Take-all disease of Poaceae is caused by Gaeumannomyces graminis (Magnaporthaceae). Four varieties are recognised in G. graminis based on ascospore size, hyphopodial morphology and host preference. The aim of the present study was to clarify boundaries among species and varieties in Gaeumannomyces by combining morphology and multi-locus phylogenetic analyses based on partial gene sequences of ITS, LSU, tef1 and rpb1. Two new genera, Falciphoriella and Gaeumannomycella were subsequently introduced in Magnaporthaceae. The resulting phylogeny revealed several cryptic species previously overlooked within Gaeumannomyces. Isolates of Gaeumannomyces were distributed in four main clades, from which 19 species could be delimited, 12 of which were new to science. Our results show that the former varieties Gaeumannomyces graminis var. avenae and Gaeumannomyces graminis var. tritici represent species phylogenetically distinct from G. graminis, for which the new combinations G. avenae and G. tritici are introduced. Based on molecular data, morphology and host preferences, Gaeumannomyces graminis var. maydis is proposed as a synonym of G. radicicola. Furthermore, an epitype for Gaeumannomyces graminis var. avenae was designated to help stabilise the application of that name.

BackgroundAncestral wheat relatives are important sources of genetic diversity for the introduction of novel traits for the improvement of modern bread wheat. In this study the aim was to assess the susceptibility of 34 accessions of the diploid wheat Triticum monococcum (A genome) to Gaeumannomyces graminis var. tritici (Ggt), the causal agent of take-all disease. The second aim was to explore the susceptibility of tetraploid wheat (T. durum) and the B genome progenitor species Aegilops speltoides to Ggt.ResultsField trials, conducted over 5 years, identified seven T. monococcum accessions with a good level of resistance to take-all when exposed to natural inoculum under UK field conditions. All other accessions were highly susceptible or did not exhibit a consistent phenotype across years. DArT marker genotyping revealed that whole genome diversity was not closely related to resistance to take-all within T. monococcum, suggesting that multiple genetic sources of resistance may exist within the species. In contrast the tetraploid wheat cultivars and Ae. speltoides were all highly susceptible to the disease, including those with known elevated levels of benzoxazinoids.ConclusionsThe diploid wheat species T. monococcum may provide a genetic source of resistance to take-all disease that could be utilised to improve the performance of T. aestivum in high disease risk situations. This represents an extremely valuable resource to achieve economic and sustainable genetic control of this root disease.

SummaryManipulation of the soil microbiota associated with crop plants has huge promise for the control of crop pathogens. However, to fully realize this potential we need a better understanding of the relationship between the soil environment and the genes and phenotypes that enable microbes to colonize plants and contribute to biocontrol. A recent 2 years of investigation into the effect of wheat variety on second year crop yield in the context of take‐all fungal infection presented the opportunity to examine soil microbiomes under closely defined field conditions. Amplicon sequencing of second year soil samples showed that P seudomonas spp. were particularly affected by the wheat cultivar grown in year one. Consequently, 318 rhizosphere‐associated P seudomonas fluorescens strains were isolated and characterized across a variety of genetic and phenotypic traits. Again, the wheat variety grown in the first year of the study was shown to exert considerable selective pressure on both the extent and nature of P seudomonas genomic diversity. Furthermore, multiple significant correlations were identified within the phenotypic/genetic structure of the Pseudomonas population, and between individual genotypes and the external wheat field environment. The approach outlined here has considerable future potential for our understanding of plant–microbe interactions, and for the broader analysis of complex microbial communities.

Biodiversity and ecosystem functioning research typically shows positive diversity- productivity relationships. However, local increases in species richness can increase competition within trophic levels, reducing the efficacy of intertrophic level population control. Pseudomonas spp. are a dominant group of soil bacteria that play key roles in plant growth promotion and control of crop fungal pathogens. Here we show that Pseudomonas spp. richness is positively correlated with take-all disease in wheat and with yield losses of ~3 t/ha in the field. We modeled the interactions between Pseudomonas and the take-all pathogen in abstract experimental microcosms, and show that increased bacterial genotypic richness escalates bacterial antagonism and decreases the ability of the bacterial community to inhibit growth of the take-all pathogen. Future work is required to determine the generality of these negative biodiversity effects on different media and directly at infection zones on root surfaces. However, the increase in competition between bacteria at high genotypic richness and the potential loss of fungal biocontrol activity highlights an important mechanism to explain the negative Pseudomonas diversity-wheat yield relationship we observed in the field. Together our results suggest that the effect of biodiversity on ecosystem functioning can depend on both the function and trophic level of interest.

The effect of wheat cultivar on the build-up of take-all inoculum during a first wheat crop was measured after harvest using a soil core bioassay in field experiments over five growing seasons (2003)(2004)(2005)(2006)(2007)(2008). Cultivar differences in individual years were explored by analysis of variance and a cross-season Residual Maximum Likelihood (REML) variance components analysis was used to compare differences in those cultivars present in all years. Differences between cultivars in the build-up of inoculum were close to or at significance in two of the five trial years (2004 P < 0AE05; 2006 P < 0AE07), and current commercially listed cultivars were represented at both extremes of the range. In 2007 and 2008, when environmental conditions were most favourable for inoculum build-up, differences were not significant (P < 0AE3). In 2005 the presence of Phialophora spp. at the trial site restricted the build-up of take-all inoculum under all cultivars. The cross season REML variance components analysis detected significant differences (range: 3AE4-47AE8% roots infected in the soil core bioassay; P < 0AE01) between the nine cultivars present in all years (excluding 2005). This is the first evidence of relatively consistent differences between hexaploid wheat cultivars in their interactions with the take-all fungus, and this could give an indication of those cultivars that could be grown as a first wheat crop, in order to reduce the risk of damaging take-all in a second wheat crop. This phenomenon has been named the take-all inoculum build-up (TAB) trait.

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Given the increasing demand for wheat which is forecast, cropping of wheat in short rotations will likely remain a common practice. However, in temperate wheat growing regions the soil-borne fungal pathogen Gaeumannomyces tritici becomes a major constraint on productivity. In cultivar rotation field experiments on the Rothamsted Farm (Hertfordshire, UK) we demonstrated a substantial reduction in take-all disease and grain yield increases of up to 2.4 tonnes/ha when a low take-all inoculum building wheat cultivar was grown in the first year of wheat cropping. Phenotyping of 71 modern elite wheat cultivars for the take-all inoculum build-up trait across six diverse trial sites identified a few cultivars which exhibited a consistent lowering of take-all inoculum build-up. However, there was also evidence of a significant interaction effect between trial site and cultivar when a pooled Residual Maximum Likelihood (REML) procedure was conducted. There was no evidence of an unusual rooting phenotype associated with take-all inoculum build-up in two independent field experiments and a sand column experiment. Together our results highlight the complex interactions between wheat genotype, environmental conditions and take-all inoculum build-up. Further work is required to determine the underlying genetic and mechanistic basis of this important phenomenon.