Defense Priming Research Articles

Defense priming in cabbage (Brassica oleracea) by insect-pathogenic fungi

Defense priming in cabbage (Brassica oleracea) by insect-pathogenic fungi

Linker histone H1 modulates defense priming and immunity in plants.

Linker H1 histones play an important role in animal and human pathogenesis, but their function in plant immunity is poorly understood. Here, we analyzed mutants of the three canonical variants of Arabidopsis H1 histones, namely H1.1, H1.2 and H1.3. We observed that double h1.1h1.2 and triple h1.1h1.2h1.3 (3h1) mutants were resistant to Pseudomonas syringae and Botrytis cinerea infections. Transcriptome analysis of 3h1 mutant plants showed H1s play a key role in regulating the expression of early and late defense genes upon pathogen challenge. Moreover, 3h1 mutant plants showed enhanced production of reactive oxygen species and activation of mitogen activated protein kinases upon pathogen-associated molecular pattern (PAMP) treatment. However, 3h1 mutant plants were insensitive to priming with flg22, a well-known bacterial PAMP which induces enhanced resistance in WT plants. The defective defense response in 3h1 upon priming was correlated with altered DNA methylation and reduced global H3K56ac levels. Our data place H1 as a molecular gatekeeper in governing dynamic changes in the chromatin landscape of defense genes during plant pathogen interaction.

Plant hormone cytokinin at the crossroads of stress priming and control of photosynthesis.

To cope with biotic and abiotic stress conditions, land plants have evolved several levels of protection, including delicate defense mechanisms to respond to changes in the environment. The benefits of inducible defense responses can be further augmented by defense priming, which allows plants to respond to a mild stimulus faster and more robustly than plants in the naïve (non-primed) state. Priming provides a low-cost protection of agriculturally important plants in a relatively safe and effective manner. Many different organic and inorganic compounds have been successfully tested to induce resistance in plants. Among the plethora of commonly used physicochemical techniques, priming by plant growth regulators (phytohormones and their derivatives) appears to be a viable approach with a wide range of applications. While several classes of plant hormones have been exploited in agriculture with promising results, much less attention has been paid to cytokinin, a major plant hormone involved in many biological processes including the regulation of photosynthesis. Cytokinins have been long known to be involved in the regulation of chlorophyll metabolism, among other functions, and are responsible for delaying the onset of senescence. A comprehensive overview of the possible mechanisms of the cytokinin-primed defense or stress-related responses, especially those related to photosynthesis, should provide better insight into some of the less understood aspects of this important group of plant growth regulators.

Increased soluble sugar accumulation in postharvest peaches in response to different defense priming elicitors

Increased soluble sugar accumulation in postharvest peaches in response to different defense priming elicitors

Molecular and Phenotypic Responses of Rhizobacteria-Treated Tomato Plants to Tomato Mosaic Virus Under Greenhouse Conditions.

Tomato mosaic disease, mainly caused by Tomato mosaic virus (ToMV), is one of the devastating viral diseases which adversely affects tomato yield, globally. Plant growth-promoting rhizobacteria (PGPR) have been recently used as bio-elicitors to induce resistance against plant viruses. The goal of this research was to apply PGPR in the tomato rhizosphere and to determine the response of plants challenged with ToMV infection, under greenhouse conditions. Two different strains of PGPR, Pseudomonas fluorescens SM90 and Bacillus subtilis DR06, in single- and double-application methods applied to evaluate their effectiveness in inducing defense-related genes, viz., NPR1, COI1, and PR1-a before (induced systemic resistance [ISR]-prime) and after (ISR-boost) ToMV challenge. Additionally, to investigate the biocontrol potential of PGPR-treated plants against viral infection, plant growth indices, ToMV accumulation, and disease severity were compared in primed and non-primed plants. Analysis of expression patterns of putative defense-related genes before and after ToMV infection indicated that studied PGPR trigger defense priming through different signaling pathways acting at the transcriptional level and in a species-dependent manner. Moreover, the biocontrol efficacy of consortium treatment did not differ significantly from the single bacteria treatments, even though their mode of action differed in transcriptional changes of ISR-induced genes. Instead, simultaneous application of Pseudomonas fluorescens SM90 and Bacillus subtilis DR06 led to more significant growth indices than the single treatments suggesting that integrated application of the PGPR could additively reduce the disease severity and virus titer and promote the growth of the tomato plant. These results suggested that enhanced defense priming via activation of the expression pattern of defense-related genes is responsible for biocontrol activity and growth promotion in PGPR-treated tomato plants challenged with ToMV compared to non-primed plants, under greenhouse conditions.

בין מלחמת "יש ברירה" לרציפות תפיסת הביטחון הישראלית: עיון מחודש בתפיסות שהובילו למלחמת לבנון הראשונה

The Israeli public largely views Israel’s invasion of Lebanon in the summer of 1982 as an unusual event, a deviation from the path that Israel had followed until then. Unlike other wars, Israel did not fight in the Lebanon War for security purposes, but for the political aims and whims of Prime Minister Menachem Begin and Defense Minister Ariel Sharon. Therefore, this war is considered a war of choice, waged by Israel when it was not under an existential threat. The claim that the Lebanon War constitutes an exceptional event in Israeli history is at the heart of this article. In order to determine whether the Lebanon War was indeed a deviation from the “right path” of the State of Israel, we examine it in light of the Israeli security doctrine that had led Israel’s decision-makers until then and that, to some extent, continues to govern them to this day. Using the security doctrine as a guide, we find that this war was not a deviation or an anomaly, but rather an application of Israeli security principles.

Trichoderma root colonization in maize triggers epigenetic changes in genes related to the jasmonic and salicylic acid pathways that prime defenses against Colletotrichum graminicola leaf infection.

Beneficial interactions between plant roots and Trichoderma species lead to both local and systemic enhancements of the plant immune system through a mechanism known as priming of defenses. Previously, we have reported a number of genes and proteins that are differentially regulated in distant tissues of maize plants following inoculation with Trichoderma atroviride. To further investigate the mechanisms involved in the systemic activation of plant responses, here we have further evaluated the regulatory aspects of a selected group of genes when priming is triggered in maize plants. Time-course experiments from the beginning of the interaction between T. atroviride and maize roots followed by leaf infection with Colletotrichum graminicola allowed us to identify a gene set regulated by priming in the leaf tissue. In the same experiment, phytohormone measurements revealed a decrease in jasmonic acid concentration while salicylic acid increased at 2 d and 6 d post-inoculation. In addition, chromatin structure and modification assays showed that chromatin was more open in the primed state compared with unprimed control conditions, and this allowed for quicker gene activation in response to pathogen attack. Overall, the results allowed us to gain insights on the interplay between the phytohormones and epigenetic regulatory events in the systemic and long-lasting regulation of maize plant defenses following Trichoderma inoculation.

Seaweed as a Natural Source against Phytopathogenic Bacteria.

Plant bacterial pathogens can be devastating and compromise entire crops of fruit and vegetables worldwide. The consequences of bacterial plant infections represent not only relevant economical losses, but also the reduction of food availability. Synthetic bactericides have been the most used tool to control bacterial diseases, representing an expensive investment for the producers, since cyclic applications are usually necessary, and are a potential threat to the environment. The development of greener methodologies is of paramount importance, and some options are already available in the market, usually related to genetic manipulation or plant community modulation, as in the case of biocontrol. Seaweeds are one of the richest sources of bioactive compounds, already being used in different industries such as cosmetics, food, medicine, pharmaceutical investigation, and agriculture, among others. They also arise as an eco-friendly alternative to synthetic bactericides. Several studies have already demonstrated their inhibitory activity over relevant bacterial phytopathogens, some of these compounds are known for their eliciting ability to trigger priming defense mechanisms. The present work aims to gather the available information regarding seaweed extracts/compounds with antibacterial activity and eliciting potential to control bacterial phytopathogens, highlighting the extracts from brown algae with protective properties against microbial attack.

Priming by Insects: Differential Effects of Sympatric and Allopatric Priming upon Plant Performance and Tolerance to Herbivory.

Plants have evolved multiple mechanisms to defend themselves from their multiple herbivores. Thus, being able to recognise among them and respond accordingly is fundamental for plant survival and reproduction. Defence priming prepares the plant to better or more rapidly respond to future damage; however, while it is considered an adaptive trait, to date, no studies have evaluated the extent and specificity of the priming recognition. To estimate the costs, benefits and specificity of priming, we used a highly specialist plant-insect system (Datura stramonium-Lema daturaphila) and performed a reciprocal transplant experiment with two populations where a priming stimulus (sympatric vs. allopatric) and a damage treatment (sympatric) were applied. We found no evidence of a fitness cost of priming, given that primed plants without damage showed no reduction in fitness. In contrast, our treatments affected the probability of bud abortion. That is, when damaged plants received no priming or the priming came from an allopatric insect, the likelihood of aborting the first bud was 1.9 times greater compared to plants being primed by their sympatric insect. We also found that damaged plants primed with an allopatric insect produced 14% fewer seeds compared to plants receiving a sympatric priming stimulus. Tolerance to herbivore damage was also the lowest when plants received the priming stimulus from an allopatric insect. Overall, these results suggest that, in our study system, plants recognise their local insect population reducing the negative effect of damage through a tolerance response.

Heritable priming by Trichoderma: A sustainable approach for wheat protection against Bipolaris sorokiniana.

Crop plants encounter a variety of biotic challenges in the field and faces significant reduction in crop yield. In the current scenario of an ever increasing global population, there is an urgent need to protect plant health by using sustainable approach to maximize the crop productivity and to mitigate the food demands. Nowadays, we mostly rely on chemical crop protection techniques, which are causing a number of environmental and health difficulties. Defence priming is a chemical-free, eco-friendly, and sustainable strategy of crop protection, which is also called "green vaccination. In the present study, for the first time, we used Trichoderma as a priming agent to protect wheat crop from spot blotch disease. We have established Trichoderma-mediated defence priming in wheat against Bipolaris sorokiniana for sustainable crop improvement. We have characterised the morphological, disease phenotype, biochemical and yield parameters of Trichoderma-primed and non-primed wheat under disease pressure. Trichoderma-primed plants were found to be more protected against B. sorokiniana as compared to non-primed plants. Biochemical studies indicated that there is no direct defence response after priming stimulus but the defence response was activated only after triggering stimulus in terms of enhanced defence metabolites in primed plants as compared to non-primed plants. In the present study, since defence was activated only when required, that is under disease pressure, there was no unnecessary allocation of resources towards defence. Hence, no yield penalty was shown in primed plants as compared to control. We further evaluated the inheritance of primed state to the next generation and found that progeny of primed parents also performed better than progeny of non-primed parents under disease pressure in terms of protection from B. sorokiniana as well as yield performance. This strategy has the potential to protect crop without any yield penalty and causing environmental degradation. Our research findings indicate that Trichoderma-mediated defence priming could be an alternative approach for improving wheat productivity under biotic stress. To be our best knowledge, this is the first documented report for the Trichoderma-mediated defence priming and induced inheritance in wheat plant. This study will open new arenas in sustainable crop protection strategies for the exploitation of defence priming in crop plants.

Plant Cell Wall Integrity Perturbations and Priming for Defense.

A plant cell wall is a highly complex structure consisting of networks of polysaccharides, proteins, and polyphenols that dynamically change during growth and development in various tissues. The cell wall not only acts as a physical barrier but also dynamically responds to disturbances caused by biotic and abiotic stresses. Plants have well-established surveillance mechanisms to detect any cell wall perturbations. Specific immune signaling pathways are triggered to contrast biotic or abiotic forces, including cascades dedicated to reinforcing the cell wall structure. This review summarizes the recent developments in molecular mechanisms underlying maintenance of cell wall integrity in plant-pathogen and parasitic interactions. Subjects such as the effect of altered expression of endogenous plant cell-wall-related genes or apoplastic expression of microbial cell-wall-modifying enzymes on cell wall integrity are covered. Targeted genetic modifications as a tool to study the potential of cell wall elicitors, priming of signaling pathways, and the outcome of disease resistance phenotypes are also discussed. The prime importance of understanding the intricate details and complete picture of plant immunity emerges, ultimately to engineer new strategies to improve crop productivity and sustainability.

Engineering Resistance against Sclerotinia sclerotiorum Using a Truncated NLR (TNx) and a Defense-Priming Gene.

The association of both cell-surface PRRs (Pattern Recognition Receptors) and intracellular receptor NLRs (Nucleotide-Binding Leucine-Rich Repeat) in engineered plants have the potential to activate strong defenses against a broad range of pathogens. Here, we describe the identification, characterization, and in planta functional analysis of a novel truncated NLR (TNx) gene from the wild species Arachis stenosperma (AsTIR19), with a protein structure lacking the C-terminal LRR (Leucine Rich Repeat) domain involved in pathogen perception. Overexpression of AsTIR19 in tobacco plants led to a significant reduction in infection caused by Sclerotinia sclerotiorum, with a further reduction in pyramid lines containing an expansin-like B gene (AdEXLB8) potentially involved in defense priming. Transcription analysis of tobacco transgenic lines revealed induction of hormone defense pathways (SA; JA-ET) and PRs (Pathogenesis-Related proteins) production. The strong upregulation of the respiratory burst oxidase homolog D (RbohD) gene in the pyramid lines suggests its central role in mediating immune responses in plants co-expressing the two transgenes, with reactive oxygen species (ROS) production enhanced by AdEXLB8 cues leading to stronger defense response. Here, we demonstrate that the association of potential priming elicitors and truncated NLRs can produce a synergistic effect on fungal resistance, constituting a promising strategy for improved, non-specific resistance to plant pathogens.

‘Priming’ protects Piper nigrum L. from Phytophthora capsici through reinforcement of phenylpropanoid pathway and possible enhancement of Piperine biosynthesis

Piper nigrum L. (black pepper), a woody perennial spice crop indigenous to India is positioned at the phylogenetically unique basal lineage of angiosperms. Cultivation of this major spice crop is constrained by rampant fungal and viral infections leading to a lack of disease-free planting material. The major disease that poses severe threat to P. nigrum plantations and nurseries is ‘quick wilt’ caused by the oomycete Phytophthora capsici, which affects the leaf, stem, spike, collar and root. In this paper, we report the consequence of priming in modulating Piper nigrum defense against Phytophthora capsici. Glycol Chitosan (GC) was used to infiltrate detached leaves of mature P. nigrum plants. It was observed that pre-treatment of GC for 24 hours resulted in significant reduction of disease symptoms in infected leaves, as evidenced by the marked decrease in the size of lesions, and also delayed the appearance of symptoms up to 72 hpi. Experiments repeated in P. nigrum seedlings under controlled growth conditions indicate that delayed disease symptoms of GC pre-treated leaves do not spread to healthy uninfiltrated leaves suggesting a priming-associated systemic defense response. An ROS-mediated manifestation of Hypersensitive Response (HR) induced by Chitosan was also evident in pre-treated leaves. A corresponding visual indication of increased lignification was observed, which correlated with an enhanced lignin content of GC-treated leaves. Enhanced callose deposition was also apparent in GC infiltrated leaves, establishing a stimulatory effect of GC in triggering HR through ROS production, enhanced lignification and callose deposition. Key genes of the core phenylpropanoid and isoprenoid pathways along with major defense signalling pathway genes of P. nigrum, including pathogenesis-related genes and hormone signalling genes showed significant transcript enrichment consequential to GC treatment. A significant quantitative enhancement in Piperine content was evident in GC-infiltrated leaves. The systemic nature of priming on disease protection was established through experiments conducted in rooted cuttings monitored for 30 days after disease infection. This is the first report that provides strong molecular evidence endorsing the twofold advantage of defense priming in P. nigrum by improving crop protection with a concomitant enhancement in Piperine biosynthesis.

Airborne cues accelerate flowering and promote photosynthesis in Brassica rapa

Abstract Volatile cues can induce and/or prime plant defences, but it is less well known if plants also respond by altering growth and reproduction‐related parameters. Here, we evaluated whether plant volatile cues can elicit changes in growth, flower production, net photosynthesis rate and defences in receiver plants and whether those responses are similar to those elicited in response to direct herbivore‐feeding. Our results demonstrate that exposure to volatiles emitted from damaged neighbours accelerated the time to first flowering, increased the number of flowers produced and the net photosynthesis rate, but did not alter the fresh harvested plant biomass and volatile defences, compared to non‐exposed plants. Earlier flowering was also observed for plants exposed to volatiles from undamaged plants. These responses differed from those of plants exposed to actual herbivore‐feeding for which feeding enhanced net photosynthesis rate but reduced growth and had no effect on reproduction and defence in response to subsequent herbivory. Synthesis. These findings document that plant airborne cues can influence floral traits of receiver plants. In particular, the flower phenology is modulated, and photosynthesis is enhanced, suggesting that carbon allocation could be directed toward reproduction.

The Green Leaf Volatile (Z)-3-Hexenyl Acetate Is Differently Emitted by Two Varieties of Tulbaghia violacea Plants Routinely and after Wounding.

While studying aromas produced by the edible flowers of Tulbaghia violacea, we noticed a different production of (Z)-3-Hexenyl acetate (a green-leaf volatile, GLV) by purple (var. ‘Violacea’) and white (var. ‘Alba’) flowers. The white Tulbaghia flowers constantly emits (Z)-3-Hexenyl acetate, which is instead produced in a lower amount by the purple-flowered variety. Thus, we moved to analyze the production of (Z)-3-Hexenyl acetate by whole plants of the two varieties by keeping them confined under a glass bell for 5 h together with a SPME (Solid Phase Micro Extraction) fiber. Results show that six main volatile compounds are emitted by T. violacea plants: (Z)-3-Hexenyl acetate, benzyl alcohol, nonanal, decanal, (Z)-3-Hexenyl-α-methylbutyrate, and one unknown compound. By cutting at half-height of the leaves, the (Z)-3-Hexenyl acetate is emitted in high quantities from both varieties, while the production of (Z)-3-Hexenyl-α-methylbutyrate increases. (Z)-3-Hexenyl acetate is a GLV capable of stimulating plant defenses, attracting herbivores and their natural enemies, and it is also involved in plant-to-plant communication and defense priming. Thus, T. violacea could represent a useful model for the study of GLVs production and a ‘signal’ plant capable of stimulating natural defenses in the neighboring plants.

Chitin-induced systemic disease resistance in rice requires both OsCERK1 and OsCEBiP and is mediated via perturbation of cell-wall biogenesis in leaves.

Chitin is a well-known elicitor of disease resistance and its recognition by plants is crucial to perceive fungal infections. Chitin can induce both a local immune response and a systemic disease resistance when provided as a supplement in soils. Unlike local immune responses, it is poorly explored how chitin-induced systemic disease resistance is developed. In this study, we report the systemic induction of disease resistance against the fungal pathogen Bipolaris oryzae by chitin supplementation of soils in rice. The transcriptome analysis uncovered genes related to cell-wall biogenesis, cytokinin signaling, regulation of phosphorylation, and defence priming in the development of chitin-induced systemic response. Alterations of cell-wall composition were observed in leaves of rice plants grown in chitin-supplemented soils, and the disease resistance against B. oryzae was increased in rice leaves treated with a cellulose biosynthesis inhibitor. The disruption of genes for lysin motif (LysM)-containing chitin receptors, OsCERK1 (Chitin elicitor receptor kinase 1) and OsCEBiP (Chitin elicitor-binding protein), compromised chitin-induced systemic disease resistance against B. oryzae and differential expression of chitin-induced genes found in wild-type rice plants. These findings suggest that chitin-induced systemic disease resistance in rice is caused by a perturbation of cell-wall biogenesis in leaves through long-distance signalling after local recognition of chitins by OsCERK1 and OsCEBiP.

Stemborer-induced rice plant volatiles boost direct and indirect resistance in neighboring plants.

Herbivore-induced plant volatiles (HIPVs) are known to be perceived by neighboring plants, resulting in induction or priming of chemical defenses. There is little information on the defense responses that are triggered by these plant-plant interactions, and the phenomenon has rarely been studied in rice. Using chemical and molecular analyses in combination with insect behavioral and performance experiments, we studied how volatiles emitted by rice plants infested by the striped stemborer (SSB) Chilo suppressalis affect defenses against this pest in conspecific plants. Compared with rice plants exposed to the volatiles from uninfested plants, plants exposed to SSB-induced volatiles showed enhanced direct and indirect resistance to SSB. When subjected to caterpillar damage, the HIPV-exposed plants showed increased expression of jasmonic acid (JA) signaling genes, resulting in JA accumulation and higher levels of defensive proteinase inhibitors. Moreover, plants exposed to SSB-induced volatiles emitted larger amounts of inducible volatiles and were more attractive to the parasitoid Cotesia chilonis. By unraveling the factors involved in HIPV-mediated defense priming in rice, we reveal a key defensive role for proteinase inhibitors. These findings pave the way for novel rice management strategies to enhance the plant's resistance to one of its most devastating pests.

A plant growth-promoting bacteria Priestia megaterium JR48 induces plant resistance to the crucifer black rot via a salicylic acid-dependent signaling pathway.

Xanthomonas campestris pv. campestris (Xcc)-induced black rot is one of the most serious diseases in cruciferous plants. Using beneficial microbes to control this disease is promising. In our preliminary work, we isolated a bacterial strain (JR48) from a vegetable field. Here, we confirmed the plant-growth-promoting (PGP) effects of JR48 in planta, and identified JR48 as a Priestia megaterium strain. We found that JR48 was able to induce plant resistance to Xcc and prime plant defense responses including hydrogen peroxide (H2O2) accumulation and callose deposition with elevated expression of defense-related genes. Further, JR48 promoted lignin biosynthesis and raised accumulation of frees salicylic acid (SA) as well as expression of pathogenesis-related (PR) genes. Finally, we confirmed that JR48-induced plant resistance and defense responses requires SA signaling pathway. Together, our results revealed that JR48 promotes plant growth and induces plant resistance to the crucifer black rot probably through reinforcing SA accumulation and response, highlighting its potential as a novel biocontrol agent in the future.

Volatile uptake, transport, perception, and signaling shape a plant's nose.

Herbivore-induced plant volatiles regulate defenses in undamaged neighboring plants. Understanding the mechanisms by which plant volatiles are taken up, perceived, and translated into canonical defense signaling pathways is an important frontier of knowledge. Volatiles can enter plants through stomata and the cuticle. They are likely perceived by membrane-associated receptors as well as intracellular receptors. The latter likely involves metabolization and transport across cell membranes by volatile transporters. Translation of volatiles into defense priming and induction typically involves mitogen-activated protein kinases (MAPKs), WRKY transcription factors, and jasmonates. We propose that the broad range of molecular processes involved in volatile signaling will likely result in substantial spatiotemporal and ontogenetic variation in plant responsiveness to volatiles, with important consequences for plant–environment interactions.

Extracellular self-RNA: A danger elicitor in pepper induces immunity against bacterial and viral pathogens in the field.

Plants and animals serve as hosts for microbes. To protect themselves from microbe-induced damage, plants and animals need to differentiate self-molecules/signals from non-self, microbe-derived molecules. Damage-associated molecular patterns (DAMPs) are danger signals released from the damaged host tissue or present on the surface of stressed cells. Although a self-extracellular DNA has previously been shown to act as a DAMP in different plant species, the existence of a self-extracellular RNA (eRNA) as a danger signal in plants remains unknown. Here, we firstly evaluated the ability of a pepper self-eRNA to activate immunity against viral and bacterial pathogens under field conditions. Pepper leaves pre-infiltrated with self-eRNA exhibited reduced titer of the naturally occurring Tomato spotted wilt virus and diminished symptoms of Xanthomonas axonopodis pv. vesicatoria infection through eliciting defense priming of abscisic acid signaling. At the end of the growing season at 90 days after transplanting, pepper plants treated with self- and non-self-eRNAs showed no difference in fruit yield. Taken together, our discovery demonstrated that self-eRNA can successfully activate plant systemic immunity without any growth penalty, indicating its potential as a novel disease management agent against a broad range of pathogenic microbes.