Defence-related Phytohormones Research Articles

The Effects of Lead and Cross-Talk Between Lead and Pea Aphids on Defence Responses of Pea Seedlings.

The main goal of this study was to investigate the effect of lead (Pb) at various concentrations, as an abiotic factor, and the cross-talk between Pb and pea aphid (Acyrthosiphon pisum (Harris)) (Hemiptera: Aphididae), as a biotic factor, on the defence responses of pea seedlings (Pisum sativum L. cv. Cysterski). The analysis of growth parameters for pea seedlings demonstrated that Pb at a low concentration, i.e., 0.025-0.0625 mM Pb(NO3)2, caused a hormesis effect, i.e., stimulation of seedling growth, whereas Pb at higher concentrations, i.e., 0.01-0.325 mM Pb(NO3)2, inhibited growth, which manifested as the inhibition of length and fresh biomass. The differences in the level of the main defence-related phytohormones, such as abscisic acid (ABA), jasmonic acid (JA) and salicylic acid (SA), and indole-3-acetic acid (IAA)-an auxin stimulating plant cell growth-depended on the dose of Pb, aphid infestation and direct contact of the stress factor with the organ. A high accumulation of soluble sugars in the organs of pea seedlings both at sublethal doses and hormetic doses at early experimental time points was observed. At 0 h and 24 h of the experiment, the hormetic doses of Pb significantly stimulated invertase activities, especially in the roots. Moreover, an increase was observed in the pisatin concentration in pea seedlings growing in the presence of different concentrations of Pb and in the case of cross-talk between Pb and A. pisum in relation to the control. Additionally, a significant induction of the expressions of isoflavone synthase (IFS) and 6α-hydroxymaackiain 3-O-methyltransferase (HMM) genes, which participate in the regulation of the pisatin biosynthesis pathway, in pea seedlings growing under the influence of sublethal 0.5 mM Pb(NO3)2 and hormetic 0.075 mM Pb(NO3)2 doses of Pb was noted. The obtained results showed that the response of P. sativum seedlings depends on the Pb dose applied, direct contact of the stress factor with the organ and the duration of contact.

Plant Protection by Tannins Depends on Defence-Related Phytohormones

AbstractPlants produce a wide variety of secondary metabolites to sustain and protect themselves against a wide range of stresses. Among these metabolites, tannins are one of the most abundant polyphenolic compounds, accounting for 25% of the dry weight of leaves, roots and bark of woody plants, but are also abundant in flowering and seed-producing plants. The presence of tannins in these organs serves to protect plants against herbivorous and pathogenic attack through their antidigestive and antimicrobial properties. In addition, tannins play a role in regulating plant growth and development by inhibiting the consumption of unripe fruits due to their astringency. In addition, several studies have also revealed various roles of them under environmental stresses. Tannins can be classified into condensed tannins (CTs), hydrolysable tannins (HTs) and phlorotannins. They are synthesised in plants via the acetate–malonate and shikimic acid pathways. Their accumulation is regulated by several transcription factors during normal development and under different stress conditions. Despite their multiple roles in plant life, information on the regulation of tannin metabolism by defence-related phytohormones is very limited. To cope with biotic and abiotic stresses, plant responses are regulated by defence-related phytohormones such as salicylic acid (SA), jasmonic acid (JA), ethylene (ET) and abscisic acid (ABA), which act as regulators of tannin production under adverse conditions. This review focuses on tannin production, moreover its occurrence, defence potential and regulation by phytohormones under different environmental and biotic stresses, based on the most recent and relevant data. Graphical Abstract

Global Change Factors Influence Plant-Epichloë Associations.

There is an increasing interest in determining the influence of global change on plant-microorganism interactions. We review the results of experiments that evaluated the effects of the global change factors carbon dioxide, ozone, temperature, drought, flooding, and salinity on plant symbioses with beneficial Epichloë endophytes. The factors affected the performance of both plants and endophytes as well as the frequency of plants symbiotic with the fungus. Elevated carbon dioxide levels and low temperatures differentially influenced the growth of plants and endophytes, which could compromise the symbioses. Furthermore, we summarise the plant stage in which the effects of the factors were quantified (vegetative, reproductive, or progeny). The factors ozone and drought were studied at all plant stages, but flooding and carbon dioxide were studied in just a few of them. While only studied in response to ozone and drought, evidence showed that the effects of these factors on symbiotic plants persisted trans-generationally. We also identified the putative mechanisms that would explain the effects of the factors on plant-endophyte associations. These mechanisms included the increased contents of reactive oxygen species and defence-related phytohormones, reduced photosynthesis, and altered levels of plant primary metabolites. Finally, we describe the counteracting mechanisms by which endophytes would mitigate the detrimental effects of the factors on plants. In presence of the factors, endophytes increased the contents of antioxidants, reduced the levels of defence-related phytohormones, and enhanced the plant uptake of nutrients and photosynthesis levels. Knowledge gaps regarding the effects of global change on plant-endophyte associations were identified and discussed.

Flowers prepare thyselves: leaf and root herbivores induce specific changes in floral phytochemistry with consequences for plant interactions with florivores.

Summary The phenotypic plasticity of flowering plants in response to herbivore damage to vegetative tissues can affect plant interactions with flower‐feeding organisms. Such induced systemic responses are probably regulated by defence‐related phytohormones that signal flowers to alter secondary chemistry that affects resistance to florivores. Current knowledge on the effects of damage to vegetative tissues on plant interactions with florivores and the underlying mechanisms is limited.We compared the preference and performance of two florivores on flowering Brassica nigra plants damaged by one of three herbivores feeding from roots or leaves. To investigate the underlying mechanisms, we quantified expression patterns of marker genes for defence‐related phytohormonal pathways, and concentrations of phytohormones and glucosinolates in buds and flowers.Florivores displayed contrasting preferences for plants damaged by herbivores feeding on roots and leaves. Chewing florivores performed better on plants damaged by folivores, but worse on plants damaged by the root herbivore. Chewing root and foliar herbivory led to specific induced changes in the phytohormone profile of buds and flowers. This resulted in increased glucosinolate concentrations for leaf‐damaged plants, and decreased glucosinolate concentrations for root‐damaged plants.The outcome of herbivore–herbivore interactions spanning from vegetative tissues to floral tissues is unique for the inducing root/leaf herbivore and receiving florivore combination.

Transcriptional Reprogramming of Arabidopsis thaliana Defence Pathways by the Entomopathogen Beauveria bassiana Correlates With Resistance Against a Fungal Pathogen but Not Against Insects.

The entomopathogenic fungus Beauveria bassiana can adopt an endophytic lifestyle by colonising a wide array of plant species. Beauveria-colonised plants can show enhanced resistance against insects and plant pathogens alike. However, little is known about the molecular and physiological mechanisms that govern such interactions. Here, we assessed the effects of two B. bassiana strains (BG11, FRh2) on the growth of Arabidopsis thaliana and its resistance against two herbivore species and a phytopathogen. Plant responses were studied on the transcriptomic and metabolic level using microarrays and by measuring changes in defence-related phytohormones and glucosinolates (GLSs). Root inoculation with B. bassiana BG11 significantly increased plant growth, while FRh2 had no such effect. Both Beauveria strains decreased leaf lesion area caused by the phytopathogen Sclerotinia sclerotiorum but did not affect population growth of the aphid Myzus persicae or the growth of Plutella xylostella caterpillars. Microarray analyses of leaves from endophyte-inoculated A. thaliana provided evidence for transcriptional reprogramming of plant defence pathways, with strain-specific changes in the expression of genes related to pathogenesis, phytoalexin, jasmonic (JA), and salicylic acid (SA) signalling pathways. However, B. bassiana colonisation did not result in higher concentrations of JA and SA or major changes in leaf GLS profiles. We conclude that the endophyte B. bassiana induces plant defence responses and hypothesise that these contribute to enhanced resistance against S. sclerotiorum.

Highly localized and persistent induction of Bx1-dependent herbivore resistance factors in maize.

The induced production of secondary metabolites in herbivore-attacked plants varies in space and time. However, the consequences of these spatiotemporal patterns for herbivore performance are not well understood. This is particularly true for 1,4-benzoxazin-3-ones (BXs), the major induced defensive metabolites of maize. Here we report on the spatiotemporal dynamics of BX induction and its consequences for the leaf feeder Spodoptera littoralis. Defence-related phytohormones and transcript levels of BX biosynthetic genes were upregulated locally at the wound site within 12h of herbivory. Within another 12h, the insecticidal BX HDMBOA-Glc started to accumulate in a highly localized manner at the feeding site. Changes in BX metabolism away from the feeding site within the same leaf were much weaker and were undetected in systemic leaves. Following the removal of the caterpillars, local HDMBOA-Glc levels remained elevated for 7days. Caterpillars that were forced to feed directly on locally induced leaf parts, but not on adjacent leaf parts, suffered from reduced growth. This effect was abolished in the BX-deficient bx1 mutant. We did not find any evidence that BXs regulate defensive phytohormones or their own accumulation. In summary, this study shows that induced herbivore resistance in maize is highly localized and dependent on BXs.

An ethylene response-related factor, GbERF1-like, from Gossypium barbadense improves resistance to Verticillium dahliae via activating lignin synthesis.

An ethylene response-related factor, GbERF1-like, from Gossypium barbadense cv. '7124' involved in the defence response to Verticillium dahliae was characterized. GbERF1-like transcripts present ubiquitously in various tissues, with higher accumulation in flower organs. GbERF1-like was also responsive to defence-related phytohormones and V. dahliae infection. The downregulation of GbERF1-like increased the susceptibility of cotton plants to V. dahliae infection, while overexpression of this gene improved disease resistance in both cotton and Arabidopsis, coupled with activation of the pathogenesis-related proteins. Further analysis revealed that genes involved in lignin synthesis, such as PAL, C4H, C3H, HCT, CCoAOMT, CCR and F5H, showed higher expression levels in the overexpressing cotton and Arabidopsis lines and lower expression levels in the RNAi cotton lines. The expression levels of these genes increased obviously when the GbERF1-like-overexpressing plants were inoculated with V. dahliae. Meanwhile, significant differences in the content of whole lignin could be found in the stems of transgenic and wild-type plants after inoculation with V. dahliae, as revealed by metabolic and histochemical analysis. More lignin could be detected in GbERF1-like-overexpressing cotton and Arabidopsis but less in GbERF1-like-silencing cotton compared with wild-type plants. The ratio of S and G monomers in GbERF1-like-overexpressing cotton and Arabidopsis increased significantly after infection by V. dahliae. Moreover, our results showed that the promoters of GhHCT1 and AtPAL3 could be transactivated by GbERF1-like in vivo based on yeast one-hybrid assays and dual-luciferase reporter assays. Knockdown of GhHCT1 in GbERF1-like over-expressing cotton decreases resistance to V. dahliae. Collectively, our results suggest that GbERF1-like acts as a positive regulator in lignin synthesis and contributes substantially to resistance to V. dahliae in plants.

Expression profiling of marker genes responsive to the defence-associated phytohormones salicylic acid, jasmonic acid and ethylene in Brachypodium distachyon.

BackgroundBrachypodium distachyon is a promising model plants for grasses. Infections of Brachypodium by various pathogens that severely impair crop production have been reported, and the species accordingly provides an alternative platform for investigating molecular mechanisms of pathogen virulence and plant disease resistance. To date, we have a broad picture of plant immunity only in Arabidopsis and rice; therefore, Brachypodium may constitute a counterpart that displays the commonality and uniqueness of defence systems among plant species. Phytohormones play key roles in plant biotic stress responses, and hormone-responsive genes are used to qualitatively and quantitatively evaluate disease resistance responses during pathogen infection. For these purposes, defence-related phytohormone marker genes expressed at time points suitable for defence-response monitoring are needed. Information about their expression profiles over time as well as their response specificity is also helpful. However, useful marker genes are still rare in Brachypodium.ResultsWe selected 34 candidates for Brachypodium marker genes on the basis of protein-sequence similarity to known marker genes used in Arabidopsis and rice. Brachypodium plants were treated with the defence-related phytohormones salicylic acid, jasmonic acid and ethylene, and their transcription levels were measured 24 and 48 h after treatment. Two genes for salicylic acid, 7 for jasmonic acid and 2 for ethylene were significantly induced at either or both time points. We then focused on 11 genes encoding pathogenesis-related (PR) 1 protein and compared their expression patterns with those of Arabidopsis and rice. Phylogenetic analysis suggested that Brachypodium contains several PR1-family genes similar to rice genes. Our expression profiling revealed that regulation patterns of some PR1 genes as well as of markers identified for defence-related phytohormones are closely related to those in rice.ConclusionWe propose that the Brachypodium immune hormone marker genes identified in this study will be useful to plant pathologists who use Brachypodium as a model pathosystem, because the timing of their transcriptional activation matches that of the disease resistance response. Our results using Brachypodium also suggest that monocots share a characteristic immune system, defined as the common defence system, that is different from that of dicots.Electronic supplementary materialThe online version of this article (doi:10.1186/s12870-016-0749-9) contains supplementary material, which is available to authorized users.

Trichoderma atroviride LU132 promotes plant growth but not induced systemic resistance to Plutella xylostella in oilseed rape

Several species of the fungus Trichoderma can promote plant health and are widely used as commercial biopesticides. Beneficial effects of this fungus are attributed to various mechanisms such as mycoparasitism, plant-growth promotion, increased stress tolerance and elicitation of induced systemic resistance against pathogens via jasmonic acid/ethylene-dependent pathways. Despite such well-established effects on pathogens, surprisingly little is known about the influence of Trichoderma on plant defences against herbivorous insects. This study investigated whether soil-supplementation of the established biocontrol agent Trichoderma atroviride LU132 affected the performance of oilseed rape (Brassica napus) and the development of Plutella xylostella caterpillars. Furthermore, induction and priming of defence-related phytohormones, genes and secondary metabolites by fungus and herbivore were assessed. Plants colonized by T. atroviride LU132 had significantly larger root and shoot biomass than controls. No effects of fungal inoculation were found on herbivore development. Leaf feeding of the herbivore induced higher jasmonic acid levels, but this was not influenced by fungal treatment. Similarly, the defence-related genes MYC2 and TPI were induced by herbivory but not primed or induced by T. atroviride. Expression of the gene PDF1.2 was repressed by herbivore feeding while no effects on the gene ACO and glucosinolates were observed. We conclude that T. atroviride LU132 has positive effects on the growth of oilseed but it does not enhance above-ground insect defences.

AtMYB44 positively modulates disease resistance to Pseudomonas syringae through the salicylic acid signalling pathway in Arabidopsis.

Plant MYB transcription factors are implicated in resistance to biotic and abiotic stresses. Here, we demonstrate that an R2-R3 MYB transcription factor, AtMYB44, plays a role in the plant defence response to the bacterial pathogen Pseudomonas syringae pv. tomato DC3000 (PstDC3000). The expression of AtMYB44 was upregulated upon pathogen infection and treatments with defence-related phytohormones. Transgenic plants overexpressing AtMYB44 (35S-Ms) exhibited greater levels of PR1 gene expression, cell death, callose deposition and hydrogen peroxide (H2O2) accumulation in leaves infected with PstDC3000. Consequently, 35S-M lines displayed enhanced resistance to PstDC3000. In contrast, the atmyb44 T-DNA insertion mutant was more susceptible to PstDC3000 and exhibited decreased PR1 gene expression upon infection. Using double mutants constructed via crosses of 35S-M lines with NahG transgenic plants and nonexpressor of pathogenesis-related genes1 mutant (npr1-1), we demonstrated that the enhanced PR1 gene expression and PstDC3000 resistance in 35S-M plants occur mainly through the salicylic acid signalling pathway.

Modulation of ethylene- and heat-controlled hyponastic leaf movement in Arabidopsis thaliana by the plant defence hormones jasmonate and salicylate

Upward leaf movement (hyponastic growth) is adopted by several plant species including Arabidopsis thaliana, as a mechanism to escape adverse growth conditions. Among the signals that trigger hyponastic growth are, the gaseous hormone ethylene, low light intensities, and supra-optimal temperatures (heat). Recent studies indicated that the defence-related phytohormones jasmonic acid (JA) and salicylic acid (SA) synthesized by the plant upon biotic infestation repress low light-induced hyponastic growth. The hyponastic growth response induced by high temperature (heat) treatment and upon application of the gaseous hormone ethylene is highly similar to the response induced by low light. To test if these environmental signals induce hyponastic growth via parallel pathways or converge downstream, we studied here the roles of Methyl-JA (MeJA) and SA on ethylene- and heat-induced hyponastic growth. For this, we used a time-lapse camera setup. Our study includes pharmacological application of MeJA and SA and biological infestation using the JA-inducing caterpillar Pieris rapae as well as mutants lacking JA or SA signalling components. The data demonstrate that MeJA is a positive, and SA, a negative regulator of ethylene-induced hyponastic growth and that both hormones repress the response to heat. Taking previous studies into account, we conclude that SA is the first among many tested components which is repressing hyponastic growth under all tested inductive environmental stimuli. However, since MeJA is a positive regulator of ethylene-induced hyponastic growth and is inhibiting low light- and heat-induced leaf movement, we conclude that defence hormones control hyponastic growth by affecting stimulus-specific signalling pathways.

Parasitism by Cuscuta pentagona sequentially induces JA and SA defence pathways in tomato

While plant responses to herbivores and pathogens are well characterized, responses to attack by other plants remain largely unexplored. We measured phytohormones and C(18) fatty acids in tomato attacked by the parasitic plant Cuscuta pentagona, and used transgenic and mutant plants to explore the roles of the defence-related phytohormones salicylic acid (SA) and jasmonic acid (JA). Parasite attachment to 10-day-old tomato plants elicited few biochemical changes, but a second attachment 10 d later elicited a 60-fold increase in JA, a 30-fold increase in SA and a hypersensitive-like response (HLR). Host age also influenced the response: neither Cuscuta seedlings nor established vines elicited a HLR in 10-day-old hosts, but both did in 20-day-old hosts. Parasites grew larger on hosts deficient in SA (NahG) or insensitive to JA [jasmonic acid-insensitive1 (jai1)], suggesting that both phytohormones mediate effective defences. Moreover, amounts of JA peaked 12 h before SA, indicating that defences may be coordinated via sequential induction of these hormones. Parasitism also induced increases in free linolenic and linoleic acids and abscisic acid. These findings provide the first documentation of plant hormonal signalling induced by a parasitic plant and show that tomato responses to C. pentagona display characteristics similar to both herbivore- and pathogen-induced responses.