Abstract
Loss-of-function alleles of MLO (Mildew Resistance Locus O) confer broad-spectrum resistance to foliar infections by powdery mildew pathogens. Like pathogens, microbes that establish mutually beneficial relationships with their plant hosts, trigger the induction of some defense responses. Initially, barley colonization by the root endophyte Serendipita indica (syn. Piriformospora indica) is associated with enhanced defense gene expression and the formation of papillae at sites of hyphal penetration attempts. This phenotype is reminiscent of mlo-conditioned immunity in barley leaf tissue and raises the question whether MLO plays a regulatory role in the establishment of beneficial interactions. Here we show that S. indica colonization was significantly reduced in plants carrying mlo mutations compared to wild type controls. The reduction in fungal biomass was associated with the enhanced formation of papillae. Moreover, epidermal cells of S. indica-treated mlo plants displayed an early accumulation of iron in the epidermal layer suggesting increased basal defense activation in the barley mutant background. Correspondingly, the induction of host cell death during later colonization stages was impaired in mlo colonized plants, highlighting the importance of the early biotrophic growth phase for S. indica root colonization. In contrast, the arbuscular mycorrhizal fungus Funneliformis mosseae displayed a similar colonization morphology on mutant and wild type plants. However, the frequency of mycorrhization and number of arbuscules was higher in mlo-5 mutants. These findings suggest that MLO differentially regulates root colonization by endophytic and AM fungi.
Highlights
Plants establish diverse beneficial interactions with fungi from different taxa
To assess the effect of the mlo-5 mutation on S. indica colonization, we quantified the relative abundance of S. indica gDNA as a proxy for fungal biomass in roots of Wild type (WT) barley cv
Similar to mlo-5, mlo-3 and -4 were less colonized by S. indica compared to WT control plants (Supplementary Figure 1), suggesting that MLO is required for barley root colonization by S. indica
Summary
Root endophytes belonging to the order Sebacinales establish long-lasting beneficial relationships with a broad range of plant species (Weiss et al, 2016). In some cases nutrient status has been reported to play a role in the interaction of the model sebacinoid fungus Serendipita indica with some plant species (Shahollari et al, 2005; Sherameti et al, 2005; Nautiyal et al, 2010; Yadav et al, 2010; Kumar et al, 2012). Following an initial biotrophic growth phase, during which the fungal hyphae remain surrounded by a plant-derived membrane, S. indica transitions to cell death-associated colonization that does not result in host disease (Deshmukh et al, 2006; Jacobs et al, 2011; Zuccaro et al, 2011; Lahrmann and Zuccaro, 2012; Qiang et al, 2012; Lahrmann et al, 2013)
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