• Chloroplast microsatellite markers were developed for Festuca rubra to examine its population g... more • Chloroplast microsatellite markers were developed for Festuca rubra to examine its population genetic characteristics, taxonomy, and coevolution with its endophyte Epichloë festucae. • Thirteen polymorphic markers were identified from the chloroplast genome of a F. ovina accession and intergenic chloroplast sequences of F. rubra accessions. They amplified a total of 65 alleles in a sample of 93 individuals of F. rubra originating from six different populations located in the Faroe Islands, Finland, Greenland, Norway, and Spain. • The developed microsatellite primer pairs can be used by researchers in population genetic and taxonomic studies, and by plant breeders in breeding programs on grasses.
We studied the effects of fungal endophyte infection of meadow ryegrass (Lolium pratense=Festuca ... more We studied the effects of fungal endophyte infection of meadow ryegrass (Lolium pratense=Festuca pratensis) on the frequency of the barley yellow dwarf virus (BYDV). The virus is transferred by aphids, which may be deterred by endophyte-origin alkaloids within the plant. In our experiment, we released viruliferous aphid vectors on endophyte-infected and endophyte-free plants in a common garden. The number of aphids and the percentage of BYDV infections were lower in endophyte-infected plants compared to endophyte-free plants, indicating that endophyte infection may protect meadow ryegrass from BYDV infections.
Grasses have been considered to primarily employ tolerance in lieu of defense in mitigating damag... more Grasses have been considered to primarily employ tolerance in lieu of defense in mitigating damage caused by herbivory. Yet a number of mechanisms have been identified in grasses, which may deter feeding by grazers. These include enhanced silicon uptake, hosting of toxin-producing endophytic fungi and induction of secondary metabolites. While these mechanisms have been individually studied, their synergistic responses to grazing, as well as their effects on grazers, are poorly known. A field experiment was carried out in 5 × 5 m outdoor enclosures to quantify phytochemical changes of either endophyte-infected (E+) or endophyte-free (E-) meadow fescue (Schedonorus pratensis) in response to medium intensity (corresponding with densities of ca. 1200 voles/ha for 5 weeks during 3 months) or heavy intensity (ca. 1200 voles/ha for 8 weeks during 3 months) grazing by a mammalian herbivore, the field vole (Microtus agrestis). A laboratory experiment was then conducted to evaluate the effects of endophyte infection status and grazing history of the grass diet on vole performance. As predicted, grazing increased foliar silicon content, by up to 13%. Grazing also increased foliar levels of phosphorous and several phenolic compounds, most notably those of the flavonols isorhamnetin-diglycoside and rhamnetin derivative. Silicon concentrations were consistently circa 16% higher in E+ grasses than in E-grasses, at all levels of grazing. Similarly, concentrations of chlorogenic acid derivative were found to be consistently higher in E+ than in E- grasses. Female voles maintained on heavily grazed grasses suffered higher mortality rates in the laboratory than female voles fed ungrazed grass, regardless of endophyte infection status. Our results conclusively demonstrate that, in addition to tolerance, grasses employ multi-tiered, effective defenses against mammalian grazers.
ABSTRACT A symbiosis between grasses and systemic fungal endophytes exists in both natural and ag... more ABSTRACT A symbiosis between grasses and systemic fungal endophytes exists in both natural and agricultural grassland communities. Our objective was to examine the effects of systemic endophytes on the competitive ability of two agronomically important grass species: meadow fescue [Festuca pratensis (Huds.) syn. Schedonorus pratensis (Huds.) P. Beauv] and tall fescue [Festuca arundinacea (Schreb.) syn. Schedonorus phoenix (Scop.)]. Plants of meadow and tall fescue were grown for 48 days in replacement series of interspecific mixture with a legume (red clover, Trifolium pratense L.) in different nutrient environments in a greenhouse. Neither of the grass species gained endophyte-promoted competitive advantage over red clover in grass–clover mixtures. Endophyte infection increased the growth of meadow fescue monocultures by 89% compared to endophyte-free monocultures in high-nutrient soils, but plant competition or the cost of endophyte infection to the meadow fescue decreased the yield in resource-limited conditions. On average, endophyte-infected and endophyte-free meadow fescues produced 0·15 and 0·17 g, and 0·14 and 0·14 g dry biomass per plant in mixtures with red clover in high- and low-nutrient soils respectively. In contrast to meadow fescue, endophyte-promoted growth of tall fescue monocultures was not detected. Endophyte-infected and endophyte-free tall fescue monocultures produced 0·76 and 0·95 g biomass per pot, respectively, in the high-nutrient environment. Endophyte infection can increase the performance of the host grass, but the positive effects depend on the host species, the species composition and soil nutrient availability.
ABSTRACT Exploring the regional pattern of variation in traits driven by symbiotic interactions m... more ABSTRACT Exploring the regional pattern of variation in traits driven by symbiotic interactions may provide insights to understand the evolutionary processes that operate over plant populations. Polyploidy, which is associated with fitness improvement, is expected to increase with latitude and altitude. However, it has never been explored in relation with the occurrence of epichloid fungal endophytes in plants. Both, variation in ploidy level and in the incidence of fungal endophytes, are known to occur in species of fine fescues. Here, we surveyed the occurrence of systemic fungal endophytes in natural Festuca vivipara populations in North European islands. In addition, we identified the fungal species associated with this grass and determined the predominant ploidy level for each population. Endophytes were found in four of six, two of three, and one of three populations for Faroe Islands, Iceland and Great Britain, respectively. With an average low incidence level of 15 % in infected populations, there was no relationship between infection level and either latitude or altitude. The phylogenetic analysis based on sequences ITS and the tub2 genes, supports that the endophytic species is Epichloë festucae, the same as in other fine fescues. We found no variation in ploidy level as all the plants were tetraploid (4X) with 28 chromosomes, a pattern which contrasts with the variation reported in previous antecedents. Our results suggest that apart from low and variable benefits of the endophyte to the plants, there would be a complex dynamics between epichloid endophytes and species of the fine fescue complex which merits further studies.
Seedborne systemic endophytic fungi of grasses are thought to be plant mutualists, because they h... more Seedborne systemic endophytic fungi of grasses are thought to be plant mutualists, because they have been shown to improve their host's resistance against biotic and abiotic stresses. The interactions in plant-endophyte associations vary from mutualistic to parasitic with environmental conditions and the genotypes of interacting species. The possible pros and cons of endophytic fungi are expected to be most evident during the seedling establishment, where host fitness is most directly affected. If this holds true, endophytes may play a focal role in local adaptation of hosts to different environments. We examined if endophyte-infected and uninfected seeds and seedlings of two native grass species, Festuca rubra and F. ovina, differ in seed germination and seedling growth rates under greenhouse conditions. The germination of F. rubra seeds was also studied in the field. This is the first time that the effects of Epichloë endophyte on seedling establishment of fine fescues from natural populations have been experimentally evaluated. Mother plant (seed family) had a marked effect on many response variables in both grass species. Length and mean biomass of tillers of endophyte-infected (E+) F. ovina seedlings were lower, but root:shoot ratios were higher than in endophyte-free (E-) seedlings. In F. rubra, the effects of the endophyte were dependent on the habitat where the seeds were collected. The E+ seeds from river banks germinated faster than E+ seeds from meadows, and E+ seedlings from the river banks produced fewer but taller and heavier tillers than the other seedlings. Our data suggest that the effects of the endophyte infection on the seedling stage of fine fescues are dependent the species of grass, host genetic background and mother plant habitat. The germination strategy and growth form of E+ red fescue seedlings from river banks may be beneficial to surviving in the harsh conditions of that habitat.
Page 1. Abstract Neotyphodium endophytes in introduced ag-ronomic grasses are well known to incre... more Page 1. Abstract Neotyphodium endophytes in introduced ag-ronomic grasses are well known to increase resistance to herbivores, but little is known of interactions between Neotyphodium endophytes and herbivores in native grass populations. ...
The effect of environmental fragmentation on the species distribution and frequency of horizontal... more The effect of environmental fragmentation on the species distribution and frequency of horizontally transmitted endophytic fungi in Betula pubescens and Betula pendula leaves was studied in an archipelago in southwestern Finland. The study system consisted of 14 islands, ranging in size and distance to the mainland, and five mainland sites. Endophytic fungi were grown out from surface-sterilized leaves. The frequency of endophytic fungi mainly depended on the size of the island, explaining 32-35% of the variation, and the distance to the mainland explaining 29-35% of the variation. The birch trees on the largest islands near the mainland had the highest endophyte frequencies. Fusicladium betulae, Gnomonia setacea and Melanconium betulinum were the most commonly isolated fungi. Foliar endophytes of birch trees are able to disperse to fairly fragmented areas, but their frequencies seem to depend on environmental isolation and size of the island.
• Chloroplast microsatellite markers were developed for Festuca rubra to examine its population g... more • Chloroplast microsatellite markers were developed for Festuca rubra to examine its population genetic characteristics, taxonomy, and coevolution with its endophyte Epichloë festucae. • Thirteen polymorphic markers were identified from the chloroplast genome of a F. ovina accession and intergenic chloroplast sequences of F. rubra accessions. They amplified a total of 65 alleles in a sample of 93 individuals of F. rubra originating from six different populations located in the Faroe Islands, Finland, Greenland, Norway, and Spain. • The developed microsatellite primer pairs can be used by researchers in population genetic and taxonomic studies, and by plant breeders in breeding programs on grasses.
We studied the effects of fungal endophyte infection of meadow ryegrass (Lolium pratense=Festuca ... more We studied the effects of fungal endophyte infection of meadow ryegrass (Lolium pratense=Festuca pratensis) on the frequency of the barley yellow dwarf virus (BYDV). The virus is transferred by aphids, which may be deterred by endophyte-origin alkaloids within the plant. In our experiment, we released viruliferous aphid vectors on endophyte-infected and endophyte-free plants in a common garden. The number of aphids and the percentage of BYDV infections were lower in endophyte-infected plants compared to endophyte-free plants, indicating that endophyte infection may protect meadow ryegrass from BYDV infections.
Grasses have been considered to primarily employ tolerance in lieu of defense in mitigating damag... more Grasses have been considered to primarily employ tolerance in lieu of defense in mitigating damage caused by herbivory. Yet a number of mechanisms have been identified in grasses, which may deter feeding by grazers. These include enhanced silicon uptake, hosting of toxin-producing endophytic fungi and induction of secondary metabolites. While these mechanisms have been individually studied, their synergistic responses to grazing, as well as their effects on grazers, are poorly known. A field experiment was carried out in 5 × 5 m outdoor enclosures to quantify phytochemical changes of either endophyte-infected (E+) or endophyte-free (E-) meadow fescue (Schedonorus pratensis) in response to medium intensity (corresponding with densities of ca. 1200 voles/ha for 5 weeks during 3 months) or heavy intensity (ca. 1200 voles/ha for 8 weeks during 3 months) grazing by a mammalian herbivore, the field vole (Microtus agrestis). A laboratory experiment was then conducted to evaluate the effects of endophyte infection status and grazing history of the grass diet on vole performance. As predicted, grazing increased foliar silicon content, by up to 13%. Grazing also increased foliar levels of phosphorous and several phenolic compounds, most notably those of the flavonols isorhamnetin-diglycoside and rhamnetin derivative. Silicon concentrations were consistently circa 16% higher in E+ grasses than in E-grasses, at all levels of grazing. Similarly, concentrations of chlorogenic acid derivative were found to be consistently higher in E+ than in E- grasses. Female voles maintained on heavily grazed grasses suffered higher mortality rates in the laboratory than female voles fed ungrazed grass, regardless of endophyte infection status. Our results conclusively demonstrate that, in addition to tolerance, grasses employ multi-tiered, effective defenses against mammalian grazers.
ABSTRACT A symbiosis between grasses and systemic fungal endophytes exists in both natural and ag... more ABSTRACT A symbiosis between grasses and systemic fungal endophytes exists in both natural and agricultural grassland communities. Our objective was to examine the effects of systemic endophytes on the competitive ability of two agronomically important grass species: meadow fescue [Festuca pratensis (Huds.) syn. Schedonorus pratensis (Huds.) P. Beauv] and tall fescue [Festuca arundinacea (Schreb.) syn. Schedonorus phoenix (Scop.)]. Plants of meadow and tall fescue were grown for 48 days in replacement series of interspecific mixture with a legume (red clover, Trifolium pratense L.) in different nutrient environments in a greenhouse. Neither of the grass species gained endophyte-promoted competitive advantage over red clover in grass–clover mixtures. Endophyte infection increased the growth of meadow fescue monocultures by 89% compared to endophyte-free monocultures in high-nutrient soils, but plant competition or the cost of endophyte infection to the meadow fescue decreased the yield in resource-limited conditions. On average, endophyte-infected and endophyte-free meadow fescues produced 0·15 and 0·17 g, and 0·14 and 0·14 g dry biomass per plant in mixtures with red clover in high- and low-nutrient soils respectively. In contrast to meadow fescue, endophyte-promoted growth of tall fescue monocultures was not detected. Endophyte-infected and endophyte-free tall fescue monocultures produced 0·76 and 0·95 g biomass per pot, respectively, in the high-nutrient environment. Endophyte infection can increase the performance of the host grass, but the positive effects depend on the host species, the species composition and soil nutrient availability.
ABSTRACT Exploring the regional pattern of variation in traits driven by symbiotic interactions m... more ABSTRACT Exploring the regional pattern of variation in traits driven by symbiotic interactions may provide insights to understand the evolutionary processes that operate over plant populations. Polyploidy, which is associated with fitness improvement, is expected to increase with latitude and altitude. However, it has never been explored in relation with the occurrence of epichloid fungal endophytes in plants. Both, variation in ploidy level and in the incidence of fungal endophytes, are known to occur in species of fine fescues. Here, we surveyed the occurrence of systemic fungal endophytes in natural Festuca vivipara populations in North European islands. In addition, we identified the fungal species associated with this grass and determined the predominant ploidy level for each population. Endophytes were found in four of six, two of three, and one of three populations for Faroe Islands, Iceland and Great Britain, respectively. With an average low incidence level of 15 % in infected populations, there was no relationship between infection level and either latitude or altitude. The phylogenetic analysis based on sequences ITS and the tub2 genes, supports that the endophytic species is Epichloë festucae, the same as in other fine fescues. We found no variation in ploidy level as all the plants were tetraploid (4X) with 28 chromosomes, a pattern which contrasts with the variation reported in previous antecedents. Our results suggest that apart from low and variable benefits of the endophyte to the plants, there would be a complex dynamics between epichloid endophytes and species of the fine fescue complex which merits further studies.
Seedborne systemic endophytic fungi of grasses are thought to be plant mutualists, because they h... more Seedborne systemic endophytic fungi of grasses are thought to be plant mutualists, because they have been shown to improve their host's resistance against biotic and abiotic stresses. The interactions in plant-endophyte associations vary from mutualistic to parasitic with environmental conditions and the genotypes of interacting species. The possible pros and cons of endophytic fungi are expected to be most evident during the seedling establishment, where host fitness is most directly affected. If this holds true, endophytes may play a focal role in local adaptation of hosts to different environments. We examined if endophyte-infected and uninfected seeds and seedlings of two native grass species, Festuca rubra and F. ovina, differ in seed germination and seedling growth rates under greenhouse conditions. The germination of F. rubra seeds was also studied in the field. This is the first time that the effects of Epichloë endophyte on seedling establishment of fine fescues from natural populations have been experimentally evaluated. Mother plant (seed family) had a marked effect on many response variables in both grass species. Length and mean biomass of tillers of endophyte-infected (E+) F. ovina seedlings were lower, but root:shoot ratios were higher than in endophyte-free (E-) seedlings. In F. rubra, the effects of the endophyte were dependent on the habitat where the seeds were collected. The E+ seeds from river banks germinated faster than E+ seeds from meadows, and E+ seedlings from the river banks produced fewer but taller and heavier tillers than the other seedlings. Our data suggest that the effects of the endophyte infection on the seedling stage of fine fescues are dependent the species of grass, host genetic background and mother plant habitat. The germination strategy and growth form of E+ red fescue seedlings from river banks may be beneficial to surviving in the harsh conditions of that habitat.
Page 1. Abstract Neotyphodium endophytes in introduced ag-ronomic grasses are well known to incre... more Page 1. Abstract Neotyphodium endophytes in introduced ag-ronomic grasses are well known to increase resistance to herbivores, but little is known of interactions between Neotyphodium endophytes and herbivores in native grass populations. ...
The effect of environmental fragmentation on the species distribution and frequency of horizontal... more The effect of environmental fragmentation on the species distribution and frequency of horizontally transmitted endophytic fungi in Betula pubescens and Betula pendula leaves was studied in an archipelago in southwestern Finland. The study system consisted of 14 islands, ranging in size and distance to the mainland, and five mainland sites. Endophytic fungi were grown out from surface-sterilized leaves. The frequency of endophytic fungi mainly depended on the size of the island, explaining 32-35% of the variation, and the distance to the mainland explaining 29-35% of the variation. The birch trees on the largest islands near the mainland had the highest endophyte frequencies. Fusicladium betulae, Gnomonia setacea and Melanconium betulinum were the most commonly isolated fungi. Foliar endophytes of birch trees are able to disperse to fairly fragmented areas, but their frequencies seem to depend on environmental isolation and size of the island.
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