Genome-wide DNA hypomethylation shapes nematode pattern-triggered immunity in plants.

A role for DNA hypomethylation has recently been suggested in the interaction between bacteria and plants; it is unclear whether this phenomenon reflects a conserved response. Treatment of plants of monocot rice and dicot tomato with nematode-associated molecular patterns from different nematode species or bacterial pathogen-associated molecular pattern flg22 revealed global DNA hypomethylation. A similar hypomethylation response was observed during early gall induction by Meloidogyne graminicola in rice. Evidence for the causal impact of hypomethylation on immunity was revealed by a significantly reduced plant susceptibility upon treatment with DNA methylation inhibitor 5-azacytidine. Whole-genome bisulphite sequencing of young galls revealed massive hypomethylation in the CHH context, while not for CG or CHG nucleotide contexts. Further, CHH hypomethylated regions were predominantly associated with gene promoter regions, which was not correlated with activated gene expression at the same time point but, rather, was correlated with a delayed transcriptional gene activation. Finally, the relevance of CHH hypomethylation in plant defence was confirmed in rice mutants of the RNA-directed DNA methylation pathway and DECREASED DNA METHYLATION 1. We demonstrated that DNA hypomethylation is associated with reduced susceptibility in rice towards root-parasitic nematodes and is likely to be part of the basal pattern-triggered immunity response in plants.

Tomato photorespiratory glycolate-oxidase-derived H2 O2 production contributes to basal defence against Pseudomonas syringae.

Despite being essential for C3 plants, photorespiration is believed to cause a significant crop yield loss even under future climates. However, how photorespiration affects plant basal defence still remains largely unknown. Here, we studied the involvement of photorespiration in tomato-Pseudomonas syringae pv. tomato DC3000 interaction focusing on three photorespiratory genes. Inoculation with P. syringae increased photorespiration rate (Pr) and expression of glycolate oxidase (GOX2), serine glyoxylate aminotransferase (SGT) and serine hydroxyl methyltransferase (SHMT1); however, inhibition of photorespiration by isonicotinic acid hydrazide decreased tomato basal defence against P. syringae. Furthermore, silencing of GOX2, SGT or SHMT1 genes in tomato decreased Pr but increased susceptibility to P. syringae, whereas transient overexpression of GOX2, SGT or SHMT1 in tobacco increased basal defence. Further study revealed that salicylic acid (SA) signalling is involved in GOX2-mediated, SGT-mediated and SHMT1-mediated defence. Moreover, H2 O2 pretreatment remarkably alleviated the GOX2 silencing-induced depression in basal defence and SA signalling, whereas it had no effect on that of SGT-silenced and SHMT1-silenced plants. Taken together, these results suggest that H2 O2 is critical for GOX2-modulated but not SGT-modulated or SHMT1-modulated SA signalling and subsequent basal defence against P. syringae. This work deepens the understanding of photorespiration-involved defence responses to bacterial attack in plants.

Solanaceous exocyst subunits are involved in immunity to diverse plant pathogens.

The exocyst, a multiprotein complex consisting of eight subunits, plays an essential role in many biological processes by mediating secretion of post-Golgi-derived vesicles towards the plasma membrane. In recent years, roles for plant exocyst subunits in pathogen defence have been uncovered, largely based on studies in the model plant Arabidopsis. Only a few studies have been undertaken to assign the role of exocyst subunits in plant defence in other plants species, including crops. In this study, predicted protein sequences from exocyst subunits were retrieved by mining databases from the Solanaceous plants Nicotiana benthamiana, tomato, and potato. Subsequently, their evolutionary relationship with Arabidopsis exocyst subunits was analysed. Gene silencing in N. benthamiana showed that several exocyst subunits are required for proper plant defence against the (hemi-)biotrophic plant pathogens Phytophthora infestans and Pseudomonas syringae. In contrast, some exocyst subunits seem to act as susceptibility factors for the necrotrophic pathogen Botrytis cinerea. Furthermore, the majority of the exocyst subunits were found to be involved in callose deposition, suggesting that they play a role in basal plant defence. This study provides insight into the evolution of exocyst subunits in Solanaceous plants and is the first to show their role in immunity against multiple unrelated pathogens.

Plant basal resistance to nematodes: an update.

Most plant-parasitic nematodes are obligate biotrophs feeding on the roots of their hosts. Whereas ectoparasites remain on the root surface and feed on the outer cell layers, endoparasitic nematodes enter the host to parasitize cells around or within the central cylinder. Nematode invasion and feeding causes tissue damage which may, in turn, lead to the activation of host basal defence responses. Hitherto, research interests in plant-nematode interaction have emphasized effector-triggered immunity rather than basal plant defence responses. However, some recent investigations suggest that basal defence pathways are not only activated but also play an important role in determining interaction outcomes. In this review we discuss the major findings and point out future directions to dissect the molecular mechanisms underlying plant basal defence to nematodes further.

The role of cell wall-based defences in the early restriction of non-pathogenic hrp mutant bacteria in Arabidopsis.

We have investigated the cause of the restricted multiplication of hrp mutant bacteria in leaves of Arabidopsis. Our focus was on early interactions leading to differentiation between virulent wild-type and non-pathogenic hrpA mutant strains of Pseudomonas syringae pv. tomato. An initial drop in recoverable bacteria detected 0-4 h after inoculation with either strain was dependent on a functional FLS2 receptor and H2O2 accumulation in challenged leaves. Wild-type bacteria subsequently multiplied rapidly whereas the hrpA mutant was restricted within 6 h. Despite the early restriction, the hrpA mutant was still viable several days after inoculation. Analysis of intercellular washing fluids (IWFs), showed that high levels of nutrients were readily available to bacteria in the apoplast and that no diffusible inhibitors were produced in response to bacterial challenge. Histochemical and immunocytochemical methods were used to detect changes in polysaccharides (callose, two forms of cellulose, and pectin), arabinogalactan proteins (AGPs), H2O2 and peroxidase. Quantitative analysis showed very similar changes in localisation of AGPs, cellulose epitopes and callose 2 and 4 h after inoculation with either strain. However from 6 to 12 h after inoculation papillae expanded only next to the hrp mutant. In contrast to the similar patterns of secretory activity recorded from mesophyll cells, accumulation of H2O2 and peroxidase was significantly greater around the hrpA mutant within the first 4h after inoculation. A striking differential accumulation of H2O2 was also found in chloroplasts in cells next to the mutant. Ascorbate levels were lower in the IWFs recovered from sites inoculated with the hrp mutant than with wild-type bacteria. The critical response, observed at the right time and place to explain the observed differential behaviour of wild-type and hrpA mutant bacteria was the accumulation of H2O2, probably generated through Type III peroxidase activity and in chloroplasts. It is proposed that H2O2 and apoplastic peroxidase cross-link secreted glycoproteins and polysaccharides to agglutinate the hrp mutant. Generation of H2O2 has been identified as a likely target for effector proteins injected into plant cells by the wild-type bacteria.