Salicylic acid-dependent and -independent impact of an RNA-binding protein on plant immunity.

Plants overexpressing the RNA-binding protein AtGRP7 (AtGRP7-ox plants) constitutively express the PR-1 (PATHOGENESIS-RELATED-1), PR-2 and PR-5 transcripts associated with salicylic acid (SA)-mediated immunity and show enhanced resistance against Pseudomonas syringae pv. tomato (Pto) DC3000. Here, we investigated whether the function of AtGRP7 in plant immunity depends on SA. Endogenous SA was elevated fivefold in AtGRP7-ox plants. The elevated PR-1, PR-2 and PR-5 levels were eliminated upon expression of the salicylate hydroxylase nahG in AtGRP7-ox plants and elevated PR-1 levels were suppressed by sid (salicylic acid deficient) 2-1 that is impaired in SA biosynthesis. RNA immunoprecipitation showed that AtGRP7 does not bind the PR-1 transcript in vivo, whereas it binds PDF1.2. Constitutive or inducible AtGRP7 overexpression increases PR-1 promoter activity, indicating that AtGRP7 affects PR-1 transcription. In line with this, the effect of AtGRP7 on PR-1 is suppressed by npr (non-expressor of PR genes) 1. Whereas AtGRP7-ox plants restricted growth of Pto DC3000 compared with wild type (wt), sid2-1 AtGRP7-ox plants allowed more growth than AtGRP7-ox plants. Furthermore, we show an enhanced hypersensitive response triggered by avirulent Pto DC3000 (AvrRpt2) in AtGRP7-ox compared with wt. In sid2-1 AtGRP7-ox, an intermediate phenotype was observed. Thus, AtGRP7 has both SA-dependent and SA-independent effects on plant immunity.

Effects of root herbivory by nematodes on the performance and preference of a leaf-infesting generalist aphid depend on nitrate fertilization.

The performance and behavior of herbivores is strongly affected by the quality of their host plants, which is determined by various environmental conditions. We investigated the performance and preference of the polyphagous shoot-infesting aphid Myzus persicae on the host-plant Arabidopsis thaliana in a two-factorial design in which nitrate fertilization was varied by 33 %, and the root-infesting cyst-nematode Heterodera schachtii was present or absent. Aphid performance was influenced by these abiotic and biotic factors in an interactive way. Nematode presence decreased aphid performance when nitrate levels were low, whereas nematode infestation did not influence aphid performance under higher nitrate fertilization. Aphids followed the "mother knows best" principle when given a choice, settling preferentially on those plants on which they performed best. Hence, they preferred nematode-free over nematode-infested plants in the low fertilization treatment but host choice was not affected by nematodes under higher nitrate fertilization. The amino acid composition of the phloem exudates was significantly influenced by fertilization but also by the interaction of the two treatments. Various glucosinolates in the leaves, which provide an estimate of phloem glucosinolates, were not affected by the individual treatments but by the combination of fertilization and herbivory. These changes in primary and secondary metabolites may be decisive for the herbivore responses. Our data demonstrate that abiotic and biotic factors can interactively affect herbivores, adding a layer of complexity to plant-mediated herbivore interactions.

Plant-mediated interactions between shoot-feeding aphids and root-feeding nematodes depend on nitrate fertilization.

The plant metabolite composition is modulated by various abiotic and biotic factors including nutrient availability and herbivory. In turn, induced changes in plant quality can affect herbivore performance and mediate indirect interactions between spatially separated herbivores sharing a host. Studies on plant-mediated herbivore interactions have been carried out at single fertilization regimes only, but we hypothesized that nutrient availability modifies these interactions. Therefore, we studied the interactions between two vascular tissue herbivores, the aboveground feeding aphid Brevicoryne brassicae and the belowground infesting nematode Heterodera schachtii, on Arabidopsis thaliana grown under two nitrate fertilization conditions (varying by 33 %). Furthermore, we investigated plant growth and primary metabolic responses to fertilization and herbivore treatments, which could potentially mediate these interactions, as the herbivores may act as metabolic sinks. Whereas nematodes had no effects on aphids, aphid presence influenced nematodes in opposite directions, depending on fertilization: at low nitrate supply, aphids had a promoting effect on nematodes, whereas at high nitrate fertilization they lowered the nematode infestation compared to control plants. Plants produced significantly more biomass under high nitrate supply but C and N contents were not altered. Primary metabolite profiles differed only marginally between roots of both fertilization treatments in plants with and without aphids, indicating that nematodes may respond to these or other metabolic modifications, which are caused by minute environmental changes, in a sensitive way. Our results highlight the need to consider the importance of plant nutrient availability on the outcome of interactions between co-occurring herbivores in future studies.

Crosstalk between above- and belowground herbivores is mediated by minute metabolic responses of the host Arabidopsis thaliana.

Plants are frequently under attack by multiple herbivores and can be infested at their shoots as well as their roots. As a consequence, plant metabolites are readily induced, mediated by phytohormones such as salicylic acid and jasmonic acid. Thereby, cross-talk between signal transduction pathways may occur if different herbivores attack the plant simultaneously. In turn, modifications in the plant metabolic pattern can affect herbivores infesting local and systemic tissue. Here, an integrative approach combining metabolomics and performance experiments was used to study the induction of plant metabolites in Arabidopsis thaliana by the specialist aphid Brevicoryne brassicae feeding on shoots and the generalist nematode Heterodera schachtii infesting root tissue. In contrast to most other studies, low infestation rates typical for the decisive early stages of infestation were used. Moreover, the consequences of induction responses on plant-mediated indirect interactions between these herbivores were investigated. In aphid-treated plants, several metabolites including glucosinolates, important defence compounds of Brassicaceae, were reduced in the shoot, but only minute changes took part in the systemic root tissue. Primary metabolites as well as phytohormones were not altered 3 days post infestation. In contrast, nematodes did not evoke significant metabolic alterations locally or systemically. In accordance, nematode presence did not affect aphid population growth, whereas aphids mediated a considerably reduced nematode infestation. These results demonstrate that plants respond in a very fine-tuned way to different challenges. Although they show only minute systemic responses to low herbivore stress, these changes can have pronounced effects on plant-mediated interactions between herbivores.

The potential of cladocerans as controphic competitors of the mosquito Culex pipiens.

This study assesses the potential of cladocerans as competitors for controlling the oviposition and larval abundances of mosquitoes. Control of mosquito larvae involving the use of antagonists has focused mostly on predators. We hypothesized that cladoceran competitors have a strong potential to control larval populations of some species of mosquitoes that can be early colonizers of newly-filled waterbodies, and should be less efficient competitors. To test this hypothesis, the establishment and development of larval populations of wild Culex pipiens were investigated in outdoor microcosms varying in terms of Daphnia magna populations. When the population was well established (i.e., high densities of D. magna), oviposition was fully inhibited and there was consequently no mosquito larvae. When the population was more recently established (i.e., lower densities of D. magna), oviposition and larval development of Cx. pipiens occurred. In the absence of D. magna, oviposition, larval biomass, and abundance of Cx. pipiens reached high values. In this situation, conspecifics were inhibiting further oviposition of Cx. pipiens. Based on these results, we suggest that competing zooplankton species, such as D. magna, could be used for the control of mosquito species such as Cx. pipiens. This approach could be beneficial for the management of wetlands.

Local and systemic transcriptional responses to crosstalk between above- and belowground herbivores in Arabidopsis thaliana.

Plants are often simultaneously infested by several herbivores at the shoots and roots. Recent results revealed that the model plant Arabidopsis thaliana shows highly challenge-specific local and systemic responses to individual and simultaneous attacks of shoot-infesting aphids and root-infesting nematodes at the metabolome level. (1) Here, we present the corresponding transcriptional changes in plants treated with Brevicoryne brassicae aphids and Heterodera schachtii nematodes individually and in combination. Overall, shoots were much less responsive than roots. Gene expression in shoots and roots was mainly altered by aphids. Nematode infestation alone had only little effect, but nematodes modified the transcript accumulation response to aphids particularly in the roots. The responding genes are involved in plant defense cascades, signaling, oxidation-reduction processes, as well as primary and secondary metabolism and degradation. These changes in transcription may become relevant for the herbivores when they are translated into changes in host plant quality.