Co-existing grass species have distinctive arbuscular mycorrhizal communities

Arbuscular mycorrhizal (AM) fungi are biotrophic symbionts colonizing the majority of land plants, and are of major importance in plant nutrient supply. Their diversity is suggested to be an important determinant of plant community structure, but the influence of host-plant and environmental factors on AM fungal community in plant roots is poorly documented. Using the terminal restriction fragment length polymorphism (T-RFLP) strategy, the diversity of AM fungi was assessed in 89 roots of three grass species (Agrostis capillaris, Festuca rubra, Poa pratensis) that co-occurred in the same plots of a field experiment. The impact of different soil amendments (nitrogen, lime, nitrogen and lime) and insecticide application on AM fungal community was also studied. The level of diversity found in AM fungal communities using the T-RFLP strategy was consistent with previous studies based on clone libraries. Our results clearly confirm that an AM fungal host-plant preference exists, even between different grass species. AM communities colonizing A. capillaris were statistically different from the others (P < 0.05). Although grass species evenness changed in amended soils, AM fungal community composition in roots of a given grass species remained stable. Conversely, in plots where insecticide was applied, we found higher AM fungal diversity and, in F. rubra roots, a statistically different AM fungal community.     

Diversity and persistence of arbuscular mycorrhizas in a low-Arctic meadow habitat

Little is known about the ecology and diversity of arbuscular mycorrhizal (AM) fungi in Arctic ecosystems. Here, the diversity and composition of the AM fungal community and its response to host plant community composition were studied in a low-Arctic meadow habitat. The natural vegetation in two low-Arctic meadow sites was manipulated. Plots with natural vegetation, monoculture and no vegetation were established. Seeds of Solidago virgaurea were sown into the plots and the AM fungal community in the seedling roots was analysed using the terminal restriction fragment length polymorphism (T-RFLP) method. The vegetation manipulation treatments affected the community composition but not the diversity of AM fungi found in S. virgaurea roots. The diversity of AM fungi was higher in S. virgaurea roots in the site with naturally higher plant species diversity. These results show that AM fungi in low-Arctic meadows are able to survive for a period of 2 yr without a host plant. This ability buffers the AM fungal community against short-term changes in host plant community composition and diversity.     

Links between tree species, symbiotic fungal diversity and ecosystem functioning in simplified tropical ecosystems

We studied the relationships among plant and arbuscular mycorrhizal (AM) fungal diversity, and their effects on ecosystem function, in a series of replicate tropical forestry plots in the La Selva Biological Station, Costa Rica. Forestry plots were 12 yr old and were either monocultures of three tree species, or polycultures of the tree species with two additional understory species. Relationships among the AM fungal spore community, host species, plant community diversity and ecosystem phosphorus-use efficiency (PUE) and net primary productivity (NPP) were assessed. Analysis of the relative abundance of AM fungal spores found that host tree species had a significant effect on the AM fungal community, as did host plant community diversity (monocultures vs polycultures). The Shannon diversity index of the AM fungal spore community differed significantly among the three host tree species, but was not significantly different between monoculture and polyculture plots. Over all the plots, significant positive relationships were found between AM fungal diversity and ecosystem NPP, and between AM fungal community evenness and PUE. Relative abundance of two of the dominant AM fungal species also showed significant correlations with NPP and PUE. We conclude that the AM fungal community composition in tropical forests is sensitive to host species, and provide evidence supporting the hypothesis that the diversity of AM fungi in tropical forests and ecosystem NPP covaries.     

Arbuscular mycorrhizal communities in tropical forests are affected by host tree species and environment

Arbuscular mycorrhizal (AM) fungi are mutualists with plant roots that are proposed to enhance plant community diversity. Models indicate that AM fungal communities could maintain plant diversity in forests if functionally different communities are spatially separated. In this study we assess the spatial and temporal distribution of the AM fungal community in a wet tropical rainforest in Costa Rica. We test whether distinct fungal communities correlate with variation in tree life history characteristics, with host tree species, and the relative importance of soil type, seasonality and rainfall. Host tree species differ in their associated AM fungal communities, but differences in the AM community between hosts could not be generalized over life history groupings of hosts. Changes in the relative abundance of a few common AM fungal species were the cause of differences in AM fungal communities for different host tree species instead of differences in the presence and absence of AM fungal species. Thus, AM fungal communities are spatially distinguishable in the forest, even though all species are widespread. Soil fertility ranging between 5 and 9 Mg/ha phosphorus did not affect composition of AM fungal communities, although sporulation was more abundant in lower fertility soils. Sampling soils over seasons revealed that some AM fungal species sporulate profusely in the dry season compared to the rainy season. On one host tree species sampled at two sites with vastly different rainfall, relative abundance of spores from Acaulospora was lower and that of Glomus was relatively higher at the site with lower and more seasonal rainfall.     

Host plant species effects on arbuscular mycorrhizal fungal communities in tallgrass prairie

Symbiotic associations between plants and arbuscular mycorrhizal (AM) fungi are ubiquitous in many herbaceous plant communities and can have large effects on these communities and ecosystem processes. The extent of species-specificity between these plant and fungal symbionts in nature is poorly known, yet reciprocal effects of the composition of plant and soil microbe communities is an important assumption of recent theoretical models of plant community structure. In grassland ecosystems, host plant species may have an important role in determining development and sporulation of AM fungi and patterns of fungal species composition and diversity. In this study, the effects of five different host plant species [Poa pratensis L., Sporobolus heterolepis (A. Gray) A. Gray, Panicum virgatum L., Baptisia bracteata Muhl. ex Ell., Solidago missouriensis Nutt.] on spore communities of AM fungi in tallgrass prairie were examined. Spore abundances and species composition of fungal communities of soil samples collected from patches within tallgrass prairie were significantly influenced by the host plant species that dominated the patch. The AM fungal spore community associated with B. bracteata showed the highest species diversity and the fungi associated with Pa. virgatum showed the lowest diversity. Results from sorghum trap cultures using soil collected from under different host plant species showed differential sporulations of AM fungal species. In addition, a greenhouse study was conducted in which different host plant species were grown in similar tallgrass prairie soil. After 4 months of growth, AM fungal species composition was significantly different beneath each host species. These results strongly suggest that AM fungi show some degree of host-specificity and are not randomly distributed in tallgrass prairie. The demonstration that host plant species composition influences AM fungal species composition provides support for current feedback models predicting strong regulatory effects of soil communities on plant community structure. Differential responses of AM fungi to host plant species may also play an important role in the regulation of species composition and diversity in AM fungal communities. Received: 29 January 1999 / Accepted: 20 October 1999     

Complexity of Semiarid Gypsophilous Shrub Communities Mediates the AMF Biodiversity at the Plant Species Level

The community composition of arbuscular mycorrhizal fungi (AMF) was analyzed in roots of Gypsophila struthium growing in gypsum soils under semiarid conditions. In order to investigate the effect of plant community degradation on the AMF biodiversity at the single species level, on the basis of the plant community complexity level, we selected four areas affected by degradation and shrub species spatial heterogeneity. The AM fungal community colonizing G. struthium was investigated from the morphological and molecular points of view. All plants were well colonized and showed a high level of infective AM propagules. Roots were analyzed by polymerase chain reaction, restriction fragment length polymorphism screening, and sequence analyses of the ribosomal DNA small subunit region. Four AM fungal types were identified and clustered into the AM fungal family: Glomeraceae, Glomus being the only taxon present. One fungal type was present in all the selected areas. Two fungal types are distinct from any previously published sequences and could be specific to gypsum soils. The chemical–physical properties of the soil were not correlated with the AMF diversity in roots. Our data show vegetation cover complexity-dependent differences in the AM fungal community composition.     

Defoliation effects on the community structure of arbuscular mycorrhizal fungi based on 18S rDNA sequences

The effects of defoliation on arbuscular mycorrhizal (AM) associations in the field were investigated in terms of the community structure of AM fungi colonizing roots of grassland plants; the carbohydrate balance of the host plants was also determined. We focused on two plant species dominating Japanese native grasslands: the grazing-intolerant species Miscanthus sinensis and the grazing-tolerant species Zoysia japonica. Community structures of AM fungi were determined from 18S rRNA gene sequences. The dominant fungal group in both plant species was the Glomus clade, which was classified into several phylogenetic groups based on genetic distances and topology. In Miscanthus roots, the Glomus-Ab, Glomus-Ac, and Glomus-Ad groups were detected almost equally. In Zoysia roots, the Glomus-Ab group was dominant. Defoliation effects on the community structure of AM fungi differed between the plant species. In Miscanthus roots, the percentage of root length colonized (%RLC) by the Glomus-Ac and Glomus-Ad groups was significantly reduced by defoliation treatment. On the other hand, AM fungal group composition in Zoysia roots was unaffected by defoliation except on the last sampling date. Decreased %RLC by Glomus-Ac and Glomus-Ad coincided with decreased non-structural carbohydrate (NSC) levels in host plants; also, significant positive correlations were found between the %RLC and some NSC levels. On the other hand, the %RLC by Glomus-Ab in both plant species was unaffected by the NSC level. These results suggest that AM fungal groups have different carbohydrate requirements from host plants.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00572-003-0286-x     

Wheat cultivars form distinctive communities of root-associated arbuscular mycorrhiza in a conventional agroecosystem

Background and aims

The effect of plant species on their root-associated arbuscular mycorrhizal (AM) fungi is well studied, but how this effect operates at the cultivar level remains poorly understood. This study investigates how wheat cultivars shape their AM fungal communities.

Methods

Twenty-one new wheat cultivars were traditionally cultivated in a dryland of northwestern China, and their agronomic traits, soil characteristics and the abundance and community composition of AM fungi were measured.

Results

Both spore community in soils and AM fungal phylotypes inside roots were significantly influenced by cultivar even though hyphal abundance, spore density and AM fungal diversity were similar across cultivars. Three out of 16 AM fungal phylotypes interacted with most cultivars, whilst some phylotypes preferred to colonize cultivars with similar agronomic traits. Six wheat cultivars, all which had hosted 6 AM fungal phylotypes, seemed to be generalists. Nestedness analysis and stochastic model fitting revealed that the AM fungal communities colonizing roots were codetermined by deterministic and stochastic processes.

Conclusions

A complex pattern of cultivar-AM fungal interactions was observed in this study, and our results highlight that the host effect on the community assembly of AM fungi could be operating on the level of plant cultivar.     

Fungal root colonization responses in natural grasslands after long-term exposure to elevated atmospheric CO2

Arbuscular mycorrhizae, ubiquitous mutualistic symbioses between plant roots and fungi in the order Glomales, are believed to be important controllers of plant responses to global change, in particular to elevated atmospheric CO₂. In order to test if any effects on the symbiosis can persist after long-term treatment, we examined root colonization by arbuscular mycorrhizal (AM) and other fungi of several plant species from two grassland communities after continuous exposure to elevated atmospheric CO₂ for six growing seasons in the field. For plant species from both a sandstone and a serpentine annual grassland there was evidence for changes in fungal root colonization, with changes occurring as a function of plant host species. We documented decreases in percentage nonmycorrhizal fungal root colonization in elevated CO₂ for several plant species. Total AM root colonization (%) only increased significantly for one out of the five plant species in each grassland. However, when dividing AM fungal hyphae into two groups of hyphae (fine endophyte and coarse endophyte), we could document significant responses of AM fungi that were hidden when only total percentage colonization was measured. We also documented changes in elevated CO₂ in the percentage of root colonized by both AM hyphal types simultaneously. Our results demonstrate that changes in fungal root colonization can occur after long-term CO₂ enrichment, and that the level of resolution of the study of AM fungal responses may have to be increased to uncover significant changes to the CO₂ treatment. This study is also one of the first to document compositional changes in the AM fungi colonizing roots of plants grown in elevated CO₂. Although it is difficult to relate the structural data directly to functional changes, possible implications of the observed changes for plant communities are discussed.     

Diversity and composition of ectomycorrhizal community on seedling roots: the role of host preference and soil origin

As the main source of inocula, ectomycorrhizal (ECM) fungal propagules are critical for root colonization and seedling survival in deforested areas. It is essential to know factors that may affect the diversity and composition of ECM fungal community on roots of seedlings planted in deforest areas during reforestation. We quantitatively evaluated the effect of host plant and soil origin on ECM fungal propagule community structure established on roots of Castanopsis fargesii, Lithocarpus harlandii, Pinus armandii, and Pinus massoniana growing in soils from local natural forests and from sites deforested by clear-cut logging in the 1950s and 1960s. ECM root tips were sampled in April, July, and October of 2006, and ECM fungal communities were determined using ECM root morphotyping, internal transcribed spacer (ITS)-RFLP, and ITS sequencing. A total of 36 ECM fungal species were observed in our study, and species richness varied with host species and soil origin. Decreased colonization rates were found in all host species except for L. harlandii, and reduced species richness was found in all host species except for P. armandii in soil from the deforested site, which implied the great changes in ECM fungal community composition. Our results showed that 33.3% variance in ECM fungal community composition could be explained by host plant species and 4.6% by soil origin. Results of indicator species analysis demonstrated that 14 out of 19 common ECM fungal species showed significant preference to host plant species, suggesting that the host preference of ECM fungi was one of the most important mechanisms in structuring ECM fungal community. Accordingly, the host plant species should be taken into account in the reforestation of deforested areas due to the strong and commonly existed host preference of ECM fungi.     

Alien plants associate with widespread generalist arbuscular mycorrhizal fungal taxa: evidence from a continental‐scale study using massively parallel 454 sequencing

Aim The biogeography of arbuscular mycorrhizal (AM) fungi is poorly understood, and consequently the potential of AM fungi to determine plant distribution has been largely overlooked. We aimed to describe AM fungal communities associating with a single host‐plant species across a wide geographical area, including the plant’s native, invasive and experimentally introduced ranges. We hypothesized that an alien AM plant associates primarily with the geographically widespread generalist AM fungal taxa present in a novel range. Location Europe, China. Methods We transplanted the palm Trachycarpus fortunei into nine European sites where it does not occur as a native species, into one site where it is naturalized (Switzerland), and into one glasshouse site. We harvested plant roots after two seasons. In addition, we sampled palms at three sites in the plant’s native range (China). Roots were subjected to DNA extraction, polymerase chain reaction (PCR) and 454 sequencing of AM fungal sequences. We analysed fungal communities with non‐metric multidimensional scaling (NMDS) ordination and cluster analysis and studied the frequency of geographically widespread fungal taxa with log‐linear analysis. We compared fungal communities in the roots of the palm with those in resident plants at one site in the introduced range (Estonia) where natural AM fungal communities had previously been studied. Results We recorded a total of 73 AM fungal taxa. AM fungal communities in the native and introduced ranges differed from one another, while those in the invasive range contained taxa present in both other ranges. Geographically widespread AM fungal taxa were over‐represented in palm roots in all regions, but especially in the introduced range. At the Estonian site, the palm was colonized by the same community of widespread AM fungal taxa as associate with resident habitat‐generalist plants; by contrast, resident forest‐specialist plants were colonized by a diverse community of widespread and other AM fungal taxa. Main conclusions AM fungal communities in the native, invasive and experimentally introduced ranges varied in taxonomic composition and richness, but they shared a pool of geographically widespread, non‐host‐specific taxa that might support the invasion of a generalist alien plant. Our dataset provides the first geographical overview of AM taxon distributions obtained using a single host‐plant species.     

Relationship between assemblages of mycorrhizal fungi and bacteria on grass roots

Soils support an enormous microbial diversity, but the ecological drivers of this diversity are poorly understood. Interactions between the roots of individual grass species and the arbuscular mycorrhizal (AM) fungi and bacteria in their rhizoplane were studied in a grazed, unimproved upland pasture. Individual root fragments were isolated from soil cores, DNA extracted and used to identify plant species and assess rhizoplane bacterial and AM fungal assemblages, by amplifying part of the small-subunit ribosomal RNA gene, followed by terminal restriction fragment length polymorphism analysis. For the first time we showed that AM fungal and bacterial assemblages are related in situ and that this relationship occurred at the community level. Principal coordinate analyses of the data show that the AM fungi were a major factor determining the bacterial assemblage on grass roots. We also report a strong influence of the composition of the plant community on AM fungal assemblage. The bacterial assemblage was also influenced by soil pH and was spatially structured, whereas AM fungi were influenced neither by the bacteria nor by soil pH. Our study shows that linkages between plant roots and their microbial communities exist in a complex web of interactions that act at individual and at community levels, with AM fungi influencing the bacterial assemblage, but not the other way round.     

Local adaptation to soil hypoxia determines the structure of an arbuscular mycorrhizal fungal community in roots from natural CO₂ springs

The processes responsible for producing and maintaining the diversity of natural arbuscular mycorrhizal (AM) fungal communities remain largely unknown. We used natural CO(2) springs (mofettes), which create hypoxic soil environments, to determine whether a long-term, directional, abiotic selection pressure could change AM fungal community structure and drive the selection of particular AM fungal phylotypes. We explored whether those phylotypes that appear exclusively in hypoxic soils are local specialists or widespread generalists able to tolerate a range of soil conditions. AM fungal community composition was characterized by cloning, restriction fragment length polymorphism typing, and the sequencing of small subunit rRNA genes from roots of four plant species growing at high (hypoxic) and low (control) geological CO(2) exposure. We found significant levels of AM fungal community turnover (β diversity) between soil types and the numerical dominance of two AM fungal phylotypes in hypoxic soils. Our results strongly suggest that direct environmental selection acting on AM fungi is a major factor regulating AM fungal communities and their phylogeographic patterns. Consequently, some AM fungi are more strongly associated with local variations in the soil environment than with their host plant's distribution.     

Interactive effects of root endophytes and arbuscular mycorrhizal fungi on an experimental plant community

Plant-soil microbial interactions have moved into focus as an important mechanism for understanding plant coexistence and composition of communities. Both arbuscular mycorrhizal (AM) as well as other root endophytic fungi co-occur in plant roots, and therefore have the potential to influence relative abundances of plant species in local assemblages. However, no study has experimentally examined how these key root endosymbiont groups might interact and affect plant community composition. Here, using an assemblage of five plant species in mesocosms in a fully factorial experiment, we added an assemblage of AM fungi and/or a mixture of root endophytic fungal isolates, all obtained from the same grassland field site. The results demonstrate that the AM fungi and root endophytes interact to affect plant community composition by changing relative species abundance, and consequently aboveground productivity. Our study highlights the need to explicitly consider interactions of root-inhabiting fungal groups in studies of plant assemblages.     

Differentiating between effects of invasion and diversity: impacts of aboveground plant communities on belowground fungal communities

Exotic plant species can affect soil microbial communities with the potential for community and ecosystem feedbacks. Yet, separating the effects of exotics from confounded changes in plant community diversity still remains a challenge. We focused on how plant diversity and native or exotic life history affected root fungi because of their significant roles in community and ecosystem processes. Specifically, we examined how fungi colonizing plant roots were affected by plant richness (one, two or four species) replicated across a range of plant community mixtures (natives, exotics, native-exotic mixtures). Fungal biomass inside roots was affected independently by plant richness and mixture, while root fungal community composition was affected only by plant richness. Extraradical networks also increased in size with plant richness. By contrast, plant biomass was a function of plant mixture, with natives consistently smaller than exotics and native-exotic mixtures intermediate. Plant invasions may have an impact on the belowground community primarily through their effects on diversity, at least in the short-term. Disentangling the effects of diversity and invasion on belowground microbial communities can help us to understand both the controllers of belowground resilience and mechanisms of successful colonization and spread of exotic plants.     

Patterns of diversity and adaptation in Glomeromycota from three prairie grasslands

Arbuscular mycorrhizal (AM) fungi are widespread root symbionts that often improve the fitness of their plant hosts. We tested whether local adaptation in mycorrhizal symbioses would shape the community structure of these root symbionts in a way that maximizes their symbiotic functioning. We grew a native prairie grass (Andropogon gerardii) with all possible combinations of soils and AM fungal inocula from three different prairies that varied in soil characteristics and disturbance history (two native prairie remnants and one recently restored). We identified the AM fungi colonizing A. gerardii roots using PCR amplification and cloning of the small subunit rRNA gene. We observed 13 operational taxonomic units (OTUs) belonging to six genera in three families. Taxonomic richness was higher in the restored than the native prairies with one member of the Gigaspora dominating the roots of plants grown with inocula from native prairies. Inoculum source and the soil environment influenced the composition of AM fungi that colonized plant roots. Correspondingly, host plants and AM fungi responded significantly to the soil–inoculum combinations such that home fungi often had the highest fitness and provided the greatest benefit to A. gerardii. Similar patterns were observed within the soil–inoculum combinations originating from two native prairies, where five sequence types of a single Gigaspora OTU were virtually the only root colonizers. Our results indicate that indigenous assemblages of AM fungi were adapted to the local soil environment and that this process occurred both at a community scale and at the scale of fungal sequence types within a dominant OTU.     

Temporal dynamics of arbuscular mycorrhizal fungi colonizing roots of representative shrub species in a semi-arid Mediterranean ecosystem

Arbuscular mycorrhizal (AM) symbiosis plays an important role in improving plant fitness and soil quality, particularly in fragile and stressed environments, as those in certain areas of Mediterranean ecosystems. AM fungal communities are usually affected by dynamic factors such as the plant community structure and composition, which in turn are imposed by seasonality. For this reason, a one-year-round time-course trial was performed by sampling the root system of two representative shrubland species (Rosmarinus officinalis and Thymus zygis) within a typical Mediterranean ecosystem from the Southeast of Spain. The 18S rDNA gene, of the AM fungal community in roots, was subjected to PCR-SSCP, sequencing, and phylogenetic analysis. Forty-three different AM fungal sequence types were found which clustered in 16 phylotypes: 14 belonged to the Glomeraceae and two to the Diversisporaceae. Surprisingly, only two of these phylotypes were related with sequences of morphologically defined species: Glomus intraradices and Glomus constrictum. Significant differences were detected for the relative abundance of some phylotypes while no effects were found for the calculated diversity indices. These results may help to design efficient mycorrhizal-based revegetation programs for this type of ecosystems.     

Molecular diversity of arbuscular mycorrhizal fungi and patterns of host association over time and space in a tropical forest

We have used molecular techniques to investigate the diversity and distribution of the arbuscular mycorrhizal (AM) fungi colonizing tree seedling roots in the tropical forest on Barro Colorado Island (BCI), Republic of Panama. In the first year, we sampled newly emergent seedlings of the understory treelet Faramea occidentalis and the canopy emergent Tetragastris panamensis, from mixed seedling carpets at each of two sites. The following year we sampled surviving seedlings from these cohorts. The roots of 48 plants were analysed using AM fungal-specific primers to amplify and clone partial small subunit (SSU) ribosomal RNA gene sequences. Over 1300 clones were screened for random fragment length polymorphism (RFLP) variation and 7% of these were sequenced. Compared with AM fungal communities sampled from temperate habitats using the same method, the overall diversity was high, with a total of 30 AM fungal types identified. Seventeen of these types have not been recorded previously, with the remainder being similar to types reported from temperate habitats. The tropical mycorrhizal population showed significant spatial heterogeneity and nonrandom associations with the different hosts. Moreover there was a strong shift in the mycorrhizal communities over time. AM fungal types that were dominant in the newly germinated seedlings were almost entirely replaced by previously rare types in the surviving seedlings the following year. The high diversity and huge variation detected across time points, sites and hosts, implies that the AM fungal types are ecologically distinct and thus may have the potential to influence recruitment and host composition in tropical forests.     

Contrasting root associated fungi of three common oak-woodland plant species based on molecular identification: host specificity or non-specific amplification?

An increasingly popular approach used to identify arbuscular mycorrhizal (AM) fungi in planta is to amplify a portion of AM fungal small subunit ribosomal DNA (SSU-rDNA) from whole root DNA extractions using the primer pair AM1-NS31, followed by cloning and sequencing. We used this approach to study the AM fungal community composition of three common oak-woodland plant species: a grass (Cynosurus echinatus), blue oak (Quercus douglasii), and a forb (Torilis arvensis). Significant diversity of AM fungi were found in the roots of C. echinatus, which is consistent with previous studies demonstrating a high degree of AM fungal diversity from the roots of various hosts. In contrast, clones from Q. douglasii and T. arvensis were primarily from non-AM fungi of diverse origins within the Ascomycota and Basidiomycota. This work demonstrates that caution must be taken when using this molecular approach to determine in planta AM fungal diversity if non-sequence based methods such as terminal restriction fragment length polymorphisms, denaturing gradient gel electrophoresis, or temperature gradient gel electrophoresis are used.