Plant Defense Signaling Research Articles

Secretions from the ventral eversible gland of Spodoptera exigua caterpillars activate defense-related genes and induce emission of volatile organic compounds in tomato, Solanum lycopersicum.

BackgroundPlant induced defense against herbivory are generally associated with metabolic costs that result in the allocation of photosynthates from growth and reproduction to the synthesis of defense compounds. Therefore, it is essential that plants are capable of sensing and differentiating mechanical injury from herbivore injury. Studies have shown that oral secretions (OS) from caterpillars contain elicitors of induced plant responses. However, studies that shows whether these elicitors originated from salivary glands or from other organs associated with feeding, such as the ventral eversible gland (VEG) are limited. Here, we tested the hypothesis that the secretions from the VEG gland of Spodoptera exigua caterpillars contain elicitors that induce plant defenses by regulating the expression of genes involved in the biosynthesis of volatile organic compounds (VOCs) and other defense-related genes. To test this hypothesis, we quantified and compared the activity of defense-related enzymes, transcript levels of defense-related genes and VOC emission in tomato plants damaged by S. exigua caterpillars with the VEG intact (VEGI) versus plants damaged by caterpillars with the VEG ablated (VEGA).ResultsThe quantified defense-related enzymes (i.e. peroxidase, polyphenol oxidase, and lipoxigenase) were expressed in significantly higher amounts in plants damaged by VEGI caterpillars than in plants damaged by VEGA caterpillars. Similarly, the genes that encode for the key enzymes involved in the biosynthesis of jasmonic acid and terpene synthase genes that regulate production of terpene VOCs, were up-regulated in plants damaged by VEGI caterpillars. Moreover, the OS of VEGA caterpillars were less active in inducing the expression of defense genes in tomato plants. Increased emissions of VOCs were detected in the headspace of plants damaged by VEGI caterpillars compared to plants damaged by VEGA caterpillars.ConclusionThese results suggest that the VEG of S. exigua caterpillars contains elicitors of late plant defense signaling in tomato which trigger defense-related enzymatic activity, regulate expression of defense-related genes, and induce emission of plant VOCs. These signaling cascades may have important ramifications for plant-insect and tritrophic interactions.

A chemical genetic approach demonstrates that MPK3/MPK6 activation and NADPH oxidase-mediated oxidative burst are two independent signaling events in plant immunity.

Plant recognition of pathogen-associated molecular patterns (PAMPs) such as bacterial flagellin-derived flg22 triggers rapid activation of mitogen-activated protein kinases (MAPKs) and generation of reactive oxygen species (ROS). Arabidopsis has at least four PAMP/pathogen-responsive MAPKs: MPK3, MPK6, MPK4 and MPK11. It was speculated that these MAPKs may function downstream of ROS in plant immunity because of their activation by exogenously added H2 O2 . MPK3/MPK6 or their orthologs in other plant species have also been reported to be involved in the ROS burst from the plant respiratory burst oxidase homolog (Rboh) of the human neutrophil gp91phox. However, detailed genetic analysis is lacking. Using a chemical genetic approach, we generated a conditional loss-of-function mpk3 mpk6 double mutant. Consistent with results obtained using a conditionally rescued mpk3 mpk6 double mutant generated previously, the results obtained using the new conditional loss-of-function mpk3 mpk6 double mutant demonstrate that the flg22-triggered ROS burst is independent of MPK3/MPK6. In Arabidopsis mutants lacking a functional AtRbohD, the flg22-induced ROS burst was completely blocked. However, activation of MPK3/MPK6 was not affected. Based on these results, we conclude that the rapid ROS burst and MPK3/MPK6 activation are two independent early signaling events in plant immunity, downstream of FLS2. We also found that MPK4 negatively affects the flg22-induced ROS burst. In addition, salicylic acid pre-treatment enhances the AtRbohD-mediated ROS burst, which is again independent of MPK3/MPK6 based on analysis of the mpk3 mpk6 double mutant. The establishment of an mpk3 mpk6 double mutant system using a chemical genetic approach provides a powerful tool to investigate the function of MPK3/MPK6 in the plant defense signaling pathway.

Expression Profiling of Four Defense-Related Buffalograss Transcripts in Response to Chinch Bug (Hemiptera: Blissidae) Feeding

Oxidative enzymes are one of many key players in plant tolerance responses and defense signaling pathways. This study evaluated gene expression of four buffalograss transcripts (two peroxidases, a catalase, and a GRAS (gibberellic acid insensitive [GAI], repressor of GAI, and scarecrow) and total peroxidase activity in response to western chinch bug (Blissus occiduus Barber) feeding in susceptible and resistant buffalograsses (Buchloë dactyloides (Nuttall) Engelmann). Basal levels of all four transcripts were consistently higher in the resistant buffalograss when compared with the susceptible genotype, which suggests important physiological differences exist between the two buffalograsses. The four defense-related transcripts also showed differential expression between infested and control plants for both the resistant and susceptible buffalograsses. Differences in total peroxidase activity were also detected between control and infested plants, and basal peroxidase activity was higher in the resistant genotype. Overall, this study indicates that elevated basal levels of specific peroxidases, catalases, and GRAS may be an effective buffalograss defense strategy against chinch bug feeding and other similar biotic stresses.

Glycerol-3-Phosphate Metabolism in Wheat Contributes to Systemic Acquired Resistance against Puccinia striiformis f. sp. tritici

Glycerol-3-phosphate (G3P) is a proposed regulator of plant defense signaling in basal resistance and systemic acquired resistance (SAR). The GLY1-encoded glycerol-3-phosphate dehydrogenase (G3PDH) and GLI1-encoded glycerol kinase (GK) are two key enzymes involved in the G3P biosynthesis in plants. However, their physiological importance in wheat defense against pathogens remains unclear. In this study, quantification analysis revealed that G3P levels were significantly induced in wheat leaves challenged by the avirulent Puccinia striiformis f. sp. tritici (Pst) race CYR23. The transcriptional levels of TaGLY1 and TaGLI1 were likewise significantly induced by avirulent Pst infection. Furthermore, knocking down TaGLY1 and TaGLI1 individually or simultaneously with barley stripe mosaic virus-induced gene silencing (BSMV-VIGS) inhibited G3P accumulation and compromised the resistance in the wheat cultivar Suwon 11, whereas the accumulation of salicylic acid (SA) and the expression of the SA-induced marker gene TaPR1 in plant leaves were altered significantly after gene silencing. These results suggested that G3P contributes to wheat systemic acquired resistance (SAR) against stripe rust, and provided evidence that the G3P function as a signaling molecule is conserved in dicots and monocots. Meanwhile, the simultaneous co-silencing of multiple genes by the VIGS system proved to be a powerful tool for multi-gene functional analysis in plants.

Aphid honeydew alters plant defence responses

Summary Plants respond to insect herbivory by ramping up the production of defence‐related jasmonic acid (JA). The pea aphid, Acyrthosiphon pisum, is able to avoid triggering JA as it feeds. The aim of our study was to examine whether or not aphids simply avoid triggering plant defence signalling or whether they are able to manipulate plant signalling in a way to alter effectual plant defences. To resolve this, we tested the ability of pea aphids to suppress the accumulation of damage‐induced JA. We then tested exogenous application of aphid honeydew as a possible mechanism for JA suppression. While broad bean plants, Vicia faba, accumulated JA in response to artificial damage, the presence of concurrent aphid feeding suppressed JA accumulation. Furthermore, exogenous application of aphid honeydew induced salicylic acid (SA) and suppressed JA accumulation. We also report the presence of SA within honeydew itself. We conclude that both aphid feeding and exogenous honeydew deposition suppress JA accumulation in response to damage. We demonstrate that honeydew application results in an increased accumulation of SA within plant tissue and find that levels of SA present within honeydew do not fully account for this increase. This finding demonstrates, for the first time, that aphid honeydew suppresses induced plant defence.

Modification of Plasma Membrane Organization in Tobacco Cells Elicited by Cryptogein

Lipid mixtures within artificial membranes undergo a separation into liquid-disordered and liquid-ordered phases. However, the existence of this segregation into microscopic liquid-ordered phases has been difficult to prove in living cells, and the precise organization of the plasma membrane into such phases has not been elucidated in plant cells. We developed a multispectral confocal microscopy approach to generate ratiometric images of the plasma membrane surface of Bright Yellow 2 tobacco (Nicotiana tabacum) suspension cells labeled with an environment sensitive fluorescent probe. This allowed the in vivo characterization of the global level of order of this membrane, by which we could demonstrate that an increase in its proportion of ordered phases transiently occurred in the early steps of the signaling triggered by cryptogein and flagellin, two elicitors of plant defense reactions. The use of fluorescence recovery after photobleaching revealed an increase in plasma membrane fluidity induced by cryptogein, but not by flagellin. Moreover, we characterized the spatial distribution of liquid-ordered phases on the membrane of living plant cells and monitored their variations induced by cryptogein elicitation. We analyze these results in the context of plant defense signaling, discuss their meaning within the framework of the "membrane raft" hypothesis, and propose a new mechanism of signaling platform formation in response to elicitor treatment.

The pepper patatin-like phospholipase CaPLP1 functions in plant cell death and defense signaling

Phospholipases hydrolyze phospholipids into fatty acids and other lipophilic substances. Phospholipid signaling is crucial for diverse cellular processes in plants. However, the precise role of phospholipases in plant cell death and defense signaling is not fully understood. Here, we identified a pepper (Capsicum annuum) patatin-like phospholipase (CaPLP1) gene that is transcriptionally induced in pepper leaves by avirulent Xanthomonas campestris pv. vesicatoria (Xcv) infection. CaPLP1 containing an N-terminal signal peptide localized to the cytoplasm and plasma membrane, leading to the secretion into the apoplastic regions. Silencing of CaPLP1 in pepper conferred enhanced susceptibility to Xcv infection. Defense responses to Xcv, including the generation of reactive oxygen species (ROS), hypersensitive cell death and the expression of the salicylic acid (SA)-dependent marker gene CaPR1, were compromised in the CaPLP1-silenced pepper plants. Transient expression of CaPLP1 in pepper leaves induced the accumulation of fluorescent phenolics, expression of the defense marker genes CaPR1 and CaSAR82A, and generation of ROS, ultimately leading to the hypersensitive cell death response. Overexpression (OX) of CaPLP1 in Arabidopsis also conferred enhanced resistance to Pseudomonas syringae pv. tomato (Pst) and Hyaloperonospora arabidopsidis infection. CaPLP1-OX leaves showed reduced Pst growth, enhanced ROS burst and electrolyte leakage, induction of the defense response genes AtPR1, AtRbohD and AtGST, as well as constitutive activation of both the SA-dependent gene AtPR1 and the JA-dependent gene AtPDF1.2. Together, these results suggest that CaPLP1 is involved in plant defense and cell death signaling in response to microbial pathogens.

Involvement of the glutamate receptorAtGLR3.3 in plant defense signaling and resistance toHyaloperonospora arabidopsidis

Like their animal counterparts, plant glutamate receptor-like (GLR) homologs are intimately associated with Ca(2+) influx through plasma membrane and participate in various physiological processes. In pathogen-associated molecular patterns (PAMP)-/elicitor-mediated resistance, Ca(2+) fluxes are necessary for activating downstream signaling events related to plant defense. In this study, oligogalacturonides (OGs), which are endogenous elicitors derived from cell wall degradation, were used to investigate the role of Arabidopsis GLRs in defense signaling. Pharmacological investigations indicated that GLRs are partly involved in free cytosolic [Ca(2+)] ([Ca(2+)]cyt) variations, nitric oxide (NO) production, reactive oxygen species (ROS) production and expression of defense-related genes by OGs. In addition, wild-type Col-0 plants treated with the glutamate-receptor antagonist 6,7-dinitriquinoxaline-2,3-dione (DNQX) had a compromised resistance to Botrytis cinerea and Hyaloperonospora arabidopsidis. Moreover, we provide genetic evidence that AtGLR3.3 is a key component of resistance against H. arabidopsidis. In addition, some OGs-triggered immune events such as defense gene expression, NO and ROS production are also to different extents dependent on AtGLR3.3. Taken together, these data provide evidence for the involvement of GLRs in elicitor/pathogen-mediated plant defense signaling pathways in Arabidopsis thaliana.

Herbivore exploits orally secreted bacteria to suppress plant defenses

Induced plant defenses in response to herbivore attack are modulated by cross-talk between jasmonic acid (JA)- and salicylic acid (SA)-signaling pathways. Oral secretions from some insect herbivores contain effectors that overcome these antiherbivore defenses. Herbivores possess diverse microbes in their digestive systems and these microbial symbionts can modify plant-insect interactions; however, the specific role of herbivore-associated microbes in manipulating plant defenses remains unclear. Here, we demonstrate that Colorado potato beetle (Leptinotarsa decemlineata) larvae exploit bacteria in their oral secretions to suppress antiherbivore defenses in tomato (Solanum lycopersicum). We found that antibiotic-untreated larvae decreased production of JA and JA-responsive antiherbivore defenses, but increased SA accumulation and SA-responsive gene expression. Beetles benefit from down-regulating plant defenses by exhibiting enhanced larval growth. In SA-deficient plants, suppression was not observed, indicating that suppression of JA-regulated defenses depends on the SA-signaling pathway. Applying bacteria isolated from larval oral secretions to wounded plants confirmed that three microbial symbionts belonging to the genera Stenotrophomonas, Pseudomonas, and Enterobacter are responsible for defense suppression. Additionally, reinoculation of these bacteria to antibiotic-treated larvae restored their ability to suppress defenses. Flagellin isolated from Pseudomonas sp. was associated with defense suppression. Our findings show that the herbivore exploits symbiotic bacteria as a decoy to deceive plants into incorrectly perceiving the threat as microbial. By interfering with the normal perception of herbivory, beetles can evade antiherbivore defenses of its host.

An S-type anion channel SLAC1 is involved in cryptogein-induced ion fluxes and modulates hypersensitive responses in tobacco BY-2 cells.

Pharmacological evidence suggests that anion channel-mediated plasma membrane anion effluxes are crucial in early defense signaling to induce immune responses and hypersensitive cell death in plants. However, their molecular bases and regulation remain largely unknown. We overexpressed Arabidopsis SLAC1, an S-type anion channel involved in stomatal closure, in cultured tobacco BY-2 cells and analyzed the effect on cryptogein-induced defense responses including fluxes of Cl− and other ions, production of reactive oxygen species (ROS), gene expression and hypersensitive responses. The SLAC1-GFP fusion protein was localized at the plasma membrane in BY-2 cells. Overexpression of SLAC1 enhanced cryptogein-induced Cl− efflux and extracellular alkalinization as well as rapid/transient and slow/prolonged phases of NADPH oxidase-mediated ROS production, which was suppressed by an anion channel inhibitor, DIDS. The overexpressor also showed enhanced sensitivity to cryptogein to induce downstream immune responses, including the induction of defense marker genes and the hypersensitive cell death. These results suggest that SLAC1 expressed in BY-2 cells mediates cryptogein-induced plasma membrane Cl− efflux to positively modulate the elicitor-triggered activation of other ion fluxes, ROS as well as a wide range of defense signaling pathways. These findings shed light on the possible involvement of the SLAC/SLAH family anion channels in cryptogein signaling to trigger the plasma membrane ion channel cascade in the plant defense signal transduction network.

Hormone defense networking in rice: tales from a different world

Recent advances in plant immunity research underpin the pivotal role of small-molecule hormones in regulating the plant defense signaling network. Although most of our understanding comes from studies of dicot plants such as Arabidopsis thaliana, new studies in monocots are providing additional insights into the defense-regulatory role of phytohormones. Here, we review the roles of both classical and more recently identified stress hormones in regulating immunity in the model monocot rice (Oryza sativa) and highlight the importance of hormone crosstalk in shaping the outcome of rice-pathogen interactions. We also propose a defense model for rice that does not support a dichotomy between the pathogen lifestyle and the effectiveness of the archetypal defense hormones salicylic acid (SA) and jasmonic acid (JA).

Methionine elicits H2O2 generation and defense gene expression in grapevine and reduces Plasmopara viticola infection

Methionine (Met) is a nutritionally essential sulfur-containing amino acid (SAA) known for its preponderant role as initiator in protein synthesis. However, other functions for Met in plants are not well described. The implication of this SAA in oxidative stress tolerance has been recently reported, however the mode of action of Met is still poorly understood. Here, we analyzed the elicitor activity of Met in grapevine as well as its effect on Plasmopara viticola resistance. The results show that Met induces hydrogen peroxide (H2O2) generation, a key element in plant defense signaling, and upregulates the expression of a battery of defense-related genes. Transcript levels of these genes were not further modulated by P. viticola inoculation of Met-pretreated plants, suggesting an elicitor role rather than a priming role for Met in grapevine. Met treatment also reduces P. viticola development in grapevine plants grown under glasshouse controlled-conditions. Fungitoxicity assays revealed that Met possesses a moderate antifungal activity compared with cysteine (Cys), another SAA known for its toxic effect to a large spectrum of fungi.

Molecular and biochemical characterization of the jasmonic acid methyltransferase gene from black cottonwood (Populus trichocarpa)

Methyl jasmonate is a metabolite known to be produced by many plants and has roles in diverse biological processes. It is biosynthesized by the action of S-adenosyl-l-methionine:jasmonic acid carboxyl methyltransferase (JMT), which belongs to the SABATH family of methyltransferases. Herein is reported the isolation and biochemical characterization of a JMT gene from black cottonwood (Populus trichocarpa). The genome of P. trichocarpa contains 28 SABATH genes (PtSABATH1 to PtSABATH28). Recombinant PtSABATH3 expressed in Escherichia coli showed the highest level of activity with jasmonic acid (JA) among carboxylic acids tested. It was therefore renamed PtJMT1. PtJMT1 also displayed activity with benzoic acid (BA), with which the activity was about 22% of that with JA. PtSABATH2 and PtSABATH4 were most similar to PtJMT1 among all PtSABATHs. However, neither of them had activity with JA. The apparent Km values of PtJMT1 using JA and BA as substrate were 175μM and 341μM, respectively. Mutation of Ser-153 and Asn-361, two residues in the active site of PtJMT1, to Tyr and Ser respectively, led to higher specific activity with BA than with JA. Homology-based structural modeling indicated that substrate alignment, in which Asn-361 is involved, plays a role in determining the substrate specificity of PtJMT1. In the leaves of young seedlings of black cottonwood, the expression of PtJMT1 was induced by plant defense signal molecules methyl jasmonate and salicylic acid and a fungal elicitor alamethicin, suggesting that PtJMT1 may have a role in plant defense against biotic stresses. Phylogenetic analysis suggests that PtJMT1 shares a common ancestor with the Arabidopsis JMT, and functional divergence of these two apparent JMT orthologs has occurred since the split of poplar and Arabidopsis lineages.

New Synthesis: Investigating Mutualisms in Virus-Vector Interactions

Most plant viruses depend on insect vectors for transmission from one host to another. Aphids are the most notable among these insect vectors, transmitting more than half of all described virus species. Virus transmission ranges from simple attachment to the tip of the aphid stylets tomore complicated interactions that involve ingestion of the viruses, passage through the haemolymph to the salivary glands, and subsequent injection into a new host plant. Plant viruses, which clearly benefit from being transported to new hosts, generally do not harm their aphid vectors. To the contrary, the frequent observation of improved aphid growth on virus-infected plants suggests that these interactions can be mutualistic, benefitting both the viruses and their insect vectors. Several studies have demonstrated that plant viruses manipulate plants, thus increasing the number of inoculated insect vectors. This can be accomplished by attracting aphids to infected plants and/or changing plant physiology in a manner that improves aphid reproduction. Increased fecundity and developmental rate of aphids on virus-infected plantswill result in greater dispersal of viruliferous aphids tonewhosts as host plant qualitydeclines andaphids becomecrowded.Crowding also induces the formation of alate aphids, which provide longer-distance virus transmission. Virus-vector interactions can be mediated indirectly by virusinduced increases in plant nutritive value, suppression of plant defense responses, or some combination of these effects. Recent publications provide evidence for active manipulation of herbivore defenses by plant-pathogenic viruses. For instance, the 2b silencing suppressor protein of cucumber mosaic virus compromises jasmonate-regulated insect defenses and improves performance of Myzus persicae (green peach aphid; Lewsey et al. 2010). Similarly, pathogenicity factor βC1, which is encoded in the betasatellite of tomato yellow leaf curl China virus, suppresses jasmonate-mediated defenses and improves growth of Bemisia tabaci (silverleaf whitefly) on Nicotiana tabacum (Zhang et al. 2012). Because arthropod-mediated transmission benefits plant viruses, and there is an apparent reproductive benefit for the insects, in many cases, the virus-vector interaction can be considered a mutualism. For instance, individual aphids that transport viruses to a new host plant would benefit from rapid virus-mediated suppression of plantdefenses. In suchasituation, it clearlywouldbeadvantageous foraphids topick up and transmit viruses. It is also possible that, by transmitting plant-pathogenic viruses, hemipteran insects can expand their ranges to include normally unsuitable host plants.Our ownunpublishedobservations show that, ifNicotiana benthamiana is infected with turnip mosaic virus, the survival of newly arrivingMyzus persicae (green peach aphids) increases from 30% to 90%. Similarly,N. tabacum is a poor host for some B. tabaci biotypes, but whitefly performance is improved greatly through prior begomovirus infection (Zhang et al. 2012). If an insect vector arriving at an uninfected plant benefits from being viruliferous, this suggests that such insects will benefit by acquiring viruses prior to moving to new host plants. For instance, aphids could be feeding from virus-infected tissue in a targetedmanner, or specifically selecting virus-infected cells as they are probing to reach the phloem. When an aphid punctures a cell infected with cauliflower mosaic virus, the virus rearranges itself rapidly, thereby promoting attachment to the aphid stylets (Martiniere et al. 2013). It has been suggested that this is due to viral recognition of some component of the plant defense signaling pathway that is triggered by aphid feeing. However, it also is possible that the aphids acquire viruses by actively manipulating plant cellular metabolism. Conversely, when a viruliferous aphid settles on a new plant, its feeding success may be promoted if viruses are deposited in a cellular location that allows rapid replication and thereby suppression of plant defenses. To date, most research on the molecular mechanisms of virus-vector interactions has focused on studying host plant manipulation by viruses that promotes aphid performance. However, aphids and other insect vectors also may promote virus transmission by altering plant physiology. To initiate progress in this research area, it will be necessary to identify conditions under which there are immediate benefits to insect vectors carrying viruses to newhost plants. Furthermore, future investigation of these ecologically important but relatively under-investigated three-way interactions should include a greater focus on discovering behavioral and molecular mechanisms by which insect vectors can actively manipulate plant-virus interactions for their own benefit.

Transcriptome Analysis of Induced Systemic Drought Tolerance Elicited by Pseudomonas chlororaphis O6 in Arabidopsis thaliana

Root colonization by Pseudomonas chlororaphis O6 induces systemic drought tolerance in Arabidopsis thaliana. Microarray analysis was performed using the 22,800-gene Affymetrix GeneChips to identify differentially-expressed genes from plants colonized with or without P. chlororaphis O6 under drought stressed conditions or normal growth conditions. Root colonization in plants grown under regular irrigation condition increased transcript accumulation from genes associated with defense, response to reactive oxygen species, and auxin- and jasmonic acid-responsive genes, but decreased transcription factors associated with ethylene and abscisic acid signaling. The cluster of genes involved in plant disease resistance were up-regulated, but the set of drought signaling response genes were down-regulated in the P. chlororaphis O6-colonized under drought stress plants compared to those of the drought stressed plants without bacterial treatment. Transcripts of the jasmonic acid-marker genes, VSP1 and pdf-1.2, the salicylic acid regulated gene, PR-1, and the ethylene-response gene, HEL, also were up-regulated in plants colonized by P. chlororaphis O6, but differed in their responsiveness to drought stress. These data show how gene expression in plants lacking adequate water can be remarkably influenced by microbial colonization leading to plant protection, and the activation of the plant defense signal pathway induced by root colonization of P. chlororaphis O6 might be a key element for induced systemic tolerance by microbes.

Abnormal Glycosphingolipid Mannosylation Triggers Salicylic Acid–Mediated Responses inArabidopsis

The Arabidopsis thaliana protein GOLGI-LOCALIZED NUCLEOTIDE SUGAR TRANSPORTER (GONST1) has been previously identified as a GDP-d-mannose transporter. It has been hypothesized that GONST1 provides precursors for the synthesis of cell wall polysaccharides, such as glucomannan. Here, we show that in vitro GONST1 can transport all four plant GDP-sugars. However, gonst1 mutants have no reduction in glucomannan quantity and show no detectable alterations in other cell wall polysaccharides. By contrast, we show that a class of glycosylated sphingolipids (glycosylinositol phosphoceramides [GIPCs]) contains Man and that this mannosylation is affected in gonst1. GONST1 therefore is a Golgi GDP-sugar transporter that specifically supplies GDP-Man to the Golgi lumen for GIPC synthesis. gonst1 plants have a dwarfed phenotype and a constitutive hypersensitive response with elevated salicylic acid levels. This suggests an unexpected role for GIPC sugar decorations in sphingolipid function and plant defense signaling. Additionally, we discuss these data in the context of substrate channeling within the Golgi.

Crosstalk between intracellular and extracellular salicylic acid signaling events leading to long-distance spread of signals

It is well recognized that salicylic acid (SA) acts as a natural signaling molecule involved in both local and systemic plant defense responses upon attacks by pathogens. Recently, cellular SA receptors and a number of SA-related phloem-mobile signals were identified. Here, we compare the old and up-to-date concepts of plant defense signaling events involving SA. Finally, the crosstalk between intracellular and extracellular SA signaling events leading to long-distance spread of signals was outlined by focusing on the modes of both the short- and long-distance signaling events involving the actions of SA. For the above purpose, two distinct conceptual models for local SA perception and signaling mechanisms in the intracellular and extracellular paths (referred to as models i and ii, respectively) were proposed. In addition to two local SA perception models, we propose that the long-distance SA action could be attributed to three different modes, namely, (iii) local increase in SA followed by transport of SA and SA intermediates, (iv) systemic propagation of SA-derived signals with both chemical and electrical natures without direct movement of SA, and (v) integrated crosstalk allowing alternately repeated secondary signal propagation and biosynthesis of SA and/or conversion of inert SA intermediates to free SA finally contributing to the systemic spread of SA-derived signals. We review here that the long-distance SA signaling events (models iii-v), inevitably involve the mechanisms described in the local signaling models (models i and ii) as the key pieces of the crosstalk.

Salicylic acid: an old hormone up to new tricks

Salicylic acid (SA) acts as a signalling molecule in plant defence against biotrophic and hemibiotrophic phytopathogens. The biosynthesis of SA on pathogen detection is essential for local and systemic acquired resistance, as well as the accumulation of pathogenesis-related (PR) proteins. SA biosynthesis can occur via several different substrates, but is predominantly accomplished by isochorismate synthase (ICS1) following pathogen recognition. The roles of BTB domain-containing proteins, NPR1, NPR3 and NPR4, in SA binding and signal transduction have been re-examined recently and are elaborated upon in this review. The pathogen-mediated manipulation of SA-dependent defences, as well as the crosstalk between the SA signalling pathway, other plant hormones and defence signals, is also discussed in consideration of recent research. Furthermore, the recent links established between SA, pathogen-triggered endoplasmic reticulum stress and the unfolded protein response are highlighted.

The contribution of Trichoderma to balancing the costs of plant growth and defense.

Trichoderma is a fungal genus of cosmopolitan distribution and high biotechnological value, with several species currently used as biological control agents. Additionally, the enzyme systems of the fungus are widely applied in industry. Species of Trichoderma protect plants against the attack of soil-borne plant pathogens by competing for nutrients and inhibiting or killing plant pathogenic fungi and oomycetes, through the production of antibiotics and/or hydrolytic enzymes. In addition to the role of Trichoderma spp. as biocontrol agents, they have other beneficial effects on plants, including the stimulation of plant defenses and the promotion of plant growth. In this review, we focus on the complex plant defense signaling network that allows the recognition of fungi as non-hostile microbes, including microbial-associated molecular patterns (MAMPs), damage-associated molecular patterns (DAMPs) and secreted elicitors. We also examine how fungal interactions with plant receptors can activate induced resistance by priming and balancing plant defense and growth responses. Our observations are integrated into a model describing Trichoderma-plant hormone signaling network interactions.

Effects of the virus satellite gene βC1 on host plant defense signaling and volatile emission

Tomato Yellow Leaf Curl China virus spreads together with its invasive vector, the silverleaf whitefly B biotype, which exhibits higher growth rates on infected plants. Previous studies indicate that the virus satellite gene βC1 accounts for the visible symptoms of infection and inhibits the constitutive expression of jasmonic acid (JA)—a phytohormone involved in plant defense against whiteflies—and of some JA-regulated genes. Here we present new details of the effects of on plant signaling and defense, obtained with (non-host) transgenic Arabidopsis thaliana and Nicotiana benthamiana plants. We found that JA induction in response to wounding was reduced in plants expressing βC1. This result implies that βC1 acts on conserved plant regulation mechanisms and might impair the entire JA defense pathway. Furthermore, transformed N. benthamiana plants exhibited elevated emissions of the volatile compound linalool, suggesting that βC1 also influences plant-derived olfactory cues available to vector and non-vector insects.