Search

Antagonistic selection between pathogens and their hosts can drive rapid evolutionary change and leave distinct molecular footprints of past and ongoing selection in the genomes of the interacting species. Despite an increasing availability of tools able to identify signatures of selection, the genetic mechanisms underlying coevolutionary interactions and the specific genes involved are still poorly understood, especially in heterogeneous natural environments. We searched the genomes of two species of Epichloe plant pathogen for evidence of recent selection. The Epichloe genus includes highly host‐specific species that can sterilize their grass hosts. We performed selection scans using genome‐wide SNP data from seven natural populations of two co‐occurring Epichloe sibling species specialized on different hosts. We found evidence of recent (and ongoing) selective sweeps across the genome in both species. However, selective sweeps were more abundant in the species with a larger effective population size. Sweep regions often overlapped with highly polymorphic AT‐rich regions supporting the role of these genome compartments in adaptive evolution. Although most loci under selection were specific to individual populations, we could also identify several candidate genes targeted by selection in sweep regions shared among populations. The genes encoded small secreted proteins typical of fungal effectors and cell wall‐degrading enzymes. By investigating the genomic signatures of selection across multiple populations and species, this study contributes to our understanding of complex adaptive processes in natural plant pathogen systems. [ABSTRACT FROM AUTHOR]

Abstract: Host specialization is a key process in ecological divergence and speciation of plant‐associated fungi. The underlying determinants of host specialization are generally poorly understood, especially in endophytes, which constitute one of the most abundant components of the plant microbiome. We addressed the genetic basis of host specialization in two sympatric subspecies of grass‐endophytic fungi from the Epichloë typhina complex: subsp. typhina and clarkii. The life cycle of these fungi entails unrestricted dispersal of gametes and sexual reproduction before infection of a new host, implying that the host imposes a selective barrier on viability of the progeny. We aimed to detect genes under divergent selection between subspecies, experiencing restricted gene flow due to adaptation to different hosts. Using pooled whole‐genome sequencing data, we combined FST and DXY population statistics in genome scans and detected 57 outlier genes showing strong differentiation between the two subspecies. Genomewide analyses of nucleotide diversity (π), Tajima's D and dN/dS ratios indicated that these genes have evolved under positive selection. Genes encoding secreted proteins were enriched among the genes showing evidence of positive selection, suggesting that molecular plant–fungus interactions are strong drivers of endophyte divergence. We focused on five genes encoding secreted proteins, which were further sequenced in 28 additional isolates collected across Europe to assess genetic variation in a larger sample size. Signature of positive selection in these isolates and putative identification of pathogenic function supports our findings that these genes represent strong candidates for host specialization determinants in Epichloë endophytes. Our results highlight the role of secreted proteins as key determinants of host specialization. [ABSTRACT FROM AUTHOR]

The article discusses the research on the identification of Piedmont white truffles also known as tuber magnatum Pico at European Alps. The research aims to study the impact of climate change on mycorrhizal fungal species. Topics include the geographic distribution and phylogeny of the truffles, the DNA extraction analysis for the fungal fruits, and the sampling of soil.

Interspecific hybrid endophytes of the genus Epichloë (Ascomycota, Clavicipitaceae) are prevalent in wild grass populations, possibly because of their larger gene variation, resulting in increased fitness benefits for host plants; however, the reasons are not yet known. We tested hypotheses regarding niche expansion mediated by hybrid endophytes, population-dependent interactions and local co-adaptation in the woodland grass Hordelymus europaeus, which naturally hosts both hybrid and non-hybrid endophyte taxa., Seedlings derived from seeds of four grass populations made endophyte free were re-inoculated with hybrid or non-hybrid endophyte strains, or left endophyte free. Plants were grown in the glasshouse with or without drought treatment., Endophyte infection increased plant biomass and tiller production by 10-15% in both treatments. Endophyte types had similar effects on growth, but opposite effects on reproduction: non-hybrid endophytes increased seed production, whereas hybrid endophytes reduced or prevented it completely., The results are consistent with the observation that non-hybrid endophytes in H. europaeus prevail at dry sites , but cannot explain the prevalence of hybrid endophytes. Thus, our results do not support the hypothesis of niche expansion of hybrid-infected plants. Moreover, plants inoculated with native relative to foreign endophytes yielded higher infections, but both showed similar growth and survival, suggesting weak co-adaptation. [ABSTRACT FROM AUTHOR]

Epichloid fungal endophytes ( Epichloë and Neotyphodium spp.) are excellent model systems for studying speciation processes because of their variable life history traits that are linked to host grass fitness. Presumed jumps to new hosts and subsequent somatic hybridizations appear to be common among epichloid endophytes resulting in increased genetic variation upon which selection can act and speciation be initiated. In this study, we explored the endophyte diversity of a rare European native woodland grass species, Hordelymus europaeus, along a latitudinal transect covering the entire distribution range of H. europaeus. From 28 populations in six countries, isolates were sampled and molecularly characterized. Based on the sequences of tubB and tefA, six distinct epichloid taxa (interspecific hybrid or cryptic haploid species) were found, of which four were novel and two have been previously reported from this host. Of the novel endophytes, two were presumed to be interspecific hybrids and two of nonhybrid origin. While previously known endophytes of H. europaeus are seed-born and strictly asexual, one of the novel nonhybrid endophytes found in the glacial refugium of the Apennine peninsula reproduced sexually in cultured plants. This is the first case of a seed-borne, but sexually reproducing endophyte of this host. We discuss the origin, and possible ancestral species, of the six epichloid taxa using phylogenetic analyses. Repeated host jumps and somatic hybridizations characterize the diversity of the endophytes. To date, no other grass species is known to host a larger diversity of endophytes than H. europaeus. [ABSTRACT FROM AUTHOR]

Epichloë spp., fungal endophytes of cool season grasses, produce collars of mycelium (stromata) on host stems that Botanophila flies visit for egg laying. Flies transfer fungal gametes among stromata and thereby serve to cross-fertilize fungi. Hence, the interaction is analogous to insect pollination in angiosperms. While most Epichloë species are not interfertile, Epichloë typhina and Epichloë clarkii can hybridize. Weinvestigated whether Botanophila flies play a role in the reproductive isolation of the two Epichloë species at a field site in southwestern Switzerland. We estimated the density of stromata and collected fly larvae and stromata occurring on plants. While most ascospores collected from both species indicated intraspecific mating, 9.3%of fungal fruiting bodies contained spores of hybrid origin. Two species of Botanophila larvae occurred on stromata and both preferred E. typhina. Yet, both fly species laid eggs on both fungal species. While preferences by Botanophila flies should influence reproductive isolation between the fungi, other mechanisms are likely more important. Our data, which show hybrid ascospores are produced, suggest postzygotic isolating mechanisms are an important means of reproductive isolation. [ABSTRACT FROM AUTHOR]

• Systemic fungal endophytes (Clavicipitaceae) of grasses reproduce sexually when the fungus forms stromata and contagious ascospores, or asexually by vertical transmission of hyphae into seeds and seedlings. Vertical transmission is predicted to favor reduced virulence compared with horizontal transmission in systems with both types of transmission. • Here, variation in vertical and horizontal transmission and its potential heritability in a host grass–endophyte interaction, Elymus hystrix infected with Epichloë elymi, were examined in natural populations and two common garden experiments using field-collected host tillers and seed progeny of maternal plants with known infection phenotypes. • Transmission mode exhibited year-to-year variation in field and common garden environments. In the common garden there were consistent differences among maternal plant families in stroma production and significant correlations between stroma production in the common garden and in natural populations. Transmission mode differed among maternal families, spanning a continuum from pure vertical transmission to a high proportion of stroma production and horizontal transmission potential. Vertical transmission to seeds occurred at high rates in all maternal families regardless of their stroma production. • Observed patterns of variation indicate that endophyte transmission mode and correlated changes in virulence can respond to selection by biotic and abiotic factors. [ABSTRACT FROM AUTHOR]

• Grass-infecting Epichloëendophytes (Ascomycota, Calvicipitaceae) depend on Botanophila flies for gamete transfer, while fly larvae feed and develop on the fertilized fungal fruiting structures. Flies are known to be attracted by volatile signals, but the exact mechanisms of chemical communication and the degree of specialization are unknown. • Headspace samples collected from five different Epichloëspecies were analysed with respect to physiologically active substances using Botanophila flies. In field bioassays using synthetic compounds, their attractiveness and the specificity of the Epichloë–Botanophila attraction were investigated. • The identification of a new natural product, methyl ( Z)-3-methyldodec-2-enoate, attracting Botanophila flies is reported here, and chokol K is confirmed as an attractive compound. Different blends of the two compounds attracted Botanophila flies under field conditions, but the three fly taxa present at the study site showed no preference for specific blends of volatiles. • Chemical communication in the Epichloë– Botanophila system relies on a few specific compounds, known as a communication system with ‘private channels’. Although ratios of emitted compounds vary in different Epichloë species, this seems not to lead to specialized attraction of Botanophila flies. Low selective pressure for specialization may have maintained a more generalist interaction between fungi and flies. [ABSTRACT FROM AUTHOR]

Species of Epichloë (Ascomycota: Clavicipitaceae), which infect grasses, maintain an obligate symbiotic relationship with flies of the genus Botanophila (Diptera: Anthomyiidae) . Sexual reproduction of the fungus usually requires a visit by Botanophila, which serves to ‘pollinate’ the fungus with spermatia of the opposite mating type; the flies in turn deposit their eggs on fungal tissues upon which the larvae feed. For a molecular phylogenetic study, a total of 108 fly larvae were collected from 10 different Epichloë species on various grasses in Europe and North America. Sequence analysis of the mitochondrial cytochrome oxidase gene ( COII) detected six distinct Botanophila taxa that were associated with Epichloë. Three were restricted to samples from Europe, two to samples from North America, and one was present on both continents. In North America the common taxon [identified as Botanophila lobata (Collin)] was found on Epichloë hosts of native grasses and on Epichloë typhina (Pers.: Fr.) Tul. and Epichloë festucae Leuchtm., Schardl and M.R. Siegel of the introduced grasses Dactylis glomerata L. and Festuca rubra L., respectively. In a maximum likelihood phylogram, one of the American taxa was most basal, while the other was nested within the European taxa, suggesting that European and American taxa of Epichloë-associated Botanophila spp. may not have separate origins. Analyses of the fly populations further indicated that up to four different Botanophila taxa can be present at a single location. On taxon level there was no specificity of flies for Epichloë host species (except taxon 6 that was found only on Epichloë glyceriae Schardl & Leuchtmann), although several taxa appeared to prefer some hosts over others. Comparison of molecular phylogenetic relationships of Botanophila species with those of the associated Epichloë hosts did not suggest co-evolution of the fungus and the fly. Thus, associations between Botanophila flies and Epichloë hosts may have arisen independently more than once. [ABSTRACT FROM AUTHOR]

• Altered environmental conditions may change populations of arbuscular mycorrhizal fungi and thereby affect mycorrhizal functioning. We investigated whether 8 yr of free-air CO2 enrichment has selected fungi that differently influence the nutrition and growth of host plants. • In a controlled pot experiment, two sets of seven randomly picked single spore isolates, originating from field plots of elevated (60 Pa) or ambient CO2 partial pressure (pCO2), were inoculated on nodulated Trifolium repens (white clover) plants. Fungal isolates belonged to the Glomus claroideum or Glomus intraradices species complex, and host plants were clonal micropropagates derived from nine genets. • Total nitrogen (N) concentration was increased in leaves of plants inoculated with fungal isolates from elevated-pCO2 plots. These isolates took up nearly twice as much N from the soil as isolates from ambient-pCO2 plots and showed much greater stimulation of biological N2 fixation. The morpho-species identity of isolates had a more pronounced effect on N2 fixation and on root length colonized than isolate identity. • We conclude that rising atmospheric pCO2 may select for fungal strains that will help their host plants to meet increased N demands. New Phytologist (2005) doi: 10.1111/j.1469-8137.2005.01440.x © New Phytologist (2005) [ABSTRACT FROM AUTHOR]

Rising atmospheric carbon dioxide partial pressure (pCO2) and nitrogen (N) deposition are important components of global environmental change. In the Swiss free air carbon dioxide enrichment (FACE) experiment, the effect of altered atmospheric pCO2 (35 vs. 60 Pa) and the influence of two different N-fertilization regimes (14 vs. 56 g N m−2 a−1) on root colonization by arbuscular mycorrhizal fungi (AMF) and other fungi (non-AMF) of Lolium perenne and Trifolium repens were studied. Plants were grown in permanent monoculture plots, and fumigated during the growth period for 7 years. At elevated pCO2 AMF and non-AMF root colonization was generally increased in both plant species, with significant effects on colonization intensity and on hyphal and non-AMF colonization. The CO2 effect on arbuscules was marginally significant ( P=0.076). Moreover, the number of small AMF spores (≤100 μm) in the soils of monocultures (at low-N fertilization) of both plant species was significantly increased, whereas that of large spores (>100 μm) was increased only in L. perenne plots. N fertilization resulted in a significant decrease of root colonization in L. perenne, including the AMF parameters, hyphae, arbuscules, vesicles and intensity, but not in T. repens. This phenomenon was probably caused by different C-sink limitations of grass and legume. Lacking effects of CO2 fumigation on intraradical AMF structures (under high-N fertilization) and no response to N fertilization of arbuscules, vesicles and colonization intensity suggest that the function of AMF in T. repens was non-nutritional. In L. perenne, however, AM symbiosis may have amended N nutrition, because all root colonization parameters were significantly increased under low-N fertilization, whereas under high-N fertilization only vesicle colonization was increased. Commonly observed P-nutritional benefits from AMF appeared to be absent under the phosphorus-rich soil conditions of our field experiment. We hypothesize that in well-fertilized agricultural ecosystems, grasses benefit from improved N nutrition and legumes benefit from increased protection against pathogens and/or herbivores. This is different from what is expected in nutritionally limited plant communities. [ABSTRACT FROM AUTHOR]

Epichloë species are self incompatible (heterothallic) fungi that must be fertilized by spermatia from individuals of opposite mating type for successful sexual reproduction to occur. Female flies of the genus Botanophila act as vectors of the fungi by ingesting and defecating spermatia (gametes) onto fungal stromata (fruiting bodies) after oviposition. Larvae feed and develop on the stromata and thus maintain a symbiotic relationship with Epichloë fungi. We hypothesized that sole dependence on fertilized stromata as a food source would promote specialization by flies to single compatible host species and that this specialization would promote reproductive isolation among Epichloë species. Analysis of progeny of ascospores from experimental field plots in Zurich, Switzerland, indicated prevalence of specific matings between stromata of the same host, and thus was consistent with the hypothesis that flies are species-specific in their visitation behaviour. Genetic analyses of spermatia contained in the faeces of individual flies also gave some support for this hypothesis. We recovered spermatia of 4 different Epichloë species from fly faeces. Comparison of spermatia found in fly faeces to those available from stromata showed flies avoided Epichloë clarkii and may have preferred Epichloë typhina. Interestingly, these are the only two Epichloë species known to be interfertile with one another. Individual flies tended to carry spermatia predominantly from one fungal species. Thus, flies may adopt a type of “majoring” and “minoring” behaviour when visiting fungi. Yet, Botanophila flies are not monolectic and often visited all hosts that were available within screened cages. In addition to any reproductive isolation flies may provide to some fungal species, differences in competitiveness among spermatia of different species deposited on the same stroma may favor intraspecific matings. [ABSTRACT FROM AUTHOR]

Compares the intraspecific competition of endophyte infected and uninfected plants of two woodland grass species. Factors influencing the plant community diversity; Projection of the beneficial effects of endophyte infection on wild grasses; Prevention of floral development.

Fungi in the genus Epichloë (Clavicipitaceae, Ascomycota) are endophytic and often mutualistic symbionts of many grasses in temperate areas. Species with a sexual cycle suppress host flowering and seed formation, whereas asexual fungi remain asymptomatic and transmit vertically by seed. Thus, the mode of reproduction may determine whether the symbiosis is mutualistic or parasitic. The level of sexual reproduction (disease expression) varies among different endophytes and on different grass hosts, but factors responsible for this variation, and evolutionary mechanisms leading to one or the other life strategy are not understood. As experimental system, we chose Brachypodium sylvaticum in which the endophyte E. sylvatica can express both reproduction modes. A field experiment was done in plots of a free air carbon dioxide enrichment (FACE) facility. We investigated the effects of three environmental factors (elevated CO2 concentration, shading and fertilisation) and one genetic factor (plant and fungal genotype combination), on plant growth and disease expression. Variation in plant growth was mainly dependent on the genotype and was increased by fertilisation. elevated CO2 and shading slightly stimulated plant growth, but only in fertilised plants. Disease expression was overwhelmingly dependent on the genotype, hence genetic factors. Fertilisation slightly stimulated disease expression in some genotypes, while the effect of elevated CO2 was negligible, and by interaction with fertilisation inconsistent in the two years. Horizontal transmissions during the experiment, presumably mediated by ascospores, confounded the original infection status of the plants. Contagious infections occurred more frequently in the shade, and in endophyte-free host plants. The latter suggests that pre-existing infections render host plants less susceptible to superinfection by choke forming strains. Although our results clearly indicate that disea... [ABSTRACT FROM AUTHOR]

Endophytes of the genus Epichloë (Clavicipitaceae, Ascomycota) are systemic symbionts of cool-season grasses. Their interactions with grass hosts may vary between mutualistic and pathogenic depending on the mode of endophyte reproduction. Sexual strains prevent flowering and seed set (choke disease) of the host and can be horizontally transmitted by ascospores, while asexual strains remain asymptomatic and are vertically transmitted through seeds. In Switzerland nearly all plants of Brachypodium sylvaticum (Huds.) P.B. are infected by Epichloeë sylvatica Leuchtmann & Schardl, but choke symptoms arc formed very rarely, and are restricted to particular locations and to a minority of plants at those locations. Earlier research has revealed that F. sylvatica is genetically differentiated into sexual and asexual subpopulations. Given the high level of infection and assuming horizontal transmission of sexual strains, multiple host infections have been predicted. In this study, 25 plants out of 63 examined by isozyme analysis were found to he infected by two or three different endophyte genotypes. In most cases endophyte genotypes appeared to he correlated with the symptom type of a particular tiller, suggesting that the fungal genome controls choke formation and that the sexual and asexual subpopulations are separated at the ramet (tiller) level rather than at the genet (plant) level. These conclusions were further supported by analyses with log-linear models of the population structure of E. sylvatica at four locations where choke symptoms were present. These analyses also revealed a geographic structure in the asexual subpopulation hut not in the sexual subpopulation which could be caused by the different dispersal ranges of their propagules. The rare occurrence of sexually reproducing strains and the dominance of a single genotype in asymptomatic plant populations may be explained by the colonization history of B. sylvaticum and its endophyte in Switzerland. [ABSTRACT FROM AUTHOR]