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A trade‐off between growth and defence against biotic stresses is common in plants. Fungal endophytes of the genus Epichloë may relieve this trade‐off in their host grasses since they can simultaneously induce plant growth and produce antiherbivore alkaloids that circumvent the need for host defence. The Epichloë ability to decouple the growth‐defence trade‐off was evaluated by subjecting ryegrass with and without Epichloë endophytes to an exogenous treatment with gibberellin (GA) followed by a challenge with Rhopalosiphum padi aphids. In agreement with the endophyte‐mediated trade‐off decoupling hypothesis, the GA‐derived promotion of plant growth increased the susceptibility to aphids in endophyte‐free plants but did not affect the insect resistance in endophyte‐symbiotic plants. In line with the unaltered insect resistance, the GA treatment did not reduce the concentration of Epichloë‐derived alkaloids. The Epichloë mycelial biomass was transiently increased by the GA treatment but at the expense of hyphal integrity. The response of the phyllosphere bacterial microbiota to both GA treatment and Epichloë was also evaluated. Only Epichloë, and not the GA treatment, altered the composition of the phyllosphere microbiota and the abundance of certain bacterial taxa. Our findings clearly demonstrate that Epichloë does indeed relieve the plant growth‐defence trade‐off. Summary statement: A debate has been raised if fungal endophytes can relieve the plant growth‐defence trade‐off. Epichloë endophytes decoupled the trade‐off: the growth‐mediated penalisation of plant defences was observed in E− but not in E+ plants. The Epichloë production of alkaloids was likely the decoupling mechanism. [ABSTRACT FROM AUTHOR]

Many temperate grasses are both hyper‐accumulators of silicon (Si) and hosts of Epichloë fungal endophytes, functional traits which may alleviate environmental stresses such as herbivore attack. Si accumulation and endophyte infection may operate synergistically, but this has not been tested in a field setting, nor in the context of changing environmental conditions. Predicted increases in atmospheric CO2 concentrations can affect both Si accumulation and endophyte function, but these have not been studied in combination.We investigated how elevated atmospheric CO2 (eCO2), Si supplementation, endophyte‐presence and insect herbivory impacted plant growth, stoichiometry (C, N, P and Si), leaf gas exchange (rates of photosynthesis, stomatal conductance, transpiration rates) and endophyte production of anti‐herbivore defences (alkaloids) of an important pasture grass (tall fescue; Lolium arundinaceum) in the field.eCO2 and Si supplementation increased shoot biomass (+52% and +31%, respectively), whereas herbivory reduced shoot biomass by at least 35% and induced Si accumulation by 24%. Shoot Si concentrations, in contrast, decreased by 17%–21% under eCO2. Si supplementation and herbivory reduced shoot C concentrations. eCO2 reduced shoot N concentrations which led to increased shoot C:N ratios. Overall, shoot P concentrations were 26% lower in endophytic plants compared to non‐endophytic plants, potentially due to decreased mass flow (i.e. observed reductions in stomatal conductance and transpiration). Alkaloid production was not discernibly affected by any experimental treatment. The negative impacts of endophytes on P uptake were particularly strong under eCO2.We show that eCO2 and insect herbivory reduce and promote Si accumulation, respectively, incorporating some field conditions for the first time. This indicates that these drivers operate in a more realistic ecological context than previously demonstrated. Reduced uptake of P in endophytic plants may adversely affect plant productivity in the future, particularly if increased demand for P due to improved plant growth under eCO2 cannot be met. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Grasses have developed a wide range of morphological and physiological mechanisms to resist herbivory. For instance, they accumulate silicon (Si) in tissue, as physical defence, and associate symbiotically with foliar Epichloë‐endophytes that provide chemical defence via antiherbivore alkaloids. Recent evidence showed that some Epichloë‐endophytes increase foliar Si in forage grasses; however, whether this impacts insect herbivores is unknown. Furthermore, while Si is primarily a physical defence, it also affects production of plant defensive secondary metabolites; Si supply might therefore affect Epichloë‐alkaloids, although this remains untested.We grew endophyte‐free (Nil) and Epichloë‐infected tall fescue and perennial ryegrass in a factorial combination with or without Si supplementation, in the absence or presence of Helicoverpa armigera. Epichloë‐endophyte strains were AR584 for tall fescue, and AR37, AR1 or Wild‐type (WT) for perennial ryegrass. We assessed how Si supply and Epichloë‐endophytes in interaction with herbivory affected foliar Si and mutualist‐derived alkaloid concentrations. Subsequently, their effects on H. armigera relative growth rates (RGRs) were evaluated.Endophytes generally increased Si concentrations in Si‐supplied plants. In tall fescue AR584 and perennial ryegrass AR37, endophytes increased constitutive (herbivore‐free) and induced (herbivore‐inoculated) Si concentrations by at least 25%; in contrast, in perennial ryegrass, the AR1 endophyte only increased constitutive levels. Si supply did not affect alkaloids produced by AR584 or AR1/WT endophytes; however, in the presence of herbivory, Si supply decreased the induction of alkaloids produced by AR37 endophytes by 33%. For tall fescue, Si supply reduced H. armigera RGR by at least 76%, regardless of endophytic status, whereas, endophyte‐alkaloids played a secondary role only reducing herbivore growth in the absence of Si supply. Conversely, in perennial ryegrass, both Si and endophyte‐alkaloids (regardless of Si supply) reduced herbivore RGR although not synergised.Novel interactions between constitutive and induced Si‐ and alkaloid‐based antiherbivore defences in grasses were observed. Overall, Si had a greater effect on the folivore than endophytes in both grasses. Endophyte defences contributed more to herbivore resistance in perennial ryegrass than tall fescue. We demonstrate that Si and endophytes were not antagonistic and highlight that the protective nature of their interaction varies with the grass‐endophyte species tested. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Tropospheric ozone is an abiotic stress of increasing importance in the context of global climate change. This greenhouse gas is a potent phytotoxic molecule with demonstrated negative effects on crop yield and natural ecosystems. Recently, oxidative stress has been proposed as a mechanism that could regulate the interaction between cool‐season grasses and Epichloë endophytes. We hypothesized that exposure of Lolium multiflorum plants, hosting endophytes to an ozone‐polluted environment at different ontogenetic phases, would impact the trans‐generational dynamics of the vertically transmitted fungal symbiont. Here, we found that the ozone‐induced stress on the mother plants did not affect the endophyte vertical transmission but it impaired the persistence of the fungus in the seed exposed to artificial ageing. Endophyte longevity in seed was reduced by exposure of the mother plant to ozone. Although ozone exposure did not influence either the endophyte mycelial concentration or their compound defences (loline alkaloids), a positive correlation was observed between host fitness and the concentration of endophyte‐derived defence compounds. This suggests that fungal defences in grass seeds were not all produced in situ but remobilized from the vegetative tissues. Our study reveals ozone trans‐generational effects on the persistence of a beneficial symbiont in a host grass. Mother plant exposure to ozone did not affect the growth and the transmission of fungal endophytes into host seeds. Seed alkaloid concentration correlated with endophyte mycelial concentration and host fitness. However, ozone‐induced stress on mother plants did impair the longevity of the endophytes in the seeds. [ABSTRACT FROM AUTHOR]

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Abstract: The plant hormone salicylic acid (SA) is recognized as an effective defence against biotrophic pathogens, but its role as regulator of beneficial plant symbionts has received little attention. We studied the relationship between the SA hormone and leaf fungal endophytes on herbivore defences in symbiotic grasses. We hypothesize that the SA exposure suppresses the endophyte reducing the fungal‐produced alkaloids. Because of the role that alkaloids play in anti‐herbivore defences, any reduction in their production should make host plants more susceptible to herbivores. Lolium multiflorum plants symbiotic and nonsymbiotic with the endophyte Epichloë occultans were exposed to SA followed by a challenge with the aphid Rhopalosiphum padi. We measured the level of plant resistance to aphids, and the defences conferred by endophytes and host plants. Symbiotic plants had lower concentrations of SA than did the nonsymbiotic counterparts. Consistent with our prediction, the hormonal treatment reduced the concentration of loline alkaloids (i.e., N‐formyllolines and N‐acetylnorlolines) and consequently decreased the endophyte‐conferred resistance against aphids. Our study highlights the importance of the interaction between the plant immune system and endophytes for the stability of the defensive mutualism. Our results indicate that the SA plays a critical role in regulating the endophyte‐conferred resistance against herbivores. [ABSTRACT FROM AUTHOR]