It is well documented that Epichloë endophytes can enhance the resistance of grasses to herbivory. However, reports on resistance to pathogenic fungi are limited, and their conclusions are variable. In this study, we chose pathogenic fungi with different trophic types, namely, the biotrophic pathogen Erysiphales species and the necrotrophic pathogen Curvularia lunata, to test the effects of Epichloë on the pathogen resistance of Achnatherum sibiricum. The results showed that, compared to Erysiphales species, C. lunata caused a higher degree of damage and lower photochemical efficiency (Fv/Fm) in endophyte−free (E−) leaves. Endophytes significantly alleviated the damage caused by these two pathogens. The leaf damaged area and Fv/Fm of endophyte−infected (E+) leaves were similar between the two pathogen treatments, indicating that the beneficial effects of endophytes were more significant when hosts were exposed to C. lunata than when they were exposed to Erysiphales species. We found that A. sibiricum initiated jasmonic acid (JA)−related pathways to resist C. lunata but salicylic acid (SA)-related pathways to resist Erysiphales species. Endophytic fungi had no effect on the content of SA but increased the content of JA and total phenolic compounds, which suggest that endophyte infection might enhance the resistance of A. sibiricum to these two different trophic types of pathogens through similar pathways.

Aims Many grasses are infected by systemic fungal endophytes that occur in aboveground plant tissues. Both aboveground endophytic fungi and belowground soil microbes can influence plant growth, but studies on their simultaneous effects on plant growth and competitiveness are limited. This study aims to investigate whether the role of aboveground endophytic fungi in the growth and competitive ability of the host grasses was influenced by soil microbes. Methods In this study, we used Epichloë endophyte-infected (EI) and endophyte-free (EF) Achnatherum sibiricum as plant materials. A wet sieving method was adopted to obtain microbial inocula with different diversities (com, combined microbe fraction, high diversity; sm, small microbe fraction, low diversity). A three-factor randomized block design was used. The first factor was the endophyte infection status of A. sibiricum. The second factor was the microbial composition of the soil inocula. The third factor was the planting type. Growth and competitive characters were measured after 16 weeks. Important Findings The results showed that a soil microbe inoculation was detrimental to the growth of A. sibiricum. Epichloë endophytes significantly mitigated the inhibitory effect of soil microbes on A. sibiricum planted alone. When A. sibiricum was planted with Stipa grandis, there was a significant interaction between Epichloë endophytes and soil microbes on the interspecific competition of A. sibiricum. When inoculated with small microbial community fraction, Epichloë endophytes significantly improved the interspecific competitive ability of host plants. When inoculated with combined microbial community fraction, however, Epichloë endophytes had no significant effect on host competition. The results showed that the interaction between Epichloë endophytes and soil microbes contributed more to the interspecific competitive ability than either Epichloë endophytes or soil microbes alone.

Epichloë endophytes may not only affect the growth and resistances of host grasses, but may also affect soil environment including soil microbes. Can Epichloë endophyte-mediated modification of soil microbes affect the competitive ability of host grasses? In this study, we tested whether Epichloë endophytes and soil microbes alter intraspecific competition between Epichloë endophyte-colonized (EI) and endophyte-free (EF) Leymus chinensis and interspecific competition between L. chinensis and Stipa krylovii. The results demonstrated that Epichloë endophyte colonization significantly enhanced the intraspecific competitive ability of L. chinensis and that this beneficial effect was not affected by soil microbes. Under interspecific competition, however, significant interactions between Epichloë endophytes and soil microbes were observed. The effect of Epichloë endophytes on interspecific competitiveness of the host changed from positive to neutral with soil microbe removal. Here higher mycorrhizal colonization rates probably contributed to interspecific competitive advantages of EI over EF L. chinensis. Our result suggests that Epichloë endophytes can influence the competitive ability of the host through plant soil feedbacks from the currently competing plant species.

Epichloë endophytes can improve the resistance of host grasses to pathogenic fungi, but the underlying mechanisms remain largely unknown. Here, we used phytohormone quantifications, gene expression analysis and pathogenicity experiments to investigate the effect of Epichloë sibirica on the resistance of Achnatherum sibiricum to Curvularia lunata pathogens. Comparison of gene expression patterns between endophyte-infected and endophyte-free leaves revealed that endophyte infection was associated with significant induction of 1758 and 765 differentially expressed genes in the host before and after pathogen inoculation, respectively. Functional analysis of the differentially expressed genes suggested that endophyte infection may activate the constitutive resistance of the host by increasing photosynthesis, enhancing the ability to scavenge reactive oxygen species, and actively regulating the expression of genes with function related to disease resistance. We found that endophyte infection was associated with induction of the expression of genes involved in the biosynthesis pathways of jasmonic acid, ethylene and pipecolic acid and amplified the defense response of jasmonic acid/ethylene co-regulated EIN/ERF1 transduction pathway and Pip-mediated TGA transduction pathway. Phytohormone quantifications showed that endophyte infection was associated with significant accumulation of jasmonic acid, ethylene and pipecolic acid after pathogen inoculation. Exogenous phytohormone treatments confirmed that the disease index of plants was negatively related to both jasmonic acid and ethylene concentrations. Our results demonstrate that endophyte infection can not only improve the constitutive resistance of the host to phytopathogens before pathogen inoculation but also be associated with enhanced systemic resistance of the host to necrotrophs after C. lunata inoculation.

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