Jasmonate Response Research Articles

Pipecolic Acid Orchestrates Plant Systemic Acquired Resistance and Defense Priming via Salicylic Acid-Dependent and -Independent Pathways.

We investigated the relationships of the two immune-regulatory plant metabolites, salicylic acid (SA) and pipecolic acid (Pip), in the establishment of plant systemic acquired resistance (SAR), SAR-associated defense priming, and basal immunity. Using SA-deficient sid2, Pip-deficient ald1, and sid2 ald1 plants deficient in both SA and Pip, we show that SA and Pip act both independently from each other and synergistically in Arabidopsis thaliana basal immunity to Pseudomonas syringae. Transcriptome analyses reveal that SAR establishment in Arabidopsis is characterized by a strong transcriptional response systemically induced in the foliage that prepares plants for future pathogen attack by preactivating multiple stages of defense signaling and that SA accumulation upon SAR activation leads to the downregulation of photosynthesis and attenuated jasmonate responses systemically within the plant. Whereas systemic Pip elevations are indispensable for SAR and necessary for virtually the whole transcriptional SAR response, a moderate but significant SA-independent component of SAR activation and SAR gene expression is revealed. During SAR, Pip orchestrates SA-dependent and SA-independent priming of pathogen responses in a FLAVIN-DEPENDENT-MONOOXYGENASE1 (FMO1)-dependent manner. We conclude that a Pip/FMO1 signaling module acts as an indispensable switch for the activation of SAR and associated defense priming events and that SA amplifies Pip-triggered responses to different degrees in the distal tissue of SAR-activated plants.

Recent insights into the molecular mechanism of jasmonate signaling during insect-plant interaction

Amongst all the phytohormones, jasmonates are most important signaling molecule associated with plant defense against herbivores as they activate the expression of both direct and indirect defenses. Jasmonates are produced at insect-infested local sites and either jasmonate itself or some Jasmonic Acid (JA)-elicited compound travels to systemic leaves and elicits defense response. Upon herbivory, receptors/sensors of the host plant recognize Herbivory Associated Molecular Patterns (HAMPs) and early signaling events start with the involvement of Mitogen Activated Protein Kinases (MAPKs). JA biosynthesis is initiated in the chloroplast with the hydrolysis of chloroplast membrane lipids by phospholipases releasing free α-linolenic acid (α-LA), which gets converted to 12-oxo-phytodienoic acid (OPDA). OPDA is subsequently transported to peroxisomes where JA is produced by β-oxidation and it conjugates with isoleucine by JA amino acid synthetase (JAR) enzymes. Once high levels of JA-Isoleucine (Ile) are achieved, JA-induced response is instigated by SCF complex (a complex consisting of Skp1, Cullin-1 and F-box protein). JA-Ile forms a complex to form JA-Ile-COI1 (Coronatine insensitive Protein1) that recognizes a target protein Jasmonate ZIM domain (JAZ1), which is ubiquitinated and subsequently subjected to proteasomal degradation releasing the repression of MYC2, thereby activating expression of JA responsive genes. Further, both Novel Interactor of JAZ (NINJA) and Topless (TPL) proteins function as negative regulators of jasmonate responses. Although there has been remarkable progress in recent years, many open questions remain to be answered regarding jasmonate signaling during herbivory.

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.

The NAC transcription factor family in maritime pine (Pinus Pinaster): molecular regulation of two genes involved in stress responses.

BackgroundNAC transcription factors comprise a large plant-specific gene family involved in the regulation of diverse biological processes. Despite the growing number of studies on NAC transcription factors in various species, little information is available about this family in conifers. The goal of this study was to identify the NAC transcription family in maritime pine (Pinus pinaster), to characterize ATAF-like genes in response to various stresses and to study their molecular regulation.MethodsWe have isolated two maritime pine NAC genes and using a transient expression assay in N. benthamiana leaves estudied the promoter jasmonate response.ResultsIn this study, we identified 37 NAC genes from maritime pine and classified them into six main subfamilies. The largest group includes 12 sequences corresponding to stress-related genes. Two of these NAC genes, PpNAC2 and PpNAC3, were isolated and their expression profiles were examined at various developmental stages and in response to various types of stress. The expression of both genes was strongly induced by methyl jasmonate (MeJA), mechanical wounding, and high salinity. The promoter regions of these genes were shown to contain cis-elements involved in the stress response and plant hormonal regulation, including E-boxes, which are commonly found in the promoters of genes that respond to jasmonate, and binding sites for bHLH proteins. Using a transient expression assay in N. benthamiana leaves, we found that the promoter of PpNAC3 was rapidly induced upon MeJA treatment, while this response disappeared in plants in which the transcription factor NbbHLH2 was silenced.ConclusionOur results suggest that PpNAC2 and PpNAC3 encode stress-responsive NAC transcription factors involved in the jasmonate response in pine. Furthermore, these data also suggest that the jasmonate signaling pathway is conserved between angiosperms and gymnosperms. These findings may be useful for engineering stress tolerance in pine via biotechnological approaches.Electronic supplementary materialThe online version of this article (doi:10.1186/s12870-015-0640-0) contains supplementary material, which is available to authorized users.

Arabidopsis thaliana AtGCN2 Kinase is Involved in Disease Resistance against Pathogens with Diverse Life Styles

The ability of the plants to detect diverse stress conditions and initiate cellular responses is vital to their survival in a constantly changing environment. General regulatory molecules often play crucial roles in controlling a multitude of cellular processes throughout the life span of an organism. GCN2 (general control nonderepressible 2) is a serine/threonine-protein kinase that acts as a global translational regulator in all eukaryotes from yeast to mammals to plants. GCN2 plays universal roles in mitigating cellular stresses by directly binding with uncharged tRNAs and phosphorylating its target, eukaryotic initiation factor 2 alpha (eIF2α). Here, we demonstrate that Arabidopsis thaliana GCN2 (AtGCN2) serves as a general regulator of salicylic acid- and jasmonic acid-mediated immune responses triggered upon infection with biotrophic and necrotrophic pathogens. Intriguingly, we found examples of both positive and negative influence of AtGCN2 on plant immunity at different developmental stages. This effect is consistent with the variable amount of abscisic acid accumulation in plants lacking functional AtGCN2 at early stages of development. Finally, we illustrate that AtGCN2 positively contributes to water loss and might also be involved in the epidermis-mediated defense responses.

The nexus between growth and defence signalling: auxin and cytokinin modulate plant immune response pathways.

Plants deploy a finely tuned balance between growth and defence responses for better fitness. Crosstalk between defence signalling hormones such as salicylic acid (SA) and jasmonates (JAs) as well as growth regulators plays a significant role in mediating the trade-off between growth and defence in plants. Here, we specifically discuss how the mutual antagonism between the signalling of auxin and SA impacts on plant growth and defence. Furthermore, the synergism between auxin and JA benefits a class of plant pathogens. JA signalling also poses growth cuts through auxin. We discuss how the effect of cytokinins (CKs) is multifaceted and is effective against a broad range of pathogens in mediating immunity. The synergism between CKs and SA promotes defence against biotrophs. Reciprocally, SA inhibits CK-mediated growth responses. Recent reports show that CKs promote JA responses; however, in a feedback loop, JA suppresses CK responses. We also highlight crosstalk between auxin and CKs and discuss their antagonistic effects on plant immunity. Efforts to minimize the negative effects of auxin on immunity and a reduction in SA- and JA-mediated growth losses should lead to better sustainable plant protection strategies.

Multilayered Organization of Jasmonate Signalling in the Regulation of Root Growth.

Physical damage can strongly affect plant growth, reducing the biomass of developing organs situated at a distance from wounds. These effects, previously studied in leaves, require the activation of jasmonate (JA) signalling. Using a novel assay involving repetitive cotyledon wounding in Arabidopsis seedlings, we uncovered a function of JA in suppressing cell division and elongation in roots. Regulatory JA signalling components were then manipulated to delineate their relative impacts on root growth. The new transcription factor mutant myc2-322B was isolated. In vitro transcription assays and whole-plant approaches revealed that myc2-322B is a dosage-dependent gain-of-function mutant that can amplify JA growth responses. Moreover, myc2-322B displayed extreme hypersensitivity to JA that totally suppressed root elongation. The mutation weakly reduced root growth in undamaged plants but, when the upstream negative regulator NINJA was genetically removed, myc2-322B powerfully repressed root growth through its effects on cell division and cell elongation. Furthermore, in a JA-deficient mutant background, ninja1 myc2-322B still repressed root elongation, indicating that it is possible to generate JA-responses in the absence of JA. We show that NINJA forms a broadly expressed regulatory layer that is required to inhibit JA signalling in the apex of roots grown under basal conditions. By contrast, MYC2, MYC3 and MYC4 displayed cell layer-specific localisations and MYC3 and MYC4 were expressed in mutually exclusive regions. In nature, growing roots are likely subjected to constant mechanical stress during soil penetration that could lead to JA production and subsequent detrimental effects on growth. Our data reveal how distinct negative regulatory layers, including both NINJA-dependent and -independent mechanisms, restrain JA responses to allow normal root growth. Mechanistic insights from this work underline the importance of mapping JA signalling components to specific cell types in order to understand and potentially engineer the growth reduction that follows physical damage.

Elevated levels of CYP94 family gene expression alleviate the jasmonate response and enhance salt tolerance in rice.

The plant hormone jasmonate and its conjugates (JAs) have important roles in growth control, leaf senescence and defense responses against insects and microbial attacks. JA biosynthesis is induced by several stresses, including mechanical wounding, pathogen attacks, drought and salinity stresses. However, the roles of JAs under abiotic stress conditions are unclear. Here we report that increased expression of the Cyt P450 family gene CYP94C2b enhanced viability of rice plants under saline conditions. This gene encodes an enzyme closely related to CYP94C1 that catalyzes conversion of bioactive jasmonate-isoleucine (JA-Ile) into 12OH-JA-Ile and 12COOH-JA-Ile. Inactivation of JA was facilitated in a rice line with enhanced CYP94C2b expression, and responses to exogenous JA and wounding were alleviated. Moreover, salt stress-induced leaf senescence but not natural senescence was delayed in the transgenic rice. These results suggest that bioactive JAs have a negative effect on viability under salt stress conditions and demonstrate that manipulating JA metabolism confers enhanced salt tolerance in rice.

RNAseq reveals weed-induced PIF3-like as a candidate target to manipulate weed stress response in soybean.

Weeds reduce yield in soybeans (Glycine max) through incompletely defined mechanisms. The effects of weeds on the soybean transcriptome were evaluated in field conditions during four separate growing seasons. RNASeq data were collected from six biological samples of soybeans growing with or without weeds. Weed species and the methods to maintain weed-free controls varied between years to mitigate treatment effects, and to allow detection of general soybean weed responses. Soybean plants were not visibly nutrient- or water-stressed. We identified 55 consistently downregulated genes in weedy plots. Many of the downregulated genes were heat shock genes. Fourteen genes were consistently upregulated. Several transcription factors including a PHYTOCHROME INTERACTING FACTOR 3-like gene (PIF3) were included among the upregulated genes. Gene set enrichment analysis indicated roles for increased oxidative stress and jasmonic acid signaling responses during weed stress. The relationship of this weed-induced PIF3 gene to genes involved in shade avoidance responses in Arabidopsis provide evidence that this gene may be important in the response of soybean to weeds. These results suggest that the weed-induced PIF3 gene will be a target for manipulating weed tolerance in soybean.

Induced jasmonate signaling leads to contrasting effects on root damage and herbivore performance.

Induced defenses play a key role in plant resistance against leaf feeders. However, very little is known about the signals that are involved in defending plants against root feeders and how they are influenced by abiotic factors. We investigated these aspects for the interaction between rice (Oryza sativa) and two root-feeding insects: the generalist cucumber beetle (Diabrotica balteata) and the more specialized rice water weevil (Lissorhoptrus oryzophilus). Rice plants responded to root attack by increasing the production of jasmonic acid (JA) and abscisic acid, whereas in contrast to in herbivore-attacked leaves, salicylic acid and ethylene levels remained unchanged. The JA response was decoupled from flooding and remained constant over different soil moisture levels. Exogenous application of methyl JA to the roots markedly decreased the performance of both root herbivores, whereas abscisic acid and the ethylene precursor 1-aminocyclopropane-1-carboxylic acid did not have any effect. JA-deficient antisense 13-lipoxygenase (asLOX) and mutant allene oxide cyclase hebiba plants lost more root biomass under attack from both root herbivores. Surprisingly, herbivore weight gain was decreased markedly in asLOX but not hebiba mutant plants, despite the higher root biomass removal. This effect was correlated with a herbivore-induced reduction of sucrose pools in asLOX roots. Taken together, our experiments show that jasmonates are induced signals that protect rice roots from herbivores under varying abiotic conditions and that boosting jasmonate responses can strongly enhance rice resistance against root pests. Furthermore, we show that a rice 13-lipoxygenase regulates root primary metabolites and specifically improves root herbivore growth.

Responses to intermittent exposure to green light during the dark period in <i>Arabidopsis thaliana</i> and possible involvement of cryptochrome 2

The effects of green light treatment during the dark period were examined in Arabidopsis thaliana as a first step to understanding the mechanism of artificial green light effects in plants. Plants were grown under long-day conditions (an 18-h light and a 6-h dark cycle) and were intermittently exposed to green light during the dark period for 2 h 3 times a week. This green light treatment suppressed the elongation of roots and hypocotyls in wild-type plants. However, the green light-induced changes were not significant in the cry2 mutant that is deficient in the blue light receptor cryptochrome 2. The green light treatment elevated both jasmonic acid and salicylic acid levels in the wild-type plants but the elevation of the jasmonic acid level was impaired in the cry2 mutant plants. These results suggest that intermittent exposure to green light triggers artificial responses in Arabidopsis plants that do not occur in the natural environment, and that cryptochrome 2-dependent and jasmonic acid-mediated responses may be partly involved in the effect of green light on plants.

Cloning and characterisation of JAZ gene family in Hevea brasiliensis.

Mechanical wounding or treatment with exogenous jasmonates (JA) induces differentiation of the laticifer in Hevea brasiliensis. JA is a key signal for latex biosynthesis and wounding response in the rubber tree. Identification of JAZ (jasmonate ZIM-domain) family of proteins that repress JA responses has facilitated rapid progress in understanding how this lipid-derived hormone controls gene expression and related physiological processes in plants. In this work, the full-length cDNAs of six JAZ genes were cloned from H.brasiliensis (termed HbJAZ). These HbJAZ have different lengths and sequence diversity, but all of them contain Jas and ZIM domains, and two of them contain an ERF-associated amphiphilic repression (EAR) motif in the N-terminal. Real-time RT-PCR analyses revealed that HbJAZ have different expression patterns and tissue specificity. Four HbJAZ were up-regulated, one was down-regulated, while two were less effected by rubber tapping treatment, suggesting that they might play distinct roles in the wounding response. A yeast two-hybrid assay revealed that HbJAZ proteins interact with each other to form homologous or heterogeneous dimer complexes, indicating that the HbJAZ proteins may expand their function through diverse JAZ-JAZ interactions. This work lays a foundation for identification of the JA signalling pathway and molecular mechanisms of latex biosynthesis in rubber trees.

Endoplasmic Reticulum-associated Inactivation of the Hormone Jasmonoyl-l-Isoleucine by Multiple Members of the Cytochrome P450 94 Family in Arabidopsis

The plant hormone jasmonate (JA) controls diverse aspects of plant immunity, growth, and development. The amplitude and duration of JA responses are controlled in large part by the intracellular level of jasmonoyl-L-isoleucine (JA-Ile). In contrast to detailed knowledge of the JA-Ile biosynthetic pathway, little is known about enzymes involved in JA-Ile metabolism and turnover. Cytochromes P450 (CYP) 94B3 and 94C1 were recently shown to sequentially oxidize JA-Ile to hydroxy (12OH-JA-Ile) and dicarboxy (12COOH-JA-Ile) derivatives. Here, we report that a third member (CYP94B1) of the CYP94 family also participates in oxidative turnover of JA-Ile in Arabidopsis. In vitro studies showed that recombinant CYP94B1 converts JA-Ile to 12OH-JA-Ile and lesser amounts of 12COOH-JA-Ile. Consistent with this finding, metabolic and physiological characterization of CYP94B1 loss-of-function and overexpressing plants demonstrated that CYP94B1 and CYP94B3 coordinately govern the majority (>95%) of 12-hydroxylation of JA-Ile in wounded leaves. Analysis of CYP94-promoter-GUS reporter lines indicated that CYP94B1 and CYP94B3 serve unique and overlapping spatio-temporal roles in JA-Ile homeostasis. Subcellular localization studies showed that CYP94s involved in conversion of JA-Ile to 12COOH-JA-Ile reside on endoplasmic reticulum (ER). In vitro studies further showed that 12COOH-JA-Ile, unlike JA-Ile, fails to promote assembly of COI1-JAZ co-receptor complexes. The double loss-of-function mutant of CYP94B3 and ILL6, a JA-Ile amidohydrolase, displayed a JA profile consistent with the collaborative action of the oxidative and the hydrolytic pathways in JA-Ile turnover. Collectively, our results provide an integrated view of how multiple ER-localized CYP94 and JA amidohydrolase enzymes attenuate JA signaling during stress responses.

Volatile methyl jasmonate is a transmissible form of jasmonate and its biosynthesis is involved in systemic jasmonate response in wounding

Volatile organic compounds (VOCs) easily diffuse due to their high hydrophobicity. Because of this physical property, VOCs are able to act as crucial signalling molecules mediating intercellular and interplant communication. Methyl jasmonate (MeJA) is a volatile ester form of jasmonic acid (JA) that is involved in interplant communication in response to biotic and abiotic stresses. Despite its function in interplant communication, the specific role of MeJA in the regulation of intercellular jasmonate responses have been poorly understood. In this study, we demonstrated that MeJA is much more effective than JA in inducing jasmonate response, and the higher efficacy of MeJA relies on its volatile property. To understand the function of MeJA in the regulation of the jasmonate response, we analysed function of JMT gene, Jasmonic acid Methyl Transferase using its knockout mutant (jmt) and overexpressing plants (35S:JMT). Mutant plants that lack JMT expression exhibited reduced jasmonate response, while JMT-overexpressing plants exhibited a higher jasmonate response to JA treatment compared to wild-type plants. In this study, we also showed that JMT is specifically expressed in the phloem, the main vascular system for the transport of phytohormones, and that JMT expression affects systemic jasmonate response in wounding. These results suggest the volatile MeJA is a transmissible form of jasmonate and that its biosynthesis is involved in systemic jasmonate response in wounding.

Different shades of JAZ during plant growth and defense.

Ever since their discovery as key regulators of the jasmonate (JA) signaling pathway, repressor proteins of the JASMONATE ZIM-domain (JAZ) family are rising stars in research on hormonal regulation of plant growth and defense. In plant cells, JAZ repressor proteins interact with an E3 ubiquitin ligase complex (SCFCOI1) that together functions as a JA receptor. In uninduced cells, JAZs block the activity of transcriptional regulators of JA responses by physically binding to them. Upon perception of bioactive JAs, JAZ proteins are rapidly degraded via the ubiquitin/26S proteasome-dependent proteolytic pathway. This releases the JAZ-bound transcription factors, resulting in the activation of downstream JA responses. JAs play a dominant role in regulating defense responses against herbivorous insects and necrotrophic pathogens, and in adaptive responses to beneficial soil-borne microbes. In addition, JAs have a signal function in a myriad of other processes, including abiotic stress reactions and plant growth responses to environmental cues. The JA pathway functions in the context of a complex network of hormone-regulated signaling pathways that, depending on the environmental or developmental condition, can act antagonistically or synergistically on each other to finely balance resource allocation between growth and defense and minimize fitness tradeoffs. In the process of balancing plant growth and defense, gibberellins (GAs) emerged as dominant antagonists of the JA signaling output. GAs regulate different aspects of plant growth via DELLA repressor proteins that block the activity of transcriptional regulators of GA responses by physically binding to them. Analogous to the role of JAZs in the JA pathway, DELLAs are degraded upon perception of GAs, resulting in the activation of downstream growth responses. Interestingly, DELLAs also interact with JAZs, thereby mutually limiting the cellular binding capacity to their cognate transcription factors. Consequently, GA-mediated degradation of DELLAs enhances the cellular binding capacity of JAZs to their cognate transcription factors, thus reducing the potential JA signaling output. This GA-mediated antagonistic effect on the JA pathway becomes apparent during the shade-avoidance response of plants that grow in dense vegetation stands. Shade-intolerant plant species respond to competition for light by increasing apical dominance and accelerating stem and petiole elongation. These growth and developmental responses occur in response to a drop in the red:far-red (R:FR) light ratio that is sensed by the phytochrome photoreceptors, predominantly phyB, and are GA-dependent through GA-mediated degradation of DELLA proteins. This allows them to outgrow neighboring plants, but at the cost of a reduced defensive capacity against necrotrophic pathogens and insect herbivores. In this issue of New Phytologist, Leone and co-workers zoomed in on the specific role of the Arabidopsis thaliana (Arabidopsis) JAZ10 protein in this process and shed light on the involvement of DELLAs in GA-JA crosstalk during the shade avoidance response. Besides growth-related hormones, also effector proteins of pathogens and plant growth-promoting mycorrhizal fungi have recently been shown to target JAZ repressor proteins, thereby changing the defense-related signaling circuitry for their own benefit. Hence, JAZ repressor proteins are emerging as central targets in the rewiring of the hormone-regulated signaling circuitry that regulates growth and defense.

A chemical inhibitor of jasmonate signaling targets JAR1 in Arabidopsis thaliana.

Jasmonates are lipid-derived plant hormones that regulate plant defenses and numerous developmental processes. Although the biosynthesis and molecular function of the most active form of the hormone, (+)-7-iso-jasmonoyl-L-isoleucine (JA-Ile), have been unraveled, it remains poorly understood how the diversity of bioactive jasmonates regulates such a multitude of plant responses. Bioactive analogs have been used as chemical tools to interrogate the diverse and dynamic processes of jasmonate action. By contrast, small molecules impairing jasmonate functions are currently unknown. Here, we report on jarin-1 as what is to our knowledge the first small-molecule inhibitor of jasmonate responses that was identified in a chemical screen using Arabidopsis thaliana. Jarin-1 impairs the activity of JA-Ile synthetase, thereby preventing the synthesis of the active hormone, JA-Ile, whereas closely related enzymes are not affected. Thus, jarin-1 may serve as a useful chemical tool in search for missing regulatory components and further dissection of the complex jasmonate signaling networks.

To grow or defend? Low red:far-red ratios reduce jasmonate sensitivity in Arabidopsis seedlings by promoting DELLA degradation and increasing JAZ10 stability.

How plants balance resource allocation between growth and defense under conditions of competitive stress is a key question in plant biology. Low red:far-red (R:FR) ratios, which signal a high risk of competition in plant canopies, repress jasmonate-induced defense responses. The mechanism of this repression is not well understood. We addressed this problem in Arabidopsis by investigating the role of DELLA and JASMONATE ZIM domain (JAZ) proteins. We showed that a quintuple della mutant and a phyB mutant were insensitive to jasmonate for several physiological readouts. Inactivation of the photoreceptor phyB by low R:FR ratios rapidly reduced DELLA protein abundance, and the inhibitory effect of FR on jasmonate signaling was missing in the gai-1 mutant, which encodes a stable version of the GAI DELLA protein. We also demonstrated that low R:FR ratios and the phyB mutation stabilized the protein JAZ10. Furthermore, we demonstrated that JAZ10 was required for the inhibitory effect of low R:FR on jasmonate responses, and that the jaz10 mutation restored jasmonate sensitivity to the phyB mutant. We conclude that, under conditions of competition for light, plants redirect resource allocation from defense to rapid elongation by promoting DELLA degradation and enhancing JAZ10 stability.

Differential control of pre-invasive and post-invasive antibacterial defense by the Arabidopsis circadian clock.

Plants show a suite of inducible defense responses against bacterial pathogens. Here we investigate in detail the effect of the circadian clock on these reactions in Arabidopsis thaliana. The magnitude of immune responses elicited by flg22, by virulent and by avirulent Pseudomonas syringae strains depends on the time of day of inoculation. The oxidative burst is stronger when flg22 is infiltrated in the morning in wild-type plants but not in the arrhythmic clock mutant lux arrhythmo/phytoclock1 (pcl1), and thus is controlled by the endogenous clock. Similarly, when bacteria are syringe-infiltrated into the leaf, defense gene induction is higher and bacterial growth is suppressed more strongly after morning inoculation in wild-type but not in pcl1 plants. Furthermore, cell death associated with the hypersensitive response was found to be under clock control. Notably, the clock effect depends on the mode of infection: upon spray inoculation onto the leaf surface, defense gene induction is higher and bacterial growth is suppressed more strongly upon evening inoculation. This different phasing of pre-invasive and post-invasive defense relates to clock-regulated stomatal movement. In particular, TIME FOR COFFEE may impact pathogen defense via clock-regulated stomata movement apart from its known role in time-of-day-dependent jasmonate responses. Taken together, these data highlight the importance of the circadian clock for the control of different immune responses at distinct times of the day.

Analyses of Catharanthus roseus and Arabidopsis thaliana WRKY transcription factors reveal involvement in jasmonate signaling.

BackgroundTo combat infection to biotic stress plants elicit the biosynthesis of numerous natural products, many of which are valuable pharmaceutical compounds. Jasmonate is a central regulator of defense response to pathogens and accumulation of specialized metabolites. Catharanthus roseus produces a large number of terpenoid indole alkaloids (TIAs) and is an excellent model for understanding the regulation of this class of valuable compounds. Recent work illustrates a possible role for the Catharanthus WRKY transcription factors (TFs) in regulating TIA biosynthesis. In Arabidopsis and other plants, the WRKY TF family is also shown to play important role in controlling tolerance to biotic and abiotic stresses, as well as secondary metabolism.ResultsHere, we describe the WRKY TF families in response to jasmonate in Arabidopsis and Catharanthus. Publically available Arabidopsis microarrays revealed at least 30% (22 of 72) of WRKY TFs respond to jasmonate treatments. Microarray analysis identified at least six jasmonate responsive Arabidopsis WRKY genes (AtWRKY7, AtWRKY20, AtWRKY26, AtWRKY45, AtWRKY48, and AtWRKY72) that have not been previously reported. The Catharanthus WRKY TF family is comprised of at least 48 members. Phylogenetic clustering reveals 11 group I, 32 group II, and 5 group III WRKY TFs. Furthermore, we found that at least 25% (12 of 48) were jasmonate responsive, and 75% (9 of 12) of the jasmonate responsive CrWRKYs are orthologs of AtWRKYs known to be regulated by jasmonate.ConclusionOverall, the CrWRKY family, ascertained from transcriptome sequences, contains approximately 75% of the number of WRKYs found in other sequenced asterid species (pepper, tomato, potato, and bladderwort). Microarray and transcriptomic data indicate that expression of WRKY TFs in Arabidopsis and Catharanthus are under tight spatio-temporal and developmental control, and potentially have a significant role in jasmonate signaling. Profiling of CrWRKY expression in response to jasmonate treatment revealed potential associations with secondary metabolism. This study provides a foundation for further characterization of WRKY TFs in jasmonate responses and regulation of natural product biosynthesis.Electronic supplementary materialThe online version of this article (doi:10.1186/1471-2164-15-502) contains supplementary material, which is available to authorized users.

Transcriptome Profiling Identified Multiple Jasmonate ZIM-Domain Proteins Involved in the Regulation of Alkaloid Biosynthesis in Tobacco BY-2 Cells

Jasmonate (JA) zinc-finger expressed in inflorescence meristem (ZIM)-domain (JAZ) proteins are key regulators of the JA response in plants. Transcriptome profiling of tobacco BY-2 cells was used to identify 17 members of the NtJAZ family, which were divided into 12 distinct groups based on their predicted amino acid sequences and conserved domains. Transcript levels of eight of the NtJAZ groups increased rapidly upon JA treatment, whereas the remaining members did not show a significant response. The majority of JA-induced NtJAZs formed homo- and heteromers and interacted with NtMYC2a (but not NtERF189) in yeast two-hybrid assays. NtJAZ1, NtJAZ3b, NtJAZ7 and NtJAZ10 were localised in the nucleus and degraded rapidly via the 26S proteasome pathway following the treatment of BY-2 with MeJA. RNAi-induced silencing of NtJAZ1, NtJAZ3, NtJAZ7a and NtJAZ10 greatly reduced the levels of NtPMT transcripts and specifically decreased the nicotine content in the four RNAi transgenic BY-2 lines. The levels of transcripts encoding other nicotine biosynthesis enzymes, NtERF189 and NtMYC2a, and other NtJAZs exhibited different expression patterns in RNAi lines with or without MeJA treatment. Our results indicate that cross-talk occurs among different NtJAZs and forms a complex transcription regulatory scheme for JA-induced nicotine biosynthesis in tobacco.