyl jasmonate induced defence against RKN in susceptible cv. DS1, which might be involved in altered activities of defence-related enzymes (chitinase and β-1,3 glucanase) and pathogenesis-related gene PR5 expression. The results indicate that exogenous application of JA might be an alternative strategy to induce soybean resistance against RKN.

Jasmonates: Multifunctional Roles in Stress Tolerance.

Jasmonates (JAs) [Jasmonic acid (JA) and methyl jasmonates (MeJAs)] are known to take part in various physiological processes. Exogenous application of JAs so far tested on different plants under abiotic stresses particularly salinity, drought, and temperature (low/high) conditions have proved effective in improving plant stress tolerance. However, its extent of effectiveness entirely depends on the type of plant species tested or its concentration. The effects of introgression or silencing of different JA- and Me-JA-related genes have been summarized in this review, which have shown a substantial role in improving crop yield and quality in different plants under stress or non-stress conditions. Regulation of JAs synthesis is impaired in stressed as well as unstressed plant cells/tissues, which is believed to be associated with a variety of metabolic events including signal transduction. Although, mitogen activated protein kinases (MAPKs) are important components of JA signaling and biosynthesis pathways, nitric oxide, ROS, calcium, ABA, ethylene, and salicylic acid are also important mediators of plant growth and development during JA signal transduction and synthesis. The exploration of other signaling molecules can be beneficial to examine the details of underlying molecular mechanisms of JA signal transduction. Much work is to be done in near future to find the proper answers of the questions like action of JA related metabolites, and identification of universal JA receptors etc. Complete signaling pathways involving MAPKs, CDPK, TGA, SIPK, WIPK, and WRKY transcription factors are yet to be investigated to understand the complete mechanism of action of JAs.

Route Scouting towards a Methyl Jasmonate Precursor

AbstractFor the synthesis of methyl jasmonate (1), via the strategic intermediates 3, 4, and 6a, we constructed a synthetic network via the diverse intermediates 7–10, 13, 14, 17, and 18. This allowed us to compare the efficiency of more than 20 novel routes. The most productive pathway with a total yield of 38% is represented by the sequence→5a→5m→13b→13a→6a→4 and proceeds via sequential bromination, basic elimination, decarbomethoxylation, isomerization, and finally Lindlar hydrogenation. The shortest selective way, 2a→[(E,E)‐12b]→3→4, is a two‐pot sequence using a modification of Naef's method, based on an aldol condensation between inexpensive cyclopentanone (2a) and crotonaldehyde, with in situ CoreyChaykovsky cyclopropanation under phase transfer conditions. The key intermediate 3 was then simply pyrolyzed to afford 4 in 27% total yield. The alternative isomerization method via the six‐step deviation→5a→5c→8c→13a→6a→4 was longer, although more efficient, with a total yield of 32%. Alternatively, a yield of 34% was obtained via the five‐step sequence→5a→5c→2h→2i→4. Another favored six‐step pathway,→5a→5c→2h→17a→14a→4 afforded the target compound in 35% total yield.

Control of Carbon Assimilation and Partitioning by Jasmonate: An Accounting of Growth-Defense Tradeoffs.

Plant growth is often constrained by the limited availability of resources in the microenvironment. Despite the continuous threat of attack from insect herbivores and pathogens, investment in defense represents a lost opportunity to expand photosynthetic capacity in leaves and absorption of nutrients and water by roots. To mitigate the metabolic expenditure on defense, plants have evolved inducible defense strategies. The plant hormone jasmonate (JA) is a key regulator of many inducible defenses. Synthesis of JA in response to perceived danger leads to the deployment of a variety of defensive structures and compounds, along with a potent inhibition of growth. Genetic studies have established an important role for JA in mediating tradeoffs between growth and defense. However, several gaps remain in understanding of how JA signaling inhibits growth, either through direct transcriptional control of JA-response genes or crosstalk with other signaling pathways. Here, we highlight recent progress in uncovering the role of JA in controlling growth-defense balance and its relationship to resource acquisition and allocation. We also discuss tradeoffs in the context of the ability of JA to promote increased leaf mass per area (LMA), which is a key indicator of leaf construction costs and leaf life span.

Physiological and Molecular Alterations Promoted by Schizotetranychus oryzae Mite Infestation in Rice Leaves.

Infestation of phytophagous mite Schizotetranychus oryzae in rice causes critical yield losses. To better understand this interaction, we employed Multidimensional Protein Identification Technology (MudPIT) approach to identify differentially expressed proteins. We detected 18 and 872 unique proteins in control and infested leaves, respectively, along with 32 proteins more abundant in control leaves. S. oryzae infestation caused decreased abundance of proteins related to photosynthesis (mostly photosystem II-related), carbon assimilation and energy production, chloroplast detoxification, defense, and fatty acid and gibberellin synthesis. On the contrary, infestation caused increased abundance of proteins involved in protein modification and degradation, gene expression at the translation level, protein partitioning to different organelles, lipid metabolism, actin cytoskeleton remodeling, and synthesis of jasmonate, amino acid, and molecular chaperones. Our results also suggest that S. oryzae infestation promotes cell-wall remodeling and interferes with ethylene biosynthesis in rice leaves. Proteomic data were positively correlated with enzymatic assays and RT-qPCR analysis. Our findings describe the protein expression patterns of infested rice leaves and suggest that the acceptor side of PSII is probably the major damaged target in the photosynthetic apparatus. These data will be useful in future biotechnological approaches aiming to induce phytophagous mite resistance in rice.

A gain-of-function mutation in Msl10 triggers cell death and wound-induced hyperaccumulation of jasmonic acid in Arabidopsis.

Jasmonates (JAs) are rapidly induced after wounding and act as key regulators for wound induced signaling pathway. However, what perceives the wound signal and how that triggers JA biosynthesis remains poorly understood. To identify components involved in Arabidopsis wound and JA signaling pathway, we screened for mutants with abnormal expression of a luciferase reporter, which is under the control of a wound-responsive promoter of an ethylene response factor (ERF) transcription factor gene, RAP2.6 (Related to APetala 2.6). The rea1 (RAP2.6 expresser in shoot apex) mutant constitutively expressed the RAP2.6-LUC reporter gene in young leaves. Along with the typical JA phenotypes including shorter petioles, loss of apical dominance, accumulation of anthocyanin pigments and constitutive expression of JA response gene, rea1 plants also displayed cell death and accumulated high levels of JA in response to wounding. The phenotype of rea1 mutant is caused by a gain-of-function mutation in the C-terminus of a mechanosensitive ion channel MscS-like 10 (MSL10). MSL10 is localized in the plasma membrane and is expressed predominantly in root tip, shoot apex and vascular tissues. These results suggest that MSL10 is involved in the wound-triggered early signal transduction pathway and possibly in regulating the positive feedback synthesis of JA.

Transcriptome sequencing and analysis of rubber tree (Hevea brasiliensis Muell.) to discover putative genes associated with tapping panel dryness (TPD).

BackgroundTapping panel dryness (TPD) involves in the partial or complete cessation of latex flow thus seriously affect latex production in rubber tree (Hevea brasiliensis). Numerous studies have been conducted to define its origin and nature, but the molecular nature and mechanism of TPD occurrence remains unknown. This study is committed to de novo sequencing and comparative analysis of the transcriptomes of healthy (H) and TPD-affected (T) rubber trees to identify the genes and pathways related to the TPD.ResultsTotal raw reads of 34,632,012 and 35,913,020 bp were obtained from H and T library, respectively using Illumina Hiseq 2000 sequencing technology. De novo assemblies yielded 141,456 and 169,285 contigs, and 96,070 and 112,243 unigenes from H and T library, respectively.Among 73597 genes, 22577 genes were identified as differential expressed genes between H and T library via comparative transcript profiling. A majority of genes involved in natural rubber biosynthesis and jasmonate synthesis with most potential relevance in TPD occurrence were found to be differentially expressed.ConclusionsIn TPD-affected trees, the expression of most genes related to the latex biosynthesis and jasmonate synthesis was severely inhibited and is probably the direct cause of the TPD. These new de novo transcriptome data sets provide a significant resource for the discovery of genes related to TPD and improve our understanding of the occurrence and maintainace of TPD.Electronic supplementary materialThe online version of this article (doi:10.1186/s12864-015-1562-9) contains supplementary material, which is available to authorized users.

Molecular cloning and sequence analysis of lipoxygenase (AsLOX1) gene in Aquilaria sinensis (Lour.) Gilg and its expression responding to MeJA and crush wounding treatment

Lipoxygenase (LOX) not only acts as a key enzyme in the octadecanoid pathway and jasmonate (JA) synthesis, but also plays a significant role in plant growth and development, especially plant defense. The LOX gene, however, remains unclear in Aquilaria sinensis, a “wounding-induced-type” medicinal plant with great economic value. Based on the transcriptome data, a full-length cDNA sequence of LOX gene (termed as AsLOX1) was cloned for A. sinensis by RT-PCR and RACE cloning strategy. The sequence analysis showed that the AsLOX1 was 3168bp in length (GenBank accession number: KJ868174), and contained a complete open frame (ORF) of 2754bp. The deduced protein is composed of 917 amino acids, with a predicted molecular weight of 103.55kDa and an isoelectric point of 6.70. The amino acid sequence of AsLOX1 was determined to have nearly 80% sequence identity with the LOX genes of several other species by multiple alignments, suggesting that LOX belongs to a kind of conservative proteins. The deduced AsLOX1 protein was predicted to possess PLAT_LH2 and the lipoxygenase superfamily domain. The phylogenetic tree analysis reveals that AsLOX1 is more closely related to Theobroma cacao LOX than to those of other plants. According to the tissue-specific expression pattern analysis, the LOX gene in A. sinensis tissues is mainly expressed in roots and stems, the main agarwood formation parts. In both the methyl jasmonate (MeJA) and crush wounding treatment, the expression of AsLOX1 was observed to be significantly induced in A. sinensis calli, which was relatively high at 1h, reached the peak at 6h, and then declined rapidly to the normal level. These results demonstrate that AsLOX1 gene is an early damage-responsive gene, and has its expression modulated in response to MeJA stress and wounding, indicating the role of AsLOX1 in defense responses to wounding-induced agarwood formation in A. sinensis.

JASMONIC ACID: ROLE IN BIOTECHNOLOGY AND THE REGULATION OF PLANTS BIOCHEMICAL PROCESSES

Published data and results of our studies concerning the involvement and role of jasmonic acid in the regulation of plant physiological and biochemical processes have been analyzed and summarized. The basic stages of jasmonates synthesis are reviewed. Properties of enzymes involved in jasmonate biosynthesis are described. Data on the jasmonic acid involvement in the regulation of seed germination, maintaining of aging processes, sex determination, cellulose synthesis, features of interaction with ABA as well as in gene expression, formation of the immune system in stress and pathogene effect conditions are presented. Jasmonic acid effects on the cell ultrustructure are discussed. Prospects of using jasmonates in biotechnology are dealt with.

Glutathione application affects the transcript profile of genes in Arabidopsis seedling

Glutathione (GSH), a tripeptide thiol compound has multiple functions in plants. Recent works suggested that GSH plays a regulatory role in signaling in plants as part of their adaptation to stress. To better understand the role of GSH as a regulatory molecule, 14 days old Arabidopsis thaliana seedlings were treated with 5mM of GSH for 4h. Changes in gene expression patterns were studied by cDNA microarray analysis. The expression of 453 genes was significantly changed compared to the untreated control, of which 261 genes were up-regulated and 192 genes were down-regulated. Genes from several groups were affected, including those of sulfur metabolism, degradation and synthesis of macromolecules and transcription factors. Up-regulation of genes involved in responses to biotic stresses, or in jasmonate or salicylic acid synthesis and their signaling, suggests that GSH triggers genes that help protect the plants during stresses. In addition, GSH down regulated genes involved in plant growth and development, like those involved in cell wall synthesis and its extension, and genes associated with auxin and cytokinins response, which are related to growth and development of the plants. The results suggest that GSH might have a role in response to biotic stress by initiating defense responses and modifying plants’ growth and development in an effort to tune their sessile lifestyle of plants to environmental constraints.