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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]

Aphids are important herbivorous insects that can cause significant crop damage, leading to yield reduction and economic loss. One avenue being explored to reduce aphid impacts is the development of aphid-resistant plants. Under projected climate scenarios, it is expected that plants will be exposed to greater biotic and abiotic stress, including increased herbivorous insect infestation and exposure to prolonged periods of environmental stress, particularly drought. In response to these projections, plant–aphid interactions under drought conditions have been a subject of growing interest; however, few studies have looked at the impact of drought stress on plant resistance to aphids despite the potential importance for plant breeding. Here, we examine the latest scientific advances regarding variation in plant resistance to aphids under drought, emphasizing underlying mechanisms and functional trade-offs and propose a conceptual model relating plant tolerance to drought with plant resistance to aphids. [ABSTRACT FROM AUTHOR]

Here, we propose that the symbiosis outcome in plants associated with endophytes can be compromised, in addition to plant resources, by the action of phytohormones and ROS that are induced by environmental stressors. Endophytes endow plant hosts with mechanisms to protect the mutualism I Epichloë i can prevent the change in the symbiosis outcome in situations of high stress levels by inducing defence-related hormone and antioxidant plant responses that increase plant fitness (e.g. [13]; [18]). Keywords: Antioxidant; endophytes; mutualism; phytohormones; plant-endophyte symbiosis; ROS EN Antioxidant endophytes mutualism phytohormones plant-endophyte symbiosis ROS 19 23 5 12/27/22 20230101 NES 230101 Plants commonly form mutualistic associations with fungal endophytes. Antioxidant, endophytes, mutualism, phytohormones, plant-endophyte symbiosis, ROS. [Extracted from the article]

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]

Plants are expected to face novel challenges as consequence of human-driven global change. Outbreaks of pests and higher incidence of contaminants are increasing. Plants can improve tolerance to stress factors through associations with symbiotic microorganisms. Certain grasses establish persistent and asymptomatic symbioses with Epichloë fungal endophytes, which are known to confer protection against Herbivores and improve plant tolerance to abiotic stress factors. Nonetheless, accumulating evidence suggests the symbiosis outcome is context-dependent. We evaluated the capacity of the endophyte fungus E. occultans in protecting the annual grass Lolium multiflorum against a spontaneous larva attack of the generalist herbivore Agrotis ipsilon under episodic exposure of plants to ozone. Symbiotic and non-symbiotic plants were individually grown outdoors and exposed to ozone at different stages resulting in four treatments: control (plant never exposed to ozone), plant exposed to ozone at vegetative stage, plant exposed to ozone at reproductive stage, and plant exposed to ozone at both stages. After the last exposure, there was an outbreak of A. ipsilon larvae. We evaluated herbivore damage, seed production per plant, and endophyte transmission to the seeds. Frequency of attacked plants was irrespective of both the endophyte and ozone exposure. However, the damage level per plant was only reduced by the endophyte. Seed production was slightly lower in endophyte-symbiotic plants and not affected by ozone. Interestingly, herbivore damaged and undamaged endophyte-symbiotic plants contributed equally to seed production. However, in plants exposed to ozone once at vegetative or reproductive stage, endophyte-free undamaged plants had higher seed production than endophyte-free damaged plants. Ozone treatments did not affect the transmission efficiency of endophytes to the seeds. Mean endophyte transmission efficiency per plant was 95%. Medium doses of ozone seem to have undetectable effects on grass-endophyte symbiosis, not affecting the defensive mutualism nor the persistence of the symbiont across generations. [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]

Background and Aims The processes that maintain variation in the prevalence of symbioses within host populations are not well understood. While the fitness benefits of symbiosis have clearly been shown to drive changes in symbiont prevalence, the rate of transmission has been less well studied. Many grasses host symbiotic fungi (Epichloë spp.), which can be transmitted vertically to seeds or horizontally via spores. These symbionts may protect plants against herbivores by producing alkaloids or by increasing tolerance to damage. Therefore, herbivory may be a key ecological factor that alters symbiont prevalence within host populations by affecting either symbiont benefits to host fitness or the symbiont transmission rate. Here, we addressed the following questions: Does symbiont presence modulate plant tolerance to herbivory? Does folivory increase symbiont vertical transmission to seeds or hyphal density in seedlings? Do plants with symbiont horizontal transmission have lower rates of vertical transmission than plants lacking horizontal transmission? Methods We studied the grass Poa autumnalis and its symbiotic fungi in the genus Epichloë. We measured plant fitness (survival, growth, reproduction) and symbiont transmission to seeds following simulated folivory in a 3-year common garden experiment and surveyed natural populations that varied in mode of symbiont transmission. Key Results Poa autumnalis hosted two Epichloë taxa, an undescribed vertically transmitted Epichloë sp. PauTG-1 and E. typhina subsp. poae with both vertical and horizontal transmission. Simulated folivory reduced plant survival, but endophyte presence increased tolerance to damage and boosted fitness. Folivory increased vertical transmission and hyphal density within seedlings, suggesting induced protection for progeny of damaged plants. Across natural populations, the prevalence of vertical transmission did not correlate with symbiont prevalence or differ with mode of transmission. Conclusions Herbivory not only mediated the reproductive fitness benefits of symbiosis, but also promoted symbiosis prevalence by increasing vertical transmission of the fungus to the next generation. Our results reveal a new mechanism by which herbivores could influence the prevalence of microbial symbionts in host populations. [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]

Tropospheric ozone is considered to be the most phytotoxic air pollutant because of its oxidizing power. The main objective of this study was to analyze the effect of intra- and interspecific competition between Triticum aestivum L. and Trifolium repens L. on the responses to high concentrations of ozone of both species, and the role of the symbiotic relationship Rhizobium - T. repens on the abovementioned responses. Monocultures and mixtures of both species in different densities were sown. Pots were transferred to open top chambers either with 90-120 ppb ozone or without ozone. Ozone had an overall negative impact on leaf area and biomass production per individual plant. These responses were dependent on species and sowing density in monocultures, but were not changed by species proportion in the mixtures. There was a positive relationship between Rhizobium nodules and plant biomass, with a tendency for smaller plants to present lower number of nodules under ozone. These results suggest that competitive and mutualistic interactions could have a greater role in determining responses to novel air pollutants than species sensitivity to the xenobiotic, per se. [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]

There is an increasing interest in determining the influence of global change on plant–microorganism interactions. We review the results of experiments that evaluated the effects of the global change factors carbon dioxide, ozone, temperature, drought, flooding, and salinity on plant symbioses with beneficial Epichloë endophytes. The factors affected the performance of both plants and endophytes as well as the frequency of plants symbiotic with the fungus. Elevated carbon dioxide levels and low temperatures differentially influenced the growth of plants and endophytes, which could compromise the symbioses. Furthermore, we summarise the plant stage in which the effects of the factors were quantified (vegetative, reproductive, or progeny). The factors ozone and drought were studied at all plant stages, but flooding and carbon dioxide were studied in just a few of them. While only studied in response to ozone and drought, evidence showed that the effects of these factors on symbiotic plants persisted trans-generationally. We also identified the putative mechanisms that would explain the effects of the factors on plant–endophyte associations. These mechanisms included the increased contents of reactive oxygen species and defence-related phytohormones, reduced photosynthesis, and altered levels of plant primary metabolites. Finally, we describe the counteracting mechanisms by which endophytes would mitigate the detrimental effects of the factors on plants. In presence of the factors, endophytes increased the contents of antioxidants, reduced the levels of defence-related phytohormones, and enhanced the plant uptake of nutrients and photosynthesis levels. Knowledge gaps regarding the effects of global change on plant–endophyte associations were identified and discussed. [ABSTRACT FROM AUTHOR]

Highlights: [•] Symbiotically modified organisms (SMOs) can improve grass yield, and resistance to pests and weeds. [•] SMOs can improve forage quality for livestock. [•] Toxicity can be caused by uncharacterized plant or fungal metabolites. [•] Feasibility of SMOs depends on the cultivar and fungal strain. [ABSTRACT FROM AUTHOR]

Tropospheric ozone is among the global change factors that pose a threat to plants and microorganisms. Symbiotic microorganisms can assist plants to cope with stress, but their role in the tolerance of plants to ozone is poorly understood. Here, we subjected endophyte-symbiotic and non-symbiotic plants of Lolium multiflorum , an annual species widely distributed in temperate grasslands, to high and low (i.e., charcoal-filtered air) ozone levels at vegetative and reproductive phases. Exposure to high ozone reduced leaf photochemical efficiency and greenness in both symbiotic and non-symbiotic plants. However, ozone-induced oxidative damage at biochemical level (i.e., lipid peroxidation) was mostly detected in symbiotic plants. Ozone exposure at the vegetative phase did not affect the reproductive investment in seeds, indicating full recovery from stress. Ozone exposure at the reproductive phase reduced biomass and seed production only in symbiotic plants indicating a symbiont-associated cost. At low ozone, endophyte-symbiotic plants showed a steeper slope in the relationship between seed number and seed weight (i.e., a number-weight trade-off) compared to non-symbiotic plants. However, when plants were treated at the reproductive phase, ozone increased the imbalance between seed number and seed weight in both endophyte-symbiotic and non-symbiotic plants. Plants with endophytes at the reproductive stage produced fewer seeds, which were not compensated by increased seed weight. Thus, fungal mycelium growing within ovaries or ozone-induced antioxidant systems may result in costs that finally depress the fitness of plants. Despite ozone pollution could destabilize plant-endophyte mutualisms and render them dysfunctional, other endophyte-mediated benefits (e.g., resistance to herbivory, tolerance to drought) could over-compensate these losses and explain the high incidence of the symbiosis in nature. • Symbiotic and non-symbiotic plants were exposed to low and high ozone (O 3). • Plant fitness was more sensitive to O 3 at reproductive than vegetative stages. • Symbiotic plants exposed to O 3 at flowering displayed reduced investment in reproduction. • O 3 caused a seed number−seed weight trade-off of non-symbiotic plants without compromising fitness. • Low frequency of symbiotic plants may be found in populations under O 3 pollution. [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]

Cool-season grasses establish symbioses with vertically transmitted Neotyphodium endophytes widespread in nature. The frequency of endophyte-infected plants in closed populations (i.e., without migrations) depends on both the differential fitness between infected and non-infected plants, and the endophyte-transmission efficiency. Most studies have been focused on the first mechanism ignoring the second. Infection frequency and endophyte transmission from vegetative tissues to seeds were surveyed in two grasses growing in vegetation units that differ in flood and grazing regimes, and soil salinity. Transmission efficiency and infection frequency for tall fescue did not vary significantly and were 0.98 and 1.00, respectively. For Italian ryegrass, transmission efficiency and infection frequency were 0.88 and 0.57 in humid prairies, and 0.96 and 0.96 in the other vegetation units. Only in humid mesophytic meadows, the observed pattern was irrespective of the presence or absence of grazers. Our results suggest that selection forces for endophyte infection are different for both species. Imperfect transmission was only compensated in tall fescue through an increased fitness of infected plants. Interpreting variations of infection frequency only in terms of differential fitness can be misleading, considering that endophyte transmission can be imperfect and variable in nature. Therefore, this study highlights the importance of measuring transmission efficiency. [ABSTRACT FROM AUTHOR]

Neotyphodium endophyte fungi are vertically transmitted symbionts of cool-season grasses. The seed phase of the grass’ life cycle appears to be critical for the persistence of the fungus. Endophyte viability decreases faster than seed viability, but little is known of the effects of this endophyte on seed viability. The endophyte could affect seed viability through changes in water content. Here, we assessed the effects of the endophyte on seed viability, the differential survival of endophyte and seed, and the effects of infection on seed water content. Viability of endophyte-infected and noninfected seeds and endophyte were evaluated over a period of 729 d under 12 controlled environmental conditions. Seed viability was reduced by the infection at high temperature and high relative humidity, but not under other conditions. Moreover, endophyte viability decreased faster than seed viability only under high humidity or high temperature. Seed water content was not affected by endophyte presence. The proportion of viable infected seeds was mainly affected by the loss in endophyte viability and secondly by the differential survival of infected and noninfected seeds. Knowledge on the relative importance of these processes is critical to understand the factors affecting the efficiency of endophyte vertical transmission and the frequency of endophyte-infected plants. [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]

• Parental environment affects functional characteristics of seeds. • Seed-transmitted fungal endophytes mediate transgenerational effects in plants. • Negative effects of transgenerational ozone were detected in endophyte-symbiotic seeds but not in non-symbiotic seeds. • Parental ozone impacted seed progeny when it occurs close to reproductive phase of the plant. • Under accelerated ageing conditions, endophyte-symbiotic seeds showed deteriorated physiological state and reduced longevity. The transition from mother plant seed production to seedlings recruitment is critical for annual plant species but also for vertically transmitted microbial symbionts. Evidence indicates that microbial symbionts play important roles in plant response to environmental changes. However, their role in plant facing gaseous tropospheric pollutants such as the highly reactive and phytotoxic ozone, have been scarcely studied. Here, we experimentally tested the consequences of maternal plant exposure to ozone on the longevity of Lolium mutiflorum seeds mediated by a vertically-transmitted fungal endophyte (Epichloë occultans). Seeds were produced by mother plants exposed to contrasting levels of ozone (low and high concentrations) in two phases of the life cycle (vegetative and reproductive). We first characterized the consequences of plant exposure to ozone on the state of the antioxidant glutathione in the seeds. Then, we examined viability dynamics of the seeds through an accelerated ageing trial and followed the physiological variables seed water content (SWC) and electrical conductivity (EC). Irrespectively of maternal ozone, seeds produced by endophyte-symbiotic plants presented higher contents of the oxidised form of glutathione. In the same direction, the fungal endophyte presence was associated with a dramatic reduction in seed longevity. The SWC was higher in endophyte-symbiotic seeds that in endophyte-free seeds. As time under accelerated ageing condition advanced, EC increased more in endophyte-symbiotic seeds than in endophyte-free seeds. Mother plant exposure to ozone tended to have a negative synergistic effect with the endophyte symbiosis. Ozone transgenerational effect was not detected on endophyte-free seeds. Our work contributes to understand the transgenerational effects of vertically transmitted fungal endophytes in plants in response to a currently important factor of stress. [ABSTRACT FROM AUTHOR]

Background and Aims In the subfamily Poöideae (Poaceae), certain grass species possess anti-herbivore alkaloids synthesized by fungal endophytes that belong to the genus Epichloë (Clavicipitaceae). The protective role of these symbiotic endophytes can vary, depending on alkaloid concentrations within specific plant–endophyte associations and plant parts. Methods We conducted a literature review to identify articles containing alkaloid concentration data for various plant parts in six important pasture species, Lolium arundinaceum , Lolium perenne , Lolium pratense , Lolium multiflorum | Lolium rigidum and Festuca rubra , associated with their common endophytes. We considered the alkaloids lolines (1-aminopyrrolizidines), peramine (pyrrolopyrazines), ergovaline (ergot alkaloids) and lolitrem B (indole-diterpenes). While all these alkaloids have shown bioactivity against insect herbivores, ergovaline and lolitrem B are harmful for mammals. Key Results Loline alkaloid levels were higher in the perennial grasses L. pratense and L. arundinaceum compared to the annual species L. multiflorum and L. rigidum , and higher in reproductive tissues than in vegetative structures. This is probably due to the greater biomass accumulation in perennial species that can result in higher endophyte mycelial biomass. Peramine concentrations were higher in L. perenne than in L. arundinaceum and not affected by plant part. This can be attributed to the high within-plant mobility of peramine. Ergovaline and lolitrem B, both hydrophobic compounds, were associated with plant parts where fungal mycelium is usually present, and their concentrations were higher in plant reproductive tissues. Only loline alkaloid data were sufficient for below-ground tissue analyses and concentrations were lower than in above-ground parts. Conclusions Our study provides a comprehensive synthesis of fungal alkaloid variation across host grasses and plant parts, essential for understanding the endophyte-conferred defence extent. The patterns can be understood by considering endophyte growth within the plant and alkaloid mobility. Our study identifies research gaps, including the limited documentation of alkaloid presence in roots and the need to investigate the influence of different environmental conditions. [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]

Information on whole community responses is needed to predict direction and magnitude of changes in plant and animal abundance under global changes. This study quantifies the effect of past ozone exposure on a weed community structure and arthropod colonization. We used the soil seed bank resulting from a long-term ozone exposure to reestablish the plant community under a new low-pollution environment. Two separate experiments using the same original soil seed bank were conducted. Plant and arthropod richness and species abundance was assessed during two years. We predicted that exposure to episodic high concentrations of ozone during a series of growing cycles would result in plant assemblies with lower diversity (lower species richness and higher dominance), due to an increase in dominance of the stress tolerant species and the elimination of the ozone-sensitive species. As a consequence, arthropod-plant interactions would also be changed. Species richness of the recruited plant communities from different exposure histories was similar (≈ 15). However, the relative abundance of the dominant species varied according to history of exposure, with two annual species dominating ozone enriched plots (90 ppb: Spergula arvensis, and 120 ppb: Calandrinia ciliata). Being consistent both years, the proportion of carnivore species was significantly higher in plots with history of higher ozone concentration (≈3.4 and ≈7.7 fold higher in 90 ppb and 120 ppb plots, respectively). Our study provides evidence that, past history of pollution might be as relevant as management practices in structuring agroecosystems, since we show that an increase in tropospheric ozone may influence biotic communities even years after the exposure. [ABSTRACT FROM AUTHOR]

Tropospheric ozone is one of the major drivers of global change. This stress factor alters plant growth and development. Ozone could act as a selection pressure on species communities composition, but also on population genetic background, thus affecting life history traits. Our objective was to evaluate the consequences of prolonged ozone exposure of a weed community on phenotypic traits of Spergula arvensis linked to persistence. Specifically, we predicted that the selection pressure exerted by high ozone concentrations as well as the concomitant changes in the weed community would drive population adaptive changes which will be reflected on seed germination, dormancy and longevity. In order to test seed viability and dormancy level, we conducted germination experiments for which we used seeds produced by S. arvensis plants grown within a weed community exposed to three ozone treatments during four years (0, 90 and 120 ppb). We also performed a soil seed bank experiment to test seed longevity with seeds coming from both the four-year ozone exposure experiment and from a short-term treatment conducted at ambient and added ozone concentrations. We found that prolonged ozone exposure produced changes in seed germination, dormancy and longevity, resulting in three S. arvensis populations. Seeds from the 90 ppb ozone selection treatment had the highest level of germination when stored at 75% RH and 25 °C and then scarified. These seeds showed the lowest dormancy level when being subjected to 5 ºC/5% RH and 25 ºC/75% followed by 5% RH storage conditions. Furthermore, ozone exposure increased seed persistence in the soil through a maternal effect. Given that tropospheric ozone is an important pollutant in rural areas, changes in seed traits due to ozone exposure could increase weed persistence in fields, thus affecting weed-crop interactions, which could ultimately reduce crop production. [ABSTRACT FROM AUTHOR]

Invasive plants may displace native species. This is the case for Poa annua, the only non-native plant species successfully established in Maritime Antarctica. Nonetheless, it is uncertain which factors drive the competitive success of P. annua in the harsh environmental conditions of the region. The ability of this plant species to establish novel mutualistic interactions with resident soil fungi may be crucial for its invasiveness. Such ability may be linked to the vertical transmission of fungal endophytes via seeds. We undertook a study to assess the role of seed fungal endophytes as promoters of the establishment and invasion of Poa annua in Maritime Antarctica. We explored the composition and diversity of fungal communities associated with different P. annua tissues (seeds, leaves and roots) and the soil. We also measured parameters including germination rate, above-ground biomass, reproductive structures, and the survival of invasive P. annua as well as of the native Colobanthus quitensis and Deschampsia antarctica grown from seeds with and without endophytes. Furthermore, we conducted inter- and intraspecific competition experiments among native and invasive plants, where chemically mediated plant-to-plant interference (allelopathy) and plant growth rate were measured to calculate a relative competition index. We found that fungal endophyte taxa associated with P. annua tissues were very different from those in the soil. Fungal endophytes in P. annua differed among seed, root and shoot tissues, which suggests low transmission among different organs. The removal of endophytes from P. annua seeds was associated with reduced seed germination, plant growth and survivorship, while the competitive ability of P. annua (assessed by accumulated biomass) relative to native species, as well as levels of allelochemicals in soils, were higher in the presence of seed fungal endophytes. Our results suggest that fungal endophytes, maternally inherited through seeds, improve host fitness and may contribute to the invasive success of P. annua in Antarctica. [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]

Plants have evolved mechanisms to survive herbivory. One such mechanism is the induction of defences upon attack that can operate intergenerationally. Cool-season grasses (sub-family Pooideae) obtain defences via symbiosis with vertically transmitted fungal endophytes (genus Epichloë) and can also show inducible responses. However, it is unknown whether these herbivore-induced responses can have intergenerational effects. We hypothesized that herbivory by aphids on maternal plants induces the intergenerational accumulation of endophyte-derived defensive alkaloids and resistance intensification in the progeny. We subjected mother plants symbiotic or not with Epichloë occultans, a species known for its production of anti-insect alkaloids known as lolines, to the aphid Rhopalosiphum padi. Then, we evaluated the progeny of these plants in terms of loline alkaloid concentration, resistance level (through herbivore performance), and shoot biomass. Herbivory on mother plants did not increase the concentration of lolines in seeds but it tended to affect loline concentration in progeny plants. There was an overall herbivore-induced intergenerational effect increasing the endophyte-conferred defence and resistance. Symbiotic plants were more resistant to aphids and had higher shoot biomass than their non-symbiotic counterparts. Since maternal herbivory did not affect the loline concentrations in seeds, the greater resistance of the progeny could have resulted from an inherited mechanism of epigenetic regulation. It would be interesting to elucidate the origin of this regulation since it could come from the host or the fungal symbiont. Thus, endophyte-driven differential fitness between symbiotic and non-symbiotic plants might be higher as generations pass on in presence of herbivores. [ABSTRACT FROM AUTHOR]

Context. Tall fescue (Schedonorus arundinaceus) is one of the most important cultivated forage grasses in temperate regions but its association with some Epichloë fungal endophytes usually makes it unsuitable for livestock feeding due to toxic alkaloids. However, re-inoculation of plants with non-toxic endophytes can result in positive effects for livestock feeding. Aims. Assess the effects of the non-toxic AR584 endophyte on seed viability and germination in two tall fescue cultivars. Methods. The effects of AR584 endophyte (presence/absence) and tall fescue cultivar (INIA Aurora/Taita) were evaluated on seed viability and germination responses across a range of temperature and water availability regimes. Response to constant temperature and water availability was characterised by the thermal-time and hydro-time models, respectively. Key results. Under accelerated aging conditions, endophyte presence reduced seed viability in cultivar Taita. For both cultivars, endophyte-infected seeds showed a lower germination rate than endophyte-free seeds, except for INIA Aurora under low incubation temperatures. Also, endophyte-infected seeds in both cultivars displayed lower germination rates and final germination as the water potential was lowered. Conclusions. A strong cultivar-by-endophyte interaction for seed germination indicates the need to understand how specific cultivarendophyte combinations best fit a given environmental condition, as defined by temperature and water availability. Implications. The information we present can be valuable not only for farmers who have to decide what to cultivate but also for seed companies that have to preserve seed quality. [ABSTRACT FROM AUTHOR]

Harsh environmental conditions derived from current climate change trends are among the main challenges for agricultural production worldwide. In the Mediterranean climatic region of central Chile, sudden occurrence of spring cold temperatures in combination with water shortage for irrigation (drought) constitutes a major limitation to highbush blueberry (Vaccinium corymbosum) plantations, as flowering and fruiting stages are highly sensitive. Hardening crops may be achievable by boosting beneficial interactions of plants with microorganisms. Inoculation with symbiotic fungi isolated from plants adapted to extreme environments could be a good strategy, if they are able to maintain functional roles with non-original hosts. Here, we evaluated the effect of two Antarctic fungal endophytes (AFE), Penicillium rubens and P. bialowienzense, on the tolerance of V. corymbosum plants to cold events in combination with drought under controlled conditions. Inoculated and uninoculated plants were exposed for a month to one event of a cold temperature (2 °C/8 h) per week with or without drought and were evaluated in physiological, biochemical, and molecular variables. A complementary set of plants was kept under the same environmental conditions for two additional months to evaluate survival as well as fruit weight and size. There was an overall positive effect of AFE on plant performance in both environmental conditions. Endophyte-inoculated plants exhibited higher gene expression of the Late Embryogenesis Abundant protein (LEA1), higher photochemical efficiency (Fv/Fm), and low oxidative stress (TBARS) than uninoculated counterparts. On the other hand, plant survival was positively affected by the presence of fungal endophytes. Similarly, fruit diameter and fruit fresh weight were improved by fungal inoculation, being this difference higher under well-watered condition. Inoculating plants with fungal endophytes isolated from extreme environments represents a promising alternative for hardening crops. This is especially relevant nowadays since agriculture is confronting great environmental uncertainties and difficulties which could became worse in the near future due to climate change. [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]

Epichloë endophytes are common symbionts living asymptomatically in pooid grasses and may provide chemical defences against herbivorous insects. While the mechanisms underlying these fungal defences have been well studied, it remains unknown whether endophyte presence affects the host's own defences. We addressed this issue by examining variation in the impact of Epichloë on constitutive and herbivore-induced emissions of volatile organic compounds (VOC), a well-known indirect plant defence, between two grass species, Schedonorus phoenix (ex. Festuca arundinacea; tall fescue) and Festuca pratensis (meadow fescue). We found that feeding by a generalist aphid species, Rhopalosiphum padi, induced VOC emissions by uninfected plants of both grass species but to varying extents, while mechanical wounding failed to do so in both species after one day of damage. Interestingly, regardless of damage treatment, Epichloë uncinata-infected F. pratensis emitted significantly lower quantities of VOCs than their uninfected counterparts. In contrast, Epichloë coenophiala-infected S. phoenix did not differ from their uninfected counterparts in constitutive VOC emissions but tended to increase VOC emissions under intense aphid feeding. A multivariate analysis showed that endophyte status imposed stronger differences in VOC profiles of F. pratensis than damage treatment, while the reverse was true for S. phoenix. Additionally, both endophytes inhibited R. padi population growth as measured by aphid dry biomass, with the inhibition appearing greater in E. uncinata-infected F. pratensis. Our results suggest, not only that Epichloë endophytes may play important roles in mediating host VOC responses to herbivory, but also that the magnitude and direction of such responses may vary with the identity of the Epichloë–grass symbiosis. Whether Epichloë-mediated host VOC responses will eventually translate into effects on higher trophic levels merits future investigation. [ABSTRACT FROM AUTHOR]

Expansins are proteins involved in cell wall metabolism that play an important role in plant growth, development, fruit ripening and abiotic stress tolerance. In the present study, we analyzed putative expansins that respond to drought stress. Five expansin genes were identified in cDNA libraries isolated from Colobanthus quitensis gown either with or without endophytic fungi under hydric stress. A differential transcript abundance was observed by qPCR analysis upon drought stress. To compare these expansin genes, and to suggest a possible mechanism of action at the molecular level, the structural model of the deduced proteins was obtained by comparative modeling methodology. The structures showed two domains and an open groove on the surface of the proteins was observed in the five structural models. The proteins were evaluated in terms of their protein-ligand interactions using four different ligands. The results suggested differences in their mode of protein-ligand interaction, in particular concerning the residues involved in the protein-ligand interaction. The presented evidence supports the participation of some members of the expansin multiprotein family in the response to drought stress in C. quitensis and suggest that the response is modulated by endophytic fungi. [Display omitted] • Antarctic plant lives in stressing environment in symbiosis with fungal endophytes. • Expansin encoding genes are differentially expressed in response to drought stress. • Expansin are differentially modulated by fungal endophytes presence. • Expansin proteins display variable substrate affinity to polysaccharides. [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]

Climatic change is pointed as one of the major challenges for global food security. Based on current models of climate change, reduction in precipitations and in turn, increase in the soil salinity will be a sharp constraint for crops productivity worldwide. In this context, root fungi appear as a new strategy to improve plant ecophysiological performance and crop yield under abiotic stress. In this study, we evaluated the impact of the two fungal endophytes Penicillium brevicompactum and P. chrysogenum isolated from Antarctic plants on nutrients and Na+ contents, net photosynthesis, water use efficiency, yield and survival in tomato and lettuce, facing salinity stress conditions. Inoculation of plant roots with fungal endophytes resulted in greater fresh and dry biomass production, and an enhanced survival rate under salt conditions. Inoculation of plants with the fungal endophytes was related with a higher up/down-regulation of ion homeostasis by enhanced expression of the NHX1 gene. The two endophytes diminished the effects of salt stress in tomato and lettuce, provoked a higher efficiency in photosynthetic energy production and an improved sequestration of Na+ in vacuoles is suggested by the upregulating of the expression of vacuolar NHX1 Na+/H+ antiporters. Promoting plant-beneficial interactions with root symbionts appears to be an environmentally friendly strategy to mitigate the impact of climate change variables on crop production. [ABSTRACT FROM AUTHOR]

Tall fescue is usually infected by a fungal endophyte, responsible of livestock intoxication due to fungal alkaloids. An endophyte fungus grows in the apoplast of aboveground tissues and is vertically transmitted through the seeds. Disinfecting plants or clones not only is useful as a way of eliminating a dangerous microorganism from cultivars, but also for experimental purposes. However, not all fungicides are effective, and if they are, a waiting period must be respected in order to avoid confounding effects of the endophyte and the treatment. We subjected plants (ramets from four genotypes) to different doses of two fungicides, Almagor® (Triazole + Imidazole) and Amistar® (Methoxy-acrylate), and evaluated the endophyte persistence. Fungicides were pipette-poured on pseudostems' bases of each plant. Endophytic status was diagnosed in tillers that received the fungicide and in the new tillers produced by the plants and seeds. While Amistar® had no detectable effect, Almagor® was 100% effective at all doses. The latter interrupted the verticaltransmission processes (5400 evaluated seeds were endophyte-free). Phytotoxic effects were not observed in seeds. Proportion of normal and abnormal seedlings depended only on plant genotype. Almagor® was effective in stopping the fungus from growing into reproductive buds when apical meristems were down on the bases. Use of Almagor® is promising on old pastures dominated by toxic tall fescue, with the purpose of enriching the soil seed-bank with endophyte-free seeds. [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]

Although the role of fungal alkaloids in protecting grasses associated with Epichloë fungal endophytes has been extensively documented, the effects of the symbiont on the host plant’s immune responses have received little attention. We propose that, in addition to producing protective alkaloids, endophytes enhance plant immunity against chewing insects by promoting endogenous defense responses mediated by the jasmonic acid (JA) pathway. We advance a model that integrates this dual effect of endophytes on plant defenses and test its predictions by means of a standard meta-analysis. This analysis supports a role of Epichloë endophytes in boosting JA-mediated plant defenses. We discuss the ecological scenarios where this effect of endophytes on plant defenses would be most beneficial for increasing plant fitness. [ABSTRACT FROM AUTHOR]

Polyploidy and symbiotic Epichloë fungal endophytes are common and heritable characteristics that can facilitate environmental range expansion in grasses. Here we examined geographic patterns of polyploidy and the frequency of fungal endophyte colonized plants in 29 Festuca rubra L. populations from eight geographic sites across latitudes from Spain to northernmost Finland and Greenland. Ploidy seemed to be positively and negatively correlated with latitude and productivity, respectively. However, the correlations were nonlinear; 84% of the plants were hexaploids (2n = 6x = 42), and the positive correlation between ploidy level and latitude is the result of only four populations skewing the data. In the southernmost end of the gradient 86% of the plants were tetraploids (2n = 4x = 28), whereas in the northernmost end of the gradient one population had only octoploid plants (2n = 8x = 56). Endophytes were detected in 22 out of the 29 populations. Endophyte frequencies varied among geographic sites, and populations and habitats within geographic sites irrespective of ploidy, latitude or productivity. The highest overall endophyte frequencies were found in the southernmost end of the gradient, Spain, where 69% of plants harbored endophytes. In northern Finland, endophytes were detected in 30% of grasses but endophyte frequencies varied among populations from 0% to 75%, being higher in meadows compared to riverbanks. The endophytes were detected in 36%, 30% and 27% of the plants in Faroe Islands, Iceland and Switzerland, respectively. Practically all examined plants collected from southern Finland and Greenland were endophyte-free, whereas in other geographic sites endophyte frequencies were highly variable among populations. Common to all populations with high endophyte frequencies is heavy vertebrate grazing. We propose that the detected endophyte frequencies and ploidy levels mirror past distribution history of F. rubra after the last glaciation period, and local adaptations to past or prevailing selection forces such as vertebrate grazing. [ABSTRACT FROM AUTHOR]