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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]

Context: Data are lacking on the effects of selected endophytes of perennial ryegrass (Lolium perenne L.) on ryegrass persistence. Aim: We aimed to determine the impact of Epichloë endophyte on the persistence of mid- and late-heading perennial ryegrass cultivars. Method: Two mid-heading (Samson, Bronsyn) and two late-heading (One50, Rohan) cultivars, infected with selected endophytes (AR37, nea2/6) or with standard toxic endophyte, were established in a replicated plot study grazed by cattle in a subtropical environment of the upper North Island of New Zealand. Persistence characteristics were quantified at least five times per annum, over 4 years. Key results: Endophyte strain had an effect on persistence; infection with standard endophyte resulted in higher ryegrass ground cover percentage, ryegrass content in pasture dry matter and autumn yield than infection with nea2/6 on many occasions, and with AR37 on some occasions. There were negligible impacts on ryegrass tiller density or nutritive value. Trends were dominated by the main effect of endophyte; interactions with heading date were inconsistent. Conclusions: Cultivars were more persistent when infected with standard endophyte than with selected endophytes, although persistence declined over 4 years for all cultivars and irrespective of heading date. Implications: Reliance on selected endophyte is unlikely to prevent persistence decline of perennial ryegrass in a subtropical environment. Other strategies will be required to maintain the persistence of high-quality pastures based on perennial ryegrass. We determined the impact of endophyte strain (standard toxic endophyte, selected strains nea2/6 and AR37), on the persistence of four perennial ryegrass cultivars. Standard endophyte resulted in greater persistence than the selected endophytes, with higher ryegrass ground cover percentage and content in pasture dry matter by the end of the study, and higher herbage yields in autumn. Persistence declined over 4 years for all cultivars, regardless of endophyte. [ABSTRACT FROM AUTHOR]

Context: Data are lacking on the effects of heading date of perennial ryegrass (Lolium perenne L.) on ryegrass persistence. Aim: We aimed to determine the impact of heading date on perennial ryegrass persistence. Method: Two mid-heading (Samson, Bronsyn) and two late-heading (One50, Rohan) cultivars, infected with Epichloë endophytes (AR37, nea2/6 or standard toxic), were established in a replicated plot study grazed by cattle in a subtropical environment of the upper North Island of New Zealand. Persistence characteristics were quantified at least five times per annum, over 4 years. Key results: Late-heading cultivars had higher yield, nutritive value, perennial ryegrass content in pasture dry matter, ground cover and tiller density than mid-heading cultivars. There were large seasonal impacts on all ryegrass characteristics, with a major decline over summer and recovery during late autumn into early spring, with less recovery in the final year. Overall, there was a strong linear decline in ryegrass content measured during spring (2018–21). Conclusions: The two late-heading cultivars were more persistent than the two mid-heading cultivars, although persistence declined over 4 years for all cultivars. Implications: Results support industry recommendations of choosing late-heading cultivars for improved pasture quality and persistence. We determined the impact of heading date on persistence of perennial ryegrass cultivars infected with a range of endophytes, in a plot study grazed by cattle. The two late-heading cultivars were more persistent than the two mid-heading cultivars, with higher yield, nutritive value, perennial ryegrass content in pasture dry matter, ground cover and tiller density, although persistence declined over 4 years for all cultivars. [ABSTRACT FROM AUTHOR]

Both adults and larvae of Listronotus bonariensis damage ryegrass (Lolium spp.), a dominant component of New Zealand pasture. Damage can be reduced by Epichloë festucae var. lolii, a biotrophic fungal symbiont of ryegrass, through the production of alkaloids with deterrent and/or toxic effects to insect herbivores including L. bonariensis. Here we report results from two pot trials and a field trial comparing endophyte strains with different alkaloid profiles that have been introduced to reduce or eliminate adverse effects on stock. Adult L. bonariensis feeding scars, number of eggs and the proportion of tillers with larval damage and the severity of that damage were recorded. Samples of pseudostem and leaf blades were analysed for alkaloids. AR1 endophyte reduced adult feeding, oviposition, and larval damage whereas AR37 had no effect on adults but was highly effective in reducing larval damage. Reduction in adult feeding and oviposition by different NEA strains was governed by concentrations of peramine and ergovaline. For plants infected with AR37, alkaloid concentrations that determined the extent of larval damage were influenced by host species and ploidy; tetraploid L. perenne was more severely damaged than its diploid equivalent; L. multiflorum cultivars were more susceptible than the L. hybridum and L. perenne cultivars. [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]

In identifying endophytes for use in pastures in New Zealand there have been two strategies used, either exclusion of ergopeptine and lolitrem alkaloids or, retaining some ergopeptine expression to enhance the stability of natural defences against invertebrate pests. Both have their strengths and weaknesses. It is a matter of balancing these to ensure the end-user has access to a product that satisfies their needs – a persistent pasture with low to nil animal toxicity in terms of animal production and welfare. The range and intensity of ergot alkaloids in grazed pasture on both pasture and ruminants is reviewed, with emphasis on New Zealand. Ergot alkaloids associated with pasture are produced by associations between certain Epichloë endophyte strains and temperate grasses. Ergot alkaloids have been shown to improve persistence of pasture through providing resistance/deterrence to insect pests as well as deterring grazing animals. However, ergovaline is toxic to grazing animals. Some commercially available ryegrass-endophyte associations can produce ergovaline concentrations close to those found in associations between ryegrass and the standard endophyte. It is feasible to eliminate ergot alkaloids from pasture grasses in New Zealand as endophyte strains are commercially available that do not express ergovaline and yet still provide excellent pest resistance. [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]

In a field study during two growing seasons in northern Sweden, we assessed meadow fescue (Festuca pratensis Huds.) with and without Neotyphodium uncinatum to determine whether the endophyte affects plant performance regarding tiller and biomass production. In addition, we measured loline alkaloids produced in infected grasses. Approximately 1000 plants grew at two different levels of nitrogen fertilization: the normal amount applied in this region and one-half that amount. Increasing the amount of nitrogen increased the total number of tillers on the plants. Endophyte-infected plants produced more tillers per plant than plants without the endophyte; there was, however, no significant difference in proportion of reproductive tillers, winter survival, or total biomass production between endophyte-infected and noninfected plants. Infected plants produced loline alkaloids in different quantities without indication of being influenced by different nitrogen fertilization levels. Key words: endophyte, forage, nitrogen fertilization. Nous avons evalue la fetuque des pres (Festuca pratensis Huds.) dans une etude sur le terrain realisee au cours de deux saisons de croissance dans le nord de la Suede, en presence ou en absence de Neotyphodium uncinatum, afin de determiner si 'endophyte affecte la performance de la plante sur le plan de la production de talles et de biomasse. Nous avons de plus mesure es alcaloides loline produits par l'herbe infectee. Pres d'un millier de plantes ont pousse sous deux conditions de fertilisation a 'azote : la quantite normale appliquee dans cette region et la moitie de cette quantite. L'augmentation de la quantite d'azote ccroissait le nombre total de talles sur les plantes. Les plantes infectes par l'endophyte produisaient davantage de talles par lante que les plantes sans endophyte; cette difference n'etait toutefois pas significative en proportion des talles reproductives, e la survie a l'hiver ou de la biomasse totale entre les plantes infectes et non infectes. Les plantes infectes produisaient des lcaloides loline en quantites differentes, sans indication que cela ait ete influence par les differents niveaux de fertilisation a 'azote. [Traduit par la Redaction] Mots-cles: endophyte, forage, fertilisation a l'azote., Introduction Meadow fescue (Festuca pratensis Huds.) is a perennial pooid grass that grows widely in Northern Europe and parts of Asia. In the Nordic and Baltic countries, it is an [...]

Background. Plants possess a sophisticated immune system to defend from herbivores. These defence responses are regulated by plant hormones including salicylic acid (SA) and jasmonic acid (JA). Sometimes, plant defences can be complemented by the presence of symbiotic microorganisms. A remarkable example of this are grasses establishing symbiotic associations with Epichloë fungal endophytes. We studied the level of resistance provided by the grass' defence hormones, and that provided by Epichloë fungal endophytes, against an introduced herbivore aphid. These fungi protect their hosts against herbivores by producing bioactive alkaloids. We hypothesized that either the presence of fungal endophytes or the induction of the plant salicylic acid (SA) defence pathway would enhance the level of resistance of the grass to the aphid. Methods. Lolium multiflorum plants, with and without the fungal endophyte Epichloë occultans, were subjected to an exogenous application of SA followed by a challenge with the aphid, Sipha maydis. Results. Our results indicate that neither the presence of E. occultans nor the induction of the plant's SA pathway regulate S. maydis populations. However, endophytesymbiotic plants may have been more tolerant to the aphid feeding because these plants produced more aboveground biomass. We suggest that this insect insensitivity could be explained by a combination between the ineffectiveness of the specific alkaloids produced by E. occultans in controlling S. maydis aphids and the capacity of this herbivore to deal with hormone-dependent defences of L. multiflorum. [ABSTRACT FROM AUTHOR]