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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. [ABSTRACT FROM AUTHOR]

Crucial to our understanding of plant ecology is the consideration of the eco‐physiological responses and constraints of plant–fungal symbioses throughout the native distribution range of their host.We examined key eco‐physiological roles of two co‐occurring fungal symbionts [Epichloë endophytes and arbuscular mycorrhizal fungi (AMF)] in the endemic grass Hordeum comosum across a wide bioclimatic gradient and contrasting grazing severity. We sampled H. comosum plants along four humid‐to‐arid transects in Patagonia, Argentina, covering its entire distribution range and determined Epichloë presence, AMF root colonization, nitrogen and phosphorus concentration, intrinsic water‐use efficiency (iWUE, the ratio of photosynthesis to stomatal conductance) and 18O‐enrichment of cellulose in shoots.Root colonization by AMF increased with Epichloë presence. All plants hosted Epichloë in the humid range of the gradient, but symbioses occurrence decreased towards arid sites which also displayed severe grazing symptoms at site level.Symbiosis with Epichloë correlated positively with shoot nitrogen concentration in the centre of the distribution range, and with shoot phosphorus concentration across the entire distribution range.The site‐level relationship of AMF colonization with 18O‐enrichment and iWUE suggested that mycorrhiza boosted stomatal conductance in humid environments but curbed it in arid environments.While the interpretation of interactions and potential causalities from observational studies should be done with caution, this study demonstrates distinct correlations between plant–fungal symbiont associations and key resource parameters (phosphorus, nitrogen and iWUE vs. 18O‐enrichment). Such correlations may suggest particular functional roles for these symbionts in the ecology of their host plant. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Bromus auleticus is a winter wild species with promissory characteristics for domestication as forage that inhabits temperate‐warm grasslands from a wide region from South America. Populations from regions with different environmental conditions are considered as ecotypes and are associated with different seedborne Epichloë fungal endophytes. Association with Epichloë provides B. auleticus with increased growth capacity and resistance against pathogens. The seeds mature at the end of spring and germinate in autumn, remaining dormant during wet and warm summer. As the persistence of Epichloë endophytes in the seed is highly sensitive to temperature and humidity, we studied how the endophytes modulate seed germination in response to these factors in two B. auleticus ecotypes named La Pampa (LP) and El Palmar (EP). We also studied how environmental factors during germination affect endophyte survival. In the LP ecotype, adapted to coldest temperatures in autumn, Epichloë improved final germination under alternating temperatures, but had no effects on germination inhibition imposed by low water availability. In the EP ecotype, adapted to warm humid summer and autumn, the endophyte promoted germination at constant 25°C and alternating 15/25°C temperatures, and prevented germination at low water potentials. At 30°C, seed germination was inhibited and whereas seeds remained viable, endophyte survival was significantly lowered. The effect of Epichloë on B. auleticus seed germination was ecotype‐specific: promoting germination at alternating temperatures in La Pampa ecotype or at high temperatures in El Palmar ecotype, while inhibiting it at low water availabilities, could be an endophyte‐mediated ecotypic strategy minimising endophyte mortality and increasing seedling establishment. [ABSTRACT FROM AUTHOR]

Ground‐level ozone is a global air pollutant with high toxicity and represents a threat to plants and microorganisms. Although beneficial microorganisms can improve host performance, their role in connecting environmentally induced maternal plant phenotypes to progeny (transgenerational effects [TGE]) is unknown. We evaluated fungal endophyte‐mediated consequences of maternal plant exposure to ozone on performance of the progeny under contrasting scenarios of the same factor (high and low) at two stages: seedling and young plant. With no variation in biomass, maternal ozone‐induced oxidative damage in the progeny that was lower in endophyte‐symbiotic plants. This correlated with an endophyte‐mediated higher concentration of proline, a defence compound associated with stress control. Interestingly, ozone‐induced TGE was not associated with reductions in plant survival. On the contrary, there was an overall positive effect on seedling survival in the presence of endophytes. The positive effect of maternal ozone increasing young plant survival was irrespective of symbiosis and only expressed under high ozone condition. Our study shows that hereditary microorganisms can modulate the capacity of plants to transgenerationally adjust progeny phenotype to atmospheric change. Ozone induced positive transgenerational effects (TGE) by increasing plant survival. It depended on fungal endophytes symbiosis, progeny size, and current ozone condition. Endophytes were determinant for seedlings, while current ozone allowed TGE to express in young plants. Our results give insights on the role of seed‐borne fungi into environmentally induced transgenerational responses of plants. [ABSTRACT FROM AUTHOR]

Our hypothesis is that I Epichloë i endophytes may uncouple the growth-defence trade-off by simultaneously inducing plant growth-related hormones and producing defence compounds (alkaloids) that circumvent the need for host defence-related hormones (Fig. As a final note, it is important to remark that we put forward that endophytes I can i eliminate the growth-defence trade-off, proving our point with I Epichloë i endophytes but acknowledging that not all endophytes would possess the ability to decouple the trade-off. Keywords: alkaloids; endophytes; growth-defence trade-off; phytohormones; symbiosis EN alkaloids endophytes growth-defence trade-off phytohormones symbiosis 388 390 3 06/20/22 20220715 NES 220715 Acknowledgements We thank Víctor M. Escobedo and Christina D. Moon for critical contributions to earlier versions of the manuscript. How and when fungal endophytes can eliminate the plant growth-defence trade-off: mechanistic perspectives. [Extracted from the article]

Tall fescue (Festuca arundinacea Schreb.) is a perennial C3 grass species usually associated with the endophyte fungus Epichloë coenophiala that enhances tolerance to biotic and abiotic stresses. Given that the proportion of infected tall fescue plants in the grasslands of the Flooding Pampa (Argentina) has been progressively increasing, it is proposed that endophytes confer tolerance to the stresses characteristic of that environment, especially flooding. Plants from a naturalized population and a commercial cultivar were grown, both with and without endophyte (wild type and AR584 respectively). The plants were subjected to two submergence treatments (i.e., control and partial submergence) in two development stages (5‐leaf stage and beginning of the elongation of internodes). Plant performance (aerial and root biomass, root aerenchyma formation, total root length and root diameter, proportion of reproductive tillers, number of panicles and seeds produced) and endophyte transmission to progeny were evaluated. Endophytes did not alter the ecological fitness of plants under partial submergence. Independently of the water condition, the symbiosis was linked to low biomass in plants from the naturalized population, while the opposite occurred in plants from the cultivar. Partial submergence did not affect the germination of the seeds produced, nor the transmission of the endophyte to the seedlings that originated from them. Our work suggests that the invasion ability of tall fescue in the Flooding Pampa grasslands is not related to an endophyte‐mediated improvement of plants to tolerate water excess. [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]

Natural grassland ecosystems are increasingly threatened by excessive loadings of nutrients and by the presence of species bred for high productivity. By manipulating grazing regimes and nutrient availability, agricultural practices facilitate the establishment and spread of certain forage plant species outside managed landscapes, challenging local biodiversity. The ecological success of some species in the invaded range sometimes seems to be associated with the symbiosis with foliar fungal endophytes. Symbiotic fungi may increase the competitiveness of host species, but also the resistance to herbivory through the production of toxic secondary compounds such as alkaloids. While progress has been made in understanding how soil nutrients modulate other benefits offered by fungal endophytes to plants (e.g. stress tolerance, competitive ability, etc), the consequences for a higher trophic level (i.e. herbivores) and the potential feedbacks on plant invasion have not been explored yet.We explored the relative and interactive importance of soil nitrogen (N) and phosphorus (P) in modulating the interaction of the invasive grass tall fescue—associated with fungal endophytes—and native herbivores in a natural grassland. We hypothesized that N and P nutrients modulate differentially leaf quality traits, namely nutritional value and fungal alkaloid contents, determining the level of damage by native insect herbivores on the exotic tall fescue.We found that only the addition of P significantly increased native caterpillar density in the field, which corresponded to a concomitant increase in leaf damage. In contrast to expectations, the concentration of the alkaloid ergovaline in leaves was not strongly related to N. It was the level of soil P which dictated the concentration of the element (P) in the leaves and reduced the level of defence against herbivores in this endophyte‐symbiotic species. Then, herbivore performance increased, and plants were more prone to be attacked.Synthesis. Our study indicates a strong control of soil P fertility on the triple interaction among plants, fungal endophytes and native herbivores. This highlights the potential role of increased soil nutrients on the invasion spread of endophyte‐symbiotic forage plants in natural grasslands. [ABSTRACT FROM AUTHOR]

We reviewed and performed a quantitative synthesis on herbicide‐resistant weeds from rain‐fed crops in Argentina. Twenty‐four weed species distributed in the main extensive crops (soybean, maize, wheat, barley, oilseed rape, sunflower, chickpea and peanut) have evolved herbicide resistance. Of the total, 54% are grasses, 88% are annual species and 63% are cross‐pollinated species. The most representative families were Poaceae with 54% resistant species, followed by Brassicaceae with 17%, and Asteraceae with 13%. Buenos Aires, Santa Fe and Córdoba were the provinces with the most documented cases of resistance (35%, 33% and 30%, respectively). The proportion of cases resistant to pre‐emergence herbicides was 10%, whereas the proportion of cases resistant to post‐emergence herbicides was 90%. Glyphosate was the herbicide with the highest incidence (92%) of resistance among weed species, followed by 29% of species that evolved resistance to ALS‐inhibiting herbicides. Whereas resistance to auxin‐like herbicides comprised 17% of the weed species, acetyl‐CoA carboxylase (8%) and protoporphyrinogen oxidase (4%) inhibiting herbicides showed the least incidence of resistance evolution among weeds. The highest number of resistant species was identified in soybean (19), followed by maize (13), wheat/barley (10) and fallow (9). Weed species with a higher number of resistant populations to a higher number of herbicide mode of action were Amaranthus hybridus, A. palmeri, Lolium multiflorum and Raphanus sativus. The change in the production system since the mid‐1990s, based on the use of herbicides (glyphosate mainly) to control weeds, is likely to account for the notorious increase in the average rate of evolution of herbicide‐resistant weeds in Argentina. [ABSTRACT FROM AUTHOR]

PREMISE OF THE STUDY:: A transgenerational effect occurs when a biotic or abiotic environmental factor acts on a parental individual and thereby affects the phenotype of progeny. Due to the importance of transgenerational effects for understanding plant ecology and evolution, their underlying mechanisms are of general interest. Here, we introduce the concept that inherited symbiotic microorganisms could act as mechanisms of transgenerational effects in plants. METHODS:: We define the criteria required to demonstrate that transgenerational effects are microbially mediated and review evidence from the well‐studied, vertically transmitted plant–fungal symbiosis (grass–Epichloë spp.) in support of such effects. We also propose a basic experimental design to test for the presence of adaptive transgenerational effects mediated by plant symbionts. KEY RESULTS:: An increasingly large body of literature shows that vertically transmitted microorganisms are common in plants, with potential to affect the phenotypes and fitness of progeny. Transgenerational effects could occur via parental modification of symbiont presence/absence, symbiont load, symbiont products, symbiont genotype or species composition, or symbiont priming. Several of these mechanisms appear likely in the grass–Epichloë endophytic symbiosis, as there is variation in the proportion of the progeny that carries the fungus, as well as variation in concentrations of mycelia and secondary compounds (alkaloids and osmolytes) in the seed. CONCLUSIONS:: Symbiont‐mediated transgenerational effects could be common in plants and could play large roles in plant adaptation to changing environments, but definitive tests are needed. We hope our contribution will spark new lines of research on the transgenerational effects of vertically transmitted symbionts in plants. [ABSTRACT FROM AUTHOR]

Ozone is an increasing tropospheric contaminant of climate change. Exposure to ozone may affect the symbiotic relationship between plants and beneficial microorganisms. In particular, the herbivore resistance mechanism conferred by fungal endophytes (defensive mutualism) may be affected, as any of the ozone-triggered effects (such as elicitation of defence mechanisms against biotrophic fungi or oxidative stress in the apoplastic space) may target the symbiont., Symbiotic and non-symbiotic Lolium multiflorum plants were exposed to ozone for two consecutive days (2 h per day), after which half were infested with 10 aphids ( Rhopalosiphum padi). We measured variables related to performance of plants, the endophyte symbiont (alkaloids) and the herbivores., Aphid populations were smaller on symbiotic plants than in non-symbiotic plants in low-ozone conditions. However, this difference disappeared in exposed plants to high-ozone conditions. Under low-ozone conditions, structure of aphid populations on endophyte-symbiotic plants was characterized by a low number of nymphs and a high number of adults. This pattern was not observed with high-ozone exposure. Level of fungal alkaloids (lolines) was not affected by either ozone or herbivory., Results indicate that ozone impairs the grass-endophyte symbiosis without affecting production of alkaloids generally linked with aphids' response to endophyte presence. Since neither plant biomass nor alkaloid level were affected by ozone, other ozone-mediated mechanisms at molecular or biochemical level may underlie plant-herbivore interaction mediated by fungal endophytes. Thus, the mechanism behind this effect must be determined in future experiments. [ABSTRACT FROM AUTHOR]

Certain species of the Pooideae subfamily develop stress tolerance and herbivory resistance through symbiosis with vertically transmitted, asexual fungi. This symbiosis is specific, and genetic factors modulate the compatibility between partners. Although gene flow is clearly a fitness trait in allogamous grasses, because it injects hybrid vigor and raw material for evolution, it could reduce compatibility and thus mutualism effectiveness. To explore the importance of host genetic background in modulating the performance of symbiosis, Lolium multiflorum plants, infected and noninfected with Neotyphodium occultans, were crossed with genetically distant plants of isolines (susceptible and resistant to diclofop-methyl herbicide) bred from two cultivars and exposed to stress. The endophyte improved seedling survival in genotypes susceptible to herbicide, while it had a negative effect on one of the genetically resistant crosses. Mutualism provided resistance to herbivory independently of the host genotype, but this effect vanished under stress. While no endophyte effect was observed on host reproductive success, it was increased by interpopulation plant crosses. Neither gene flow nor herbicide had an important impact on endophyte transmission. Host fitness improvements attributable to gene flow do not appear to result in direct conflict with mutualism while this seems to be an important mechanism for the ecological and contemporary evolution of the symbiotum. [ABSTRACT FROM AUTHOR]

Neotyphodium endophytic fungi, the asexual state of Epichloë species, protect cool-season grasses against stresses. The outcomes of Neotyphodium-grass symbioses are agronomically relevant as they may affect the productivity of pastures. It has been suggested that the mutualism is characteristic of agronomic grasses and that differential rates of gene flow between both partners' populations are expected to disrupt the specificity of the association and, thus, the mutualism in wild grasses. We propose that compatibility is necessary but not sufficient to explain the outcomes of Neotyphodium-grass symbiosis, and advance a model that links genetic compatibility, mutualism effectiveness, and endophyte transmission efficiency. For endophytes that reproduce clonally and depend on allogamous hosts for reproduction and dissemination, we propose that this symbiosis works as an integrated entity where gene flow promotes its fitness and evolution. Compatibility between the host plant and the fungal endophyte would be high in genetically close parents; however, mutualism effectiveness and transmission efficiency would be low in fitness depressed host plants. Increasing the genetic distance of mating parents would increase mutualism effectiveness and transmission efficiency. This tendency would be broken when the genetic distance between parents is high (out-breeding depression). Our model allows for testable hypotheses that would contribute to understand the coevolutionary origin and future of the endophyte-grass mutualism. [ABSTRACT FROM AUTHOR]

The authors comment on the study about the relationship between the pattern of Neotyphodium spp. endophyte infection frequency and the gradient of grazing pressure by pikas and hoary marmots in Canada. They propose that the effect of herbivores on infection frequency might be through phenotypic traits and interaction with other environmental factors. They also show the relation of relative fitness to herbivory deterrence.

Neotyphodium frequently occurs as an endophyte in grasses. Evidence shows enhanced fitness of endophyte infected grasses relative to non-infected ones. Some studies of seed germination show endophyte enhancement of plant fitness in various environments, but inconsistent results indicate that further studies are needed. So far, experiments have failed to separate the confounded effects of population origin and seed management. For this reason, we evaluated the effects of endophyte infection on seed dormancy and germination in Lolium multiflorum using an experimental design controlling these factors. Depending on the year of seed production, endophyte infection modified seed response to light quality, affecting predominantly seed dormancy levels. Nevertheless, the endophyte did not affect base temperature or thermal time of germination. We concluded that endophytes were not a strong influence on germination behaviour. We speculate from our results that the presence of the endophyte changes germination by an indirect effect, in extending growth of the maternal plant during seed development and ripening. The direct effect of hyphae in the seed on seed behaviour was disregarded, because the difference between infected and non-infected seed varied within the year of seed production. Future experiments should focus on effects of the endophyte on the canopy of parent plants during seed production and ripening, and, hence, on subsequent dormancy and germination of the seeds. [ABSTRACT FROM AUTHOR]

Lolium multiflorumis a successful invader of postagricultural succession in the Inland Pampa grasslands in Argentina, becoming a dominant species in the plant community. Individual plants of this annual species are naturally highly infected with fungal endophytes (Neotyphodiumsp.) from early successional stages. We assessed the effect ofNeotyphodiuminfection on the biology ofL. multiflorum. We evaluated growth attributes between endophyte infected (E+) and uninfected (E–) plants under non-competitive conditions during the normal growing season. E+ plants produced significantly more vegetative tillers and allocated more biomass to roots and seeds. Although seed germination rates were greater in endophyte free plants, the rate of emergence and the final proportion of emerged seedlings were similar between the biotypes. The greater production of vegetative tillers, and the greater resource allocation to roots and seeds are likely to confer an ecological advantage to E+ plants, thus enabling their dominance over the E– individuals in natural grasslands. [ABSTRACT FROM AUTHOR]

Summary: A trade‐off between growth and defence functions is commonly observed in plants. We propose that the association of plants with Epichloë fungal endophytes may eliminate this trade‐off. This would be a consequence of the double role of these endophytes in host plants: the stimulation of plant growth hormones (e.g. gibberellins) and the fungal production of antiherbivore alkaloids. We put forward a model that integrates this dual effect of endophytes on plant growth and defence and test its predictions by means of meta‐analysis of published literature. Our results support the notion that the enhanced plant resistance promoted by endophytes does not compromise plant growth. The limits and ecological benefits of this endophyte‐mediated lack of plant growth–defence trade‐off are discussed. [ABSTRACT FROM AUTHOR]

In this review, we examine the functional roles of microbial symbionts in plant tolerance to cold and freezing stresses. The impacts of symbionts on antioxidant activity, hormonal signaling and host osmotic balance are described, including the effects of the bacterial endosymbionts Burkholderia, Pseudomonas and Azospirillum on photosynthesis and the accumulation of carbohydrates such as trehalose and raffinose that improve cell osmotic regulation and plasma membrane integrity. The influence of root fungal endophytes and arbuscular mycorrhizal fungi on plant physiology at low temperatures, for example their effects on nutrient acquisition and the accumulation of indole‐3‐acetic acid and antioxidants in tissues, are also reviewed. Meta‐analyses are presented showing that aspects of plant performance (shoot biomass, relative water content, sugar and proline concentrations and Fv/Fm) are enhanced in symbiotic plants at low (−1 to 15 °C), but not at high (20–26 °C), temperatures. We discuss the implications of microbial symbionts for plant performance at low and sub‐zero temperatures in the natural environment and propose future directions for research into the effects of symbionts on the cold and freezing tolerances of plants, concluding that further studies should routinely incorporate symbiotic microbes in their experimental designs. [ABSTRACT FROM AUTHOR]

Plants inhabiting environments with stressful conditions often exhibit a low number of flowers, which can be attributed to the energetic cost associated with reproduction. One of the most stressful environments for plants is the Antarctic continent, characterized by limited soil water availability and low temperatures. Induction of dehydrins like those from the COR gene family and auxin transcriptional response repressor genes (IAAs), which are involved in floral repression, has been described in response to water stress. Here, we investigated the relationship between the water deficit‐induced stress response and the number of flowers in Colobanthus quitensis plants collected from populations along a latitudinal gradient. The expression levels of COR47 and IAA12 genes in response to water deficit were found to be associated with the number of flowers. The relationship was observed both in the field and growth chambers. Watering the plants in the growth chambers alleviated the stress and stimualted flowering, thereby eliminating the trade‐off observed in the field. Our study provides a mechanistic understanding of the ecological constraints on plant reproduction along a water availability gradient. However, further experiments are needed to elucidate the primary role of water availability in regulating resource allocation to reproduction in plants inhibiting extreme environments. [ABSTRACT FROM AUTHOR]

Environmental stresses can compromise the interactions of plants with beneficial microbes. In the present review, experimental results showing that stresses negatively affect the abundance and/or functionality of plant beneficial microbes are summarized. It is proposed that the environmental interference of these plant−microbe interactions is explained by the stress‐mediated induction of plant signalling pathways associated with defence hormones and reactive oxygen species. These plant responses are recognized to regulate beneficial microbes within plants. The direct negative effect of stresses on microbes may also contribute to the environmental regulation of these plant mutualisms. It is also posited that, in stress situations, beneficial microbes harbour mechanisms that contribute to maintain the mutualistic associations. Beneficial microbes produce effector proteins and increase the antioxidant levels in plants that counteract the detrimental effects of plant stress responses on them. In addition, they deliver specific stress‐protective mechanisms that assist to their plant hosts to mitigate the negative effects of stresses. Our study contributes to understanding how environmental stresses affect plant−microbe interactions and highlights why beneficial microbes can still deliver benefits to plants in stressful environments. SUMMARY STATEMENT: Here, it is proposed that the environmental interference of plant‐microbe interactions is explained by the stress‐mediated induction of certain plant signalling pathways and that specific mechanisms harboured by beneficial microbes assist in maintaining the mutualisms in stress situations. [ABSTRACT FROM AUTHOR]

1. The strength of many interactions between plants and other organisms changes across regional gradients. For example, the relevance of plant-herbivore interactions increases with primary production. Likewise, biotic interactions collectively become more intense from the poles to the equator. Yet, the regional variation of the interaction between grasses and systemic fungal endophytes, which provide resistance to biotic and abiotic environmental factors (i.e. herbivory and drought), is poorly understood. 2. We compiled 1008 records of the incidence level of fungal endophytes (Epichloë, Ascomycetes: Clavicipitaceae) on wild populations of 48 cool season grasses, encompassing 10 biomes across a broad latitudinal expanse and primary production gradient. Symbiosis incidence was analysed as a function of mean primary production, precipitation, temperature and latitude of each site, which in turn were obtained from climatic and satellital sources. 3. Across a 30-fold variation of mean primary production, average symbiosis incidence increased from 20% to 70%. The pattern became stronger when the analysis was restricted to the single grass genus Festuca, which accounted for half of the total data. 4. The number of grass populations showing no symbiosis incidence (0%) decreased as primary production increased, whereas those with 100% of incidence increased. 5. Primary production at the regional scale was negatively correlated with latitude but positively with mean annual temperature and precipitation. Symbiosis incidence was similarly correlated with latitude and temperature, and it was not with mean annual precipitation. 6. Synthesis. Different descriptors of this grass-fungus symbiosis show that average incidence in wild populations world-wide increases with mean primary production. As at large spatial scales herbivory and temperature increase and aridity decreases with primary production, our results suggest that, at broad-scales, these biotic and abiotic factors may be important drivers of the symbiosis success. [ABSTRACT FROM AUTHOR]

Symbiosis between cool-season grasses and vertically transmitted fungal endophytes are common and significantly impact on ecosystem function. This makes the understanding of the underlying mechanisms to symbiotic individuals frequency in local populations much more interesting. Most studies have been focused on the differential fitness between symbiotic and non-symbiotic counterparts (relative fitness), barely considering other mechanisms. We performed a microcosms experiment to evaluate whether grazing alters the dynamics of the endophyte Neotyphodium occultans in the annual grass Lolium multiflorum by simultaneously modifying the relative fitness and the endophyte efficiency to be transmitted from host plants to seeds. Grazing was simulated by means of clipping and trampling on symbiotic and non-symbiotic plants growing separately, in soils obtained from paddocks, differing in their agronomic management history (natural grassland vs. ryegrass promotion). Seed production showed a complex pattern as it depended on the symbiotic status of the plants, the level of grazing and the agro-ecological context. Grazed plants produced three times fewer seeds than ungrazed plants only in microcosms with endophyte-symbiotic plants in soils from ryegrass promotion. Endophyte benefits on seed production were exclusively observed in ungrazed plants in the same soil. Symbiotic plants produced symbiotic and non-symbiotic seeds in all the treatments. While the production of non-symbiotic seeds by these plants was not affected by grazing and the soil, grazing reduced the production of symbiotic seeds in both contexts. Grazing negative effect on the density of fully infected spikes determined a significant increment in the transmission failures which were not modified by agro-ecological contexts. Therefore, grazing can modulate symbiosis dynamics through reducing seed production and endophyte transmission efficiency. Transmission has been disregarded, but it is a context-dependent process that could lead to a gradual reduction in the symbiotic plants frequency in a population if the mutualism effectiveness does not outweigh transmission failures. [ABSTRACT FROM AUTHOR]

The most studied mechanism of protection against herbivores in grasses associated with Epichloë fungal endophytes has been the fungal production of alkaloids. However, the contribution of the plant immune response on the level of resistance to herbivores in symbiotic grasses has been poorly explored. We studied the relationship between the plant hormone, jasmonic acid (JA) and Epichloë fungal endophytes on herbivore defences in symbiotic grasses. We hypothesized that an exogenous application of methyl jasmonate (MeJA), an activator of the plant JA defence response, would increase the level of resistance of endophyte‐symbiotic and non‐symbiotic plants to a chewing insect. As Epichloë endophytes produce alkaloids, an enhancement of the JA defence would complement the resistance given by these alkaloids.Lolium multiflorum plants symbiotic and non‐symbiotic with the endophyte Epichloë occultans were subjected to an exogenous application of MeJA followed by a challenge with the generalist chewing insect Spodoptera frugiperda. We measured the level of plant resistance to chewing insects, and the defences conferred by host plants and fungal endophytes.Symbiotic plants were more resistant to S. frugiperda than their non‐symbiotic counterparts. However, despite the fact that the concentration of JA significantly increased in all plants exposure to MeJA, neither endophyte‐symbiotic nor non‐symbiotic plants showed an enhanced resistance to insects. Unexpectedly, the exposure of endophyte‐symbiotic plants to MeJA led to a reduction in the concentration of loline alkaloids (i.e. N‐formyllolines and N‐acetylnorlolines), consequently decreasing the level of plant resistance to the herbivore.Synthesis. Our results suggest that, rather than complementing the alkaloid‐based defence, the jasmonic acid hormone weakens the anti‐herbivore mechanism conferred by Epichloë endophytes. The present study highlights that the interaction between the jasmonic acid hormone and the presence of leaf fungal endophytes can be of importance for the effectiveness of the anti‐herbivore defences of symbiotic plants. Our results suggest that, rather than complementing the alkaloid‐based defence, the jasmonic acid hormone weakens the anti‐herbivore mechanism conferred by Epichloë endophytes. The present study highlights that the interaction between the jasmonic acid hormone and the presence of leaf fungal endophytes can be of importance for the effectiveness of the anti‐herbivore defences of symbiotic plants. [ABSTRACT FROM AUTHOR]

Abstract: Temperate grasses frequently acquire resistance to herbivores through a symbiosis with epichloid fungi that produces alkaloids of variable deterrent effects. However, in those cases without apparent endophyte negative effects on domestic herbivores, it is not clear whether plant consumption or preference is affected or not. We performed three experiments with 1‐year‐old steers (Bos taurus, Aberdeen Angus) and the annual grass Lolium multiflorum, infected or not by Epichloë occultans to evaluate preference and to identify the underlying tolerance mechanisms. The first experiment evaluated steer preference for L. multiflorum cultivated in plots with three endophyte infection frequencies (low, medium and high), and investigated the role of canopy structure and plant nutritional traits on preference. The second experiment evaluated preference for chopped grass, offered in individual trays with contrasting infection frequencies (low and high), to discard possible effects associated with canopy structure and to focus on nutritional traits. The third experiment was performed with a tray + basket design that separated visual and olfactory stimuli from nutritional traits. High endophyte infection frequencies reduced consistently animal preference, even after short (~10 min) feeding events. However, we did not find significant evidence of plant structural, nutritional, visual or olfactory traits. Our results discarded several potential mechanisms; therefore, the dissuasive effect of fungal endophytes on animal consumption might be related to other mechanisms, including, likely, alkaloids and changes on grass metabolome. [ABSTRACT FROM AUTHOR]

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]