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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]

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]

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. Key words: endophyte infection, seed longevity, endophyte longevity, endophyte transmission. Les champignons endophytes Neotyphodium sont des symbiotes des herbacees de saisons froides qui se transmettent verticalement. La phase seminale du cycle vital des herbacees semble critique pour la persistance de ce champignon. La viabilite de l'endophyte diminue plus rapidement que la viabilite de la semence, mais on connat peu de chose des effets de cet endophyte sur la viabilite des graines. L'endophyte pourrait affecter la viabilite des graines par des modifications de la teneur en eau. Les auteurs evaluent les effets de l'endophyte sur la viabilite des graines, sur la survie differentielle de l'endophyte et de la graine, ainsi que les effets de la colonisation sur la teneur en eau de la graine. Ils ont evalue la viabilite de graines colonisees ou non colonisees par l'endophyte, ainsi que celle de l'endophyte, sur une periode de 729 jours sous 12 conditions environnementales controlees. La temperature et l'humidite relative elevees reduisent la viabilite de la graine, mais pas les autres conditions. De plus, la viabilite de l'endophyte diminue plus rapidement que celle de la graine, seulement sous des temperatures et des humidites elevees. La presence de l'endophyte n'affecte pas la teneur en eau des graines. La proportion des graines viables colonisees est surtout affectee par la perte de viabilite de l'endophyte et secondairement par la survie differentielle des graines colonisees et non colonisees. On doit connatre l'importance relative de ces processus pour comprendre les facteurs qui affectent l'efficacite de la transmission verticale de l'endophyte et la frequence des plantes colonisees par l'endophyte. Mots-cles : colonisation endophytique, longevite des graines, longevite de l'endophyte, transmission de l'endophyte. [Traduit par la Redaction], Introduction Many cool-season grasses are symbiotically associated with nonpathogenic systemic and endophytic fungi of the genus Neotyphodium (Clay and Schardl 2002; Faeth and Sullivan 2003). These microorganisms are solely vertically [...]

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]

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]

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]

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

BACKGROUND: The effect of plant size (seedlings versus young plants versus adult plants) on the phenotypic level of glyphosate resistance and selection intensity (SI) in Sorghum halepense with and without a reduced glyphosate translocation resistance mechanism was evaluated. RESULTS: Resistance parameters [the 50% lethal dose (LD50) and the dose required to cause a 50% reduction in plant growth (GR50)] in adult plants were notably higher than in seedlings regardless of the resistance status. However, under similar plant size increases, populations comprised of glyphosate‐resistant (R) individuals showed higher survival and growth when glyphosate treated compared with glyphosate‐susceptible (S) plants. An increase in SI was always evident with increasing glyphosate doses. However, the rate of increase in SI was higher under glyphosate selection of young R and S plants, followed by seedlings and adult R and S plants. However, in conditions of R seedlings coexisting with adult S plants under glyphosate treatment (1000–4000 g ha−1), selection against glyphosate resistance was observed. CONCLUSION: Any increase in size from the seedling stage of R plants translates into an amplification of resistance. Depending on the particular size combinations of spatially coexisting R and S plants, selection for glyphosate resistance may be faster, slower or even not evident. © 2018 Society of Chemical Industry Depending on the particular size combination of spatially coexisting glyphosate‐resistant (R) and susceptible (S) plants, the intensity of glyphosate selection may increase or decrease under increasing glyphosate doses. [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]