Leaf endophyte Neotyphodium coenophialum modifies mineral uptake in tall fescue

Neotyphodium coenophialum (Morgan-Jones and Gams) Glenn, Bacon and Hanlin, a fungal endophyte found primarily in shoots of tall fescue (Festuca arundinacea Shreb.), can modify rhizosphere activity in response to phosphorus (P) deficiency. In a controlled environment experiment, two cloned tall fescue genotypes (DN2 and DN4) free (E-) and infected (E+) with their naturally occurring endophyte strains were grown in nutrient solutions at low P (3.1 ppm) or high P (31 ppm) concentrations for 21 d. Endophyte infection increased root dry matter (DM) of DN4 by 21% but did not affect root DM of DN2. Under P deficiency, shoot and total DM were not affected by endophyte but relative growth rate was greater in E+ than E- plants. In high P nutrient solution, E+ plants produced 13% less (DN2) or 29% more (DN4) shoot DM than E- plants. Endophyte affected mineral concentrations in roots more than in shoots. Regardless of P concentration in nutrient solution, E+ DN2 accumulated more P, Ca, Zn and Cu but less K in roots than E- plants. When grown in high P nutrient solution, concentrations of Fe and B in roots of E+ DN2 plants were reduced compared with those of E- plants. Concentrations of P, Ca and Cu in roots of DN4 were less, but K was greater in E+ than E- plants. In shoots, E+ DN2 had greater concentrations of Fe and Cu than E- DN2, regardless of P concentration in nutrient solution. Genotype DN4 responded to endophyte infection by reducing B concentration in shoots. Nutrient uptake rates were affected by endophyte infection in plants grown in low P nutrient solution. A greater uptake rate of most nutrients and their transport to shoots was observed in DN2, but responses of DN4 were not consistent. Results suggest that endophyte may elicit different modes of tall fescue adaptation to P deficiency. This revised version was published online in June 2006 with corrections to the Cover Date.     

Endophytic fungi (Neotyphodium coenophialum) affect the growth and mineral uptake, transport and efficiency ratios in tall fescue (Festuca arundinacea)

Neotyphodium coenophialuminteracts mutualistically with its host grasses. Tall fescue (Festuca arundinacea Schreb.) plants infected by the fungal endophyte,Neotyphodium coenophialum(Morgan-Jones and Gams) Glenn, Bacon and Hanlin, often perform better than non-infected plants, especially in limited resource environments. However, there is a scarcity of information about endophyte-grass ecotypes interaction in Andisols of temperate regions. Clones of three tall fescue ecotypes (Fukaura, Koiwai and Showa) either infected with N. coenophialum (E+) or noninfected (E–) were grown in Andisols (Black Andisol: naturally low content of phosphorus, high in other nutrients; Red Andisol: naturally high content of phosphorus, low in other nutrients) for 133 days in a controlled environment. Cumulative shoot dry weight, daily regrowth rates (tiller number, plant height and shoot dry matter) after clippings and nutrient uptake, transport and efficiency ratios were measured. In Black Andisol, E+ plants had significantly higher cumulative shoot dry weight as well as daily regrowth rates than E– plants, while in Red Andisol the reverse was true. Among the ecotypes studied, Showa had the highest shoot growth. Significantly higher phosphorus (P), potassium (K), calcium (Ca) and magnesium (Mg) uptake as well as transport were identified in E+ vs. E– plants grown in Black Andisol. With few exceptions, values for nutrient efficiency ratios were not significantly different between E+ and E– plants grown in both soils. Significant three-way interaction (endophyte × ecotype × soil) for cumulative shoot dry weight and regrowth rate revealed that the ecotype specific regrowth responses to endophyte infection were depended on soil nutrient conditions. Vegetative growth and nutrient acquisition in tall fescue varied with ecotype and were modified by abiotic (soil fertility status) as well as biotic (endophyte infection) factors.     

How does the Fungal Endophyte Neotyphodium coenophialum Affect Tall Fescue (Festuca arundinacea) Rhizodeposition and Soil Microorganisms?

The goal of our study was to investigate the impact of fungal endophytes in tall fescue (Festuca arundinacea) on rhizodeposition and in turn, the soil microbial community. Sand-based, aseptic microlysimeter units were constructed for the collection of rhizodeposit solutions for chemical analyses from the roots of endophyte-free (E−) and endophyte-infected (E+) tall fescue plants. E+ plants were infected with Neotyphodium coenophialum, the most common endophyte found in tall fescue. Rhizodeposit solutions collected over nine weeks from E+ grass contained more organic carbon and carbohydrates than E−. These solutions were allowed to percolate through columns of plant-free soils to assess the response of the soil microbial communities. Soils to which solutions from E+ grass were applied had significantly higher respiration rates than those receiving solutions from E− grass, suggesting that microbial activity was stimulated by changes in the rhizodeposits. Culture-based assays of the soil microbial community (plate counts and community-level physiological profiling) suggest that the basic structure of the microbial community was not affected by application of rhizodeposit solutions from E+ plants as compared to E−. Our results indicate that the presence of a fungal endophyte may enhance rhizodeposition by tall fescue and could consequently influence microbial mineralization processes in the soil. In grasslands where nutrients may be limiting, hosting a fungal endophyte has the potential to enhance plant nutrient supply indirectly via a stimulatory effect on the soil microbial biomass. Megan M. Van Hecke and Amy M. Treonis - Both authors contributed equally to this work.     

Does Endophyte Influence Regrowth of Tall Fescue?

The influence of ecological and environmental factors on theresponse of the mutualistic symbionts Acremonium coenophialum(Morgan-Jones and Gams), and tall fescue ( Festuca arundinacea,Schreb.) has received substantial attention. However, much ofwhat is known about developmental and physiological responseshas been obtained from static or mature canopies and the influenceof defoliation on the symbiota has not been considered in detail.A controlled environment experiment was conducted to assessthe influence of defoliation and endophyte on the regrowth anddevelopment of two tall fescue genotypes and their respectiveendophytes, known to differ in morphology and alkaloid productioncapacity. The response of endophyte infected relative to non-infectedplants was evaluated for measured and calculated parametersfor each defoliation. While the influence of genotype was substantialon virtually all parameters, endophyte interacted with genotypeand defoliation resulting in enhanced plasticity of mutualistsas a response to varying conditions. Endophyte infection influencedleaf mass depending upon genotype, while the relative benefitof endophyte on pseudostem mass was affected by defoliation.Endophyte interacted with genotype to influence relative growthrate and productivity relative to nitrogen concentration whencanopies were clipped, but was less important when canopiesremained uncut. In some instances endophyte gave growth andsize advantage to the host and did not in others. Generalitiesregarding symbiont response to extrinsic factors may be misleading,since responses depend upon the specific mutualism consideredand conditions imposed. Acremonium coenophialum ; Festuca arundinacea ; non-structural carbohydrate; relative growth rate     

Influence of Endophyte and Water Regime Upon Tall Fescue Accessions. II. Pyrrolizidine and Ergopeptine Alkaloids

Alkaloids, along with specific environmental conditions, havebeen associated with both detrimental and beneficial aspectsof endophyte (Acremonium coenophialum Morgan-Jones et Gams)infected tall fescue (Festuca arundinacea Schreb.) associations.Benefits to the plant accrue through reduced herbivory, whereasdetriment to the animal occurs as altered grazing behaviourand reduced productivity. A controlled environment study wasconducted to examine pyrrolizidine and ergopeptine alkaloidconcentration of four tall fescue accessions as influenced byendophyte status and water regime. Endophyte-free plants weredevoid of ergopeptine alkaloid and contained little, if any,pyrrolizidine alkaloid. Leaf blade tissue of endophyte-infectedisolines contained a range of both ergopeptine (256 to 1633ng g^–1) and pyrrolizidine (92 to 450 µg g^–1)alkaloid concentrations. Water deficit generally increased alkaloidconcentration. Alkaloid yield, based upon concentration andtissue d. wt, showed that significant increase in ergopeptineand pyrrolizidine alkaloid in leaf tissue was associated withwater deficit and was due to actual increased synthesis andnot simply decreased phytomass. Leaf and pseudostem (leaf sheathand stem base) tissue alkaloid concentrations indicated differentaccumulation patterns for ergopeptine and pyrrolizidine alkaloids.Ergopeptine alkaloid yield increased in water-stressed pseudostem,whereas pyrrolizidine alkaloid yield decreased in some, butnot all accessions. The range of host genotype/endophyte biotyperesponse offers the possibility to select associations whichproduce few deleterious effects in animals yet maintain highforage productivity and persistence. Festuca arundinacea, Acremonium coenophialum, tall fescue genotypes, water stress, N-formyl and N-acetyl loline, ergovaline     

Defoliation and Leaf Age Influence on Ergot Alkaloids in Tall Fescue

A controlled environment experiment was conducted to determinethe influence of defoliation on the regrowth and developmentof two tall fescue (Festuca arundinacea Schreb.) host-endophyte(Neotyphodium coenophialum Morgan-Jones and Gams (Glenn, Bacon,Price and Hanlincomb. nov. ) associations (DN2 and DN11), knownto differ in morphology and alkaloid production capacity. Defoliationtreatments included an uncut control, and clipping to a 5- or10-cm residue height. In a separate experiment, leaf age effectswere determined on an uncut plant canopy. Ergot alkaloid concentrationwas greatest in pseudostem and least in harvested leaf. Non-infectedplants were devoid of alkaloid. Alkaloid production (expressedas a function of dry matter) and yield were greater in uncutthan clipped plants, and were greater in DN11 than DN2 plants.Alkaloid production increased with increasing N concentrationin both associations; however, the rate of production was influencedby concentration of non-structural carbohydrate. Leaf age influencedleaf mass and alkaloid concentration and yield. Oldest leaves(>6 weeks after appearance) of DN2 and DN11 had the lowestconcentrations of ergot alkaloid, while leaves that were 2 to4 weeks old had the greatest alkaloid concentration and yield(alkaloid concentrationxleaf mass). Repeated defoliation reducedthe production of alkaloids, but did so as a function of non-structuralcarbohydrate. Our data suggest that alkaloid production canbe modified by defoliation as well as by host-endophyte association. Neotyphodium coenophialum ; Festuca arundinacea ; N; non-structural carbohydrate     

Temporal and spatial patterns of soil water following wildfire-induced changes in plant communities in the Great Basin in Nevada,USA

Infection of tall fescue (Festuca arundinacea Schreb.) with its endemicNeotyphodium coenophialum-endophyte (Morgan-Jones and Gams) Glenn, Bacon and Hanlin appears to reduce copper (Cu) concentrations in forage and serum of grazing animals, contributing to a range of immune-related disorders. A greenhouse experiment was conducted to identify effects of novel endophyte strains on Cu acquisition by tall fescue (Festuca arundinacea Schreb.) varieties Grasslands Flecha and Jesup infected with a novel, non ergot producing endophyte strain AR542, and two perennial ryegrass (Lolium perenne L.) varieties Aries and Quartet infected with a novel, non lolitrem B producing strain AR1, and their noninfected (E−) forms. Individual endophyte/grass associations were cultivated in nutrient solutions at 1.0 (P+) and 0.0 mM (P−) phosphorus concentrations. The Cu²⁺-binding activity of extracellular root exudates, and concentrations of Cu and other heavy metals in roots and shoots were measured. Extracellular root exudates of AR542-infected vs. E− tall fescue had higher Cu²⁺-binding activity only in P− nutrient solution as shown by lower concentration of free Cu²⁺ (0.096 vs. 0.188 mmol Cu²⁺ g⁻¹ root DM, respectively). The Cu²⁺-binding activity by root exudates of perennial ryegrass was not affected by endophyte infection, but was higher (i.e., lower concentration of free Cu²⁺) in P− vs. P+ nutrient solution (0.068 vs. 0.114 mmol Cu²⁺ g⁻¹ root DM). In this hydroponic experiment, Cu concentrations in shoots of both grasses were not a function of Cu²⁺-binding activity and endophyte effects on heavy metal concentrations in shoots and roots were specific for each variety. The Cu²⁺-binding activity of extracellular root exudates may affect Cu accumulation by field-grown, endophyte-infected tall fescue under P-limiting growth conditions and warrants verification by more specific methods.     

Soil organic carbon fractions differ in two contrasting tall fescue systems

The value of tall fescue (Festuca arundinacea Schreb.) for C sequestration in addition to forage production and soil conservation is of current interest. However, studies relating to the impacts of endophyte infected (E+) and endophyte free (E?) tall fescue on soil organic matter fractions are few. This study examined how E+ and E? growth affected soil C fractions 4 years after establishment. The study site was at the University of Kentucky research farm, Lexington, Kentucky, USA. From soil cores in replicated fields we measured total C, particulate organic matter C (POM C), microbial biomass C (MBC), mineralizable C (Min C), C in aggregates, and aggregate distribution at depths of 0 to 15 and 15 to 30 cm. Significant effects between E+ and E- fescue were sometimes observed for MBC, Min C, C in micro-aggregates, and aggregate distribution, but not for total C, and POM C. At 0?C15 cm MBC (E+ 26% greater than E?), Min C (E+ 43% lower than E?), C associated with micro-aggregates (E+ 15% lower than E?), and micro-aggregates (46% more micro-aggregates in E+ than E?), were affected by endophyte infection, confirming hypotheses that early changes in soil properties were reflected in labile C fractions and soil structure. Endophyte infection status in tall fescue has quantifiable effects on C sequestration and soil structure, achievable in a relatively short period that can be used to monitor conservation efforts and the consequences of pasture renovation strategies.     

Fungal endophyte presence and genotype affect plant diversity and soil-to-atmosphere trace gas fluxes

Aims

Novel fungal endophyte (Neotyphodium coenophialum; Latch, Christensen and Samuels; Glenn, Bacon, and Hanlin) genotypes in symbiosis with tall fescue (Lolium arundinaceum; Schreb. Darbysh.) have been recently introduced to agricultural seed markets. These novel endophytes do not produce the full suite of toxins that the ‘common toxic’ form does, and therefore, may not have the same consequences on plant and soil processes. Here, we evaluated the effects of endophyte presence and genotype on ecosystem processes of tall fescue stands.

Methods

We quantified the effects of the presence of the common toxic endophyte (CT), two novel endophyte genotypes (AR-542, AR-584), no endophyte (endophyte free, E-), and a mixture of all endophyte statuses (mix) within a single genotype of tall fescue (PDF) on various soil and plant parameters.

Results

Endophyte presence and genotype affected tall fescue cover and plant species diversity: cover—CT, AR-542, AR -584, mix > E- and species diversity—E- > AR-542, AR -584 > CT, mix. Most measured soil parameters had significant endophyte effects. For example, higher fluxes of soil CO₂ and N₂O were measured from stands of AR-542 than from the other endophyte treatments.

Conclusions

These results indicate that endophyte presence and genetic identity are important in understanding the ecosystem-scale effects of this agronomically important grass-fungal symbiosis.     

Short-term Responses of Soil C and N Fractions to Tall Fescue Endophyte Infection

Tall fescue (Festuca arundinacea Schreb.) is naturally infected with a fungal endophyte, Neotyphodium coenophialum, which produces toxic ergot alkaloids that negatively affect herbivores and that may alter soil organic matter dynamics. A 60-week mesocosm study with a factorial arrangement of soil type (clay loam and loamy sand) and endophyte infection (with and without) was conducted to determine potential changes in soil C and N fractions. Forage and root dry-matter production were greater with than without endophyte infection, while forage C and N concentrations were unaffected. Total, particulate, mineralizable, and aggregate-associated C and N fractions increased several fold during the course of the experiment due to large rhizosphere inputs in all treatments. The fraction of total C and N in water-stable macroaggregates (>0.25 mm) was initially 0.43 ± 0.10 and 0.46 ± 0.16, respectively, and increased during the course of the experiment to 0.68 ± 0.06 and 0.56 ± 0.15 when averaged across soil type and endophyte infection level as a result of organic matter cycling and deposition in this active biophysical fraction. Changes in soil C and N fractions due to endophyte infection were minimal. The lack of detectable changes in soil C and N fractions due to endophyte infection may have been a result of the overwhelming input of C from roots and/or the relatively short-term nature (60 weeks). Greater plant productivity of endophyte-infected tall fescue is likely a contributing mechanism for eventual changes in total and active C and N fractions that have been observed in long-term pastures.     

Susceptibility of tall fescue to Rhizoctonia zeae infection as affected by endophyte symbiosis

Endophytic fungi belonging to the genus Neotyphodium often form symbiotic associations with grasses. The host plants usually benefit from the association with an endophyte. Presence of the symbiont may increase host resistance to infection by some pathogens. However, the exact mechanism of the lower susceptibility of endophyte‐infected plants to diseases is still unclear. Growth chamber trials were conducted to determine whether (a) tall fescue plants infected with the endophyte Neotyphodium coenophialum (E+) are more resistant to sheath and leaf spot disease caused by Rhizoctonia zeae than endophyte‐free (E?) plants, and (b) R. zeae growth inhibition is associated with endophyte presence. Tall fescue genotypes, each symbiotic with a genetically different native endophyte strain, were inoculated with isolates of R. zeae. The tillers infection by R. zeae, density of endophyte hyphae and content of total phenolic compounds in tillers were studied. Antifungal activity of the N. coenophialum towards R. zeae, Rhizoctonia solani, Bipolaris sorokiniana and Curvularia lunata was also investigated in dual‐culture assays. For Tf3, Tf4, TfA2 and TfA9 tall fescue genotypes, the E+ plants had reduced R. zeae infection. In the Tf9 and Tf8085 genotypes, R. zeae infection was similar for both E+ and E? plants. The strongest effect was observed for the Tf4 endophyte. A strongly positive correlation (r = 0.94) occurred between endophyte hyphal density and disease index across all tall fescue genotypes. Dual‐culture assays showed no inhibitory interaction between the seven endophyte strains and the R. zeae isolates; however, some endophytes inhibited R. solani, B. sorokiniana and C. lunata. Endophyte presence increased the production of phenolic compounds by the host grasses. The level of phenolics also differed significantly depending on the time of analysis after inoculation of plants by R. zeae. The results indicate that N. coenophialum can suppress disease severity caused by R. zeae infection. The mechanism of higher resistance of E+ plants is likely not based on direct inhibition such as antibiosis or competition. Thus, the induction of specific mechanisms in the host plant, for example, production of phenolic compounds, seems to be the main way of providing resistance to the grass by the endophyte.     

Mealybug, Phenococcus solani, and barley aphid, Sipha maydis, response to endophyte-infected tall and meadow fescues

Mealybugs and aphids are insects which damage grass species. The effects of fungal endophytes on the feeding of the mealybug, Phenococcus solani Ferris (Homoptera: Pseudococcidae), and barley aphid, Sipha maydis Passerini (Homoptera: Aphididae), on tall fescue, Festuca arundinacea Schreb. and meadow fescue, Festuca pratensis Huds., were studied under greenhouse conditions. Mealybugs preferred endophyte‐free (E–) clones over their endophyte‐infected (E+) counterparts. E+ plants had a significantly lower number of mealybugs than E– plants. A mixture of E+ and E– plants supported intermediate mealybug numbers, between pure plantings of E+ and E– grasses. Barley aphids released on to plant materials were deterred from feeding and could not persist on E+ plants. E– plants did not survive because of aphid damage, while E+ plants generally re‐grew, but were damaged to some degree. The results showed that the use of pure stands of endophyte‐infected grasses or a mixed stand of infected and non‐infected plants may increase the persistence and durability of turf and forage grass species in the presence of foliar damaging insects.     

The effect ofAcremonium coenophialum on the growth and nematode infestation of tall fescue

The presence of the endophytic fungusAcremonium coenophialum Morgan-Jones et Gams in tall fescue (Festuca arundinacea Schreb.) induces toxicity when this grass is grazed by cattle; however, there is evidence that removing the endophyte reduces the stand vigor and longevity of fescue. A field trial was conducted to determine the effects of water supply and the presence of the endophytic fungus on plant growth, drought tolerance, and soil nematode populations in Kentucky 31 tall fescue. The design included two factors, level of endophyte infection (0 and 75%) and irrigation regime (none, low, and high). Where water deficits occurred, herbage yield and leaf area were lower, and percentage dead tissue and canopy minus air temperature were greater in endophyte-free compared with endophyte-infected fescue. Soil populations ofPratylenchus scribneri andTylenchorhynchus acutus were substantially higher in the noninfected than in the endophyte-infected plots. The endophyte apparently confers drought tolerance to Kentucky 31 tall fescue, and this effect may be at least partially mediated through enhanced resistance to soil-borne nematodes.Published with the approval of the Director of the Ark. Agric. Exp. Stn.     

Short-day and Low-temperature Control of Floral Induction in Festuca

The conditions necessary for floral induction to occur in tallfescue (Festuca arundinacea), meadow fescue (Festuca pratensis),and red fescue (Festuca rubra), have been investigated. Onlya Tunisian ecotype of tall fescue produced inflorescences undershort-day conditions when air temperatures were above 8 °C.Under short days with low temperatures nearly all plants ofS. 170 tall fescue and S. 215 meadow fescue produced inflorescencesafter 15 weeks' exposure, but S. 59 red fescue showed only asmall response. Evidence was obtained for the existence in bothtall fescue and meadow fescue of a juvenile stage during whichplants showed a reduced response to inductive conditions. Avariation of 35 days in the required length of exposure to inductiveconditions was demonstrated between families within the S. 170variety of tall fescue, indicating the possibility of selectingfor larger or smaller inductive requirements. A second generationof seed was produced within a 12-month period from inflorescenceswhich had developed in a heated glasshouse during the wintermonths.     

Variation in Nitrogen Utilization in Acremonium coenophialum Isolates

Acremonium coenophialum, a fungal endosymbiont in tall fescue, is responsible for the production of alkaloid toxins. Animals grazing endophyte-infected tall fescue often show toxicosis. In marginal environments, the endophyte is important for long-term survival of tall fescue. Few differences in endophyte isolates from individual tall fescue plants have been reported. To aid development of a toxicosis-free tall fescue, it is important to identify differences in endophyte isolates. This report describes variation in nitrogen utilization in a defined culture medium by endophyte isolates from Kentucky-31 tall fescue. Overall, the best nitrogen sources for dry weight production of mycelium were proline and potassium nitrate. Thirty-four isolates grown on agar-solidified defined media with single nitrogen sources showed variation in nitrogen utilization. Fifty percent of the isolates were unable to utilize two or more amino acids. Manipulation of endophyte variation could lead to development of a toxicosis-free tall fescue cultivar.     

Effect of Ploidy on Quality of Tall Fescue, Italian Ryegrass x Tall Fescue and Tall Fescue x Giant Fescue Hybrids

Selected quality parameters were measured for forage leaf tissuefrom a spaced-plant nursery. The genotypes used were Ky 31 tallfescue and hybrids of Italian ryegrass (Lolium multiflorum Lam.)x tall fescue (Festuca arundinacea Schreb.) and tall fescuex giant fescue [Fescue gigantea (L ) Vill.]. Hybrid ploidy rangedfrom 2n = 28 to 84 chromosomes. Forage quality was characterizedby neutral detergent fibre (NDF), acid detergent fibre (ADF),total soluble carbohydrates (TSC) nutritive value index (NVI),hemicellulose, and in vitro dry matter disappearance (DMD). Quality of tall fescue, as measured by increased DMD, was improvedby hybridization with giant fescue. Improved DMD and NVI correlatedwith lower NDF and ADF in the hybrids. A few hybrids of Italianryegrass x tall fescue (2n = 28) were higher in some qualityparameters than Ky 31. Tall fescue x giant fescue hybrids (2n= 80 to 84), as a group, had significant quality improvementover Ky 31 in higher DMD and NVI and lower NDF and ADF. Whilesome individual hybrids within each group were significantlyhigher in quality, only the 2n = 80 to 84 chromosome group wasconsistently higher than Ky 31. Prediction equations for DMD,NDF, and ADF were established based on solvent extraction withnear-infrared reflectance spectroscopy (NIRS). Linear correlationcoefficients between chemical measurement and NIRS for eachquality parameter were 0–95 or higher. Acid detergent fibre, neutral detergent fibre, dry matter disappearance, hemicellulose, nutritive value index, Festuca arundinacea, Festuca gigantea, Lolium multiflorum     

Influence of phosphorus on the growth and ergot alkaloid content of Neotyphodium coenophialum-infected tall fescue (Festuca arundinacea Schreb.)

Tall fescue (Festuca arundinacea Schreb.) plants infected by the fungal endophyte Neotyphodium coenophialum (Morgan-Jones & Gams) (Glenn et al., 1996) often perform better than noninfected plants, especially in marginal resource environments. There is a lack of information about endophyte related effects on the rhizosphere of grasses. In a greenhouse experiment, four endophyte-infected (E+) tall fescue clones (DN2, DN4, DN7, DN11) and their endophyte-free (E–) forms were grown in limed (pH 6.3) Porter soil (low fertility, acidic, high aluminum and low phosphorus content, coarse-loamy mixed mesic Umbric Dystrochrept) at three soil P levels (17, 50, and 96 mg P kg^-1 soil) for five months. Excluding the genotype effect, endophyte infection significantly increased cumulative herbage DM yield by 8% at 17 mg P kg^-1 soil but reduced cumulative herbage DM yield by 12% at 96 mg P kg^-1 soil. With increased P availability in the soil, shoot and root DM, and root/shoot ratio in E+ plants were significantly less when compared to E– plants. Endophyte infection increased specific root length at 17 and 50 mg P kg^-1soil. At soil P level of 17 mg P kg^-1soil, E+ plants had significantly higher P concentrations both in roots and shoots. Similar relationships were found for Mg and Ca. E+ plants had significantly higher Zn, Fe, and Al concentration in roots, and lower Mn and Al concentration in shoots when compared to E– plants. Ergot alkaloid concentration and content in shoot of E+ plants increased with increasing P availability in the soil from 17 to 50 mg P kg^-1 but declined again at 96 mg P kg^-1 soil. Ergot alkaloid accumulation in roots increased linearly with P availability in the soil. Results suggest that endophyte infection affects uptake of phosphorus and other mineral nutrients and may benefit tall fescue grown on P-deficient soils. Phosphorus seems also to be involved in ergot alkaloid accumulation in endophyte-infected tall fescue.     

Shoot specific fungal endophytes alter soil phosphorus (P) fractions and potential acid phosphatase activity but do not increase P uptake in tall fescue

Aims

An experiment was performed to test how different fungal endophyte strains influenced tall fescue’s ability to access P from four P sources varying in solubility.

Methods

Novel endophyte infected (AR542E+ or AR584E+), common toxic endophyte infected (CTE+), or endophyte-free (E-) tall fescues were grown for 90 days in acidic soils amended with 30 mg kg^?1 P of potassium dihydrogen phosphate (KH₂PO₄), iron phosphate (FePO₄), aluminum phosphate (AlPO₄), or tricalcium phosphate ((Ca₃(PO₄)₂), respectively.

Results

Phosphorus form strongly influenced plant biomass, P acquisition, agronomic P use efficiency, microbial communities, P fractions. P uptake and vegetative biomass were similar for plants grown with AlPO₄, Ca₃(PO₄)₂, and KH₂PO₄ but greater than in control and FePO₄ soils. Infection with AR542E+ resulted in significantly less shoot biomass than CTE+ and E- varieties; there was no influence of endophyte on root biomass. The biomarker for arbuscular mycorrhizal fungi (AM fungi, 16:1ω5c) was selected as an effective predictor of variations in P uptake and tall fescue biomass. Potential acid phosphatase activity was strongly influenced by endophyte x P form interaction.

Conclusions

Endophyte infection in tall fescue significantly affected the NaOH-extractable inorganic P fraction, but had little detectable influence on soil microbial community structure, root biomass, or P uptake.

     

Insect Feeding Deterrents in Endophyte-Infected Tall Fescue

The presence of an endophytic fungus, Acremonium coenophialum, in tall fescue (Festuca arundinacea) deterred aphid feeding by Rhopalosiphum padi and Schizaphis graminum. Both species of aphid were unable to survive when confined to endophyte-infected tall fescue plants. Feeding deterrents and toxic factors to R. padi and Oncopeltus fasciatus, large milkweed bug, were primarily associated with a methanol extract obtained when endophyte-infected tall fescue seed was serially extracted with hexane, ethyl acetate, and methanol. The concentrations of pyrrolizidine alkaloids were determined to be 30 to 100 times greater in the methanol extract than in the hexane and ethyl acetate extracts.     

Insect oviposition preference between Epichloë‐symbiotic and Epichloë‐free grasses does not necessarily reflect larval performance

Variation in plant communities is likely to modulate the feeding and oviposition behavior of herbivorous insects, and plant‐associated microbes are largely ignored in this context. Here, we take into account that insects feeding on grasses commonly encounter systemic and vertically transmitted (via seeds) fungal Epichloë endophytes, which are regarded as defensive grass mutualists. Defensive mutualism is primarily attributable to alkaloids of fungal origin. To study the effects of Epichloë on insect behavior and performance, we selected wild tall fescue (Festuca arundinacea) and red fescue (Festuca rubra) as grass–endophyte models. The plants used either harbored the systemic endophyte (E+) or were endophyte‐free (E?). As a model herbivore, we selected the Coenonympha hero butterfly feeding on grasses as larvae. We examined both oviposition and feeding preferences of the herbivore as well as larval performance in relation to the presence of Epichloë endophytes in the plants. Our findings did not clearly support the female's oviposition preference to reflect the performance of her offspring. First, the preference responses depended greatly on the grass–endophyte symbiotum. In F. arundinacea, C. hero females preferred E+ individuals in oviposition‐choice tests, whereas in F. rubra, the endophytes may decrease exploitation, as both C. hero adults and larvae preferred E? grasses. Second, the endophytes had no effect on larval performance. Overall, F. arundinacea was an inferior host for C. hero larvae. However, the attraction of C. hero females to E+ may not be maladaptive if these plants constitute a favorable oviposition substrate for reasons other than the plants' nutritional quality. For example, rougher surface of E+ plant may physically facilitate the attachment of eggs, or the plants offer greater protection from natural enemies. Our results highlight the importance of considering the preference of herbivorous insects in studies involving the endophyte‐symbiotic grasses as host plants.