Arbuscular Mycorrhizal Communities Research Articles

Evidence for ecological matching of whole AM fungal communities to the local plant–soil environment

The range of ecological roles exhibited by arbuscular mycorrhizal (AM) fungi depends on functional differences among naturally occurring local assemblages of AM species. While functional differences have been demonstrated among AM fungal species and among geographic isolates of the same species, almost nothing is known about functional differences among whole communities of naturally occurring AM fungi. In the greenhouse, we reciprocally transplanted whole AM fungal communities between plant-soil systems representing a serpentine grassland and a tallgrass prairie, using as hosts two grasses common to both systems. For Sorghastrum nutans, native fungi consistently enhanced plant growth more than fungi switched from the alternate system. For Schizachyrium scoparium, foreign and native fungi promoted plant growth similarly in both the serpentine and prairie systems. Thus, the use of foreign inoculum in restoration could change the relative performance, and potentially the competitive abilities, of co-occurring plant species. Moving AM fungal inocula into foreign environments also caused changes in the taxonomic composition of the resultant spore communities, demonstrating their response to environmental influences. These results provide strong evidence for functional differences among naturally occurring AM communities and suggest that a particular AM fungal community may be better matched ecologically to its local habitat than communities taken from other locations.

Order of plant host establishment alters the composition of arbuscular mycorrhizal communities

The causes of local diversity and composition remain a central question in community ecology. Numerous studies have attempted to understand community assembly, both within and across trophic levels. However, little is known about how community assembly aboveground influences soil microbial communities belowground. We hypothesized that plant establishment order can affect the community of arbuscular mycorrhizal fungi (AMF) in roots, with the strength of this effect dependent on both host plant identity and neighboring plant identity. Such priority effects of plants on AMF may act through host-specific filters of the initial species pool that limit the available pool for plants that established second. In a greenhouse experiment with four plant hosts, we found that the strength of the priority effect on AMF communities reflected both host plant characteristics and interactions between host and neighbor plant species, consistent with differential host specificity among plants. These patterns were independent of plant biomass and root colonization. Functional studies of AMF associated with a wide array of host plants will be required to further understand this potential driver of community dynamics.

Plant neighborhood control of arbuscular mycorrhizal community composition

Arbuscular mycorrhizal fungi (AMF) are important root symbionts that can provide benefits to plant hosts, yet we understand little about how neighboring hosts in a plant community contribute to the composition of the AMF community. We hypothesized that the composition of the plant neighborhood, including the identities of both host and neighbor, would alter AMF community composition. We tested this in a glasshouse experiment in which a native perennial grass (Nassella pulchra) and three annual grasses (Avena barbata, Bromus hordeaceaous and Vulpia microstachys) were grown in two neighborhoods: conspecific monocultures and heterospecific perennial-annual mixtures. To identify AMF taxa colonizing plant roots, we used a combination of terminal restriction fragment length polymorphism and cloning. Both host and neighbor were important in structuring AMF communities. Unique AMF communities were associated with each plant host in monoculture. In heterospecific neighborhoods, the annual neighbors V. microstachys, A. barbata, and B. hordeaceus influenced N. pulchra AMF in different ways (synergistic, controlling, or neutral) and the reciprocal effect was not always symmetric. Our findings support a community approach to AMF studies, which can be used to increase our understanding of processes such as invasion and succession.

Molecular analysis of arbuscular mycorrhizal community structure and spores distribution in tree-based intercropping and forest systems

Due to their potential to enhance soil biodiversity and fertility, tree-based intercropping (TBI) systems are recognized as promising agrotechnologies. However, few studies have examined the effects of TBI on the diversity of arbuscular mycorrhizal (AM) fungi. To investigate this aspect, and to compare TBI with a more typical forest plantation (FO) system, a field experiment using soybean ( Glycine max (L.) Merr. cv. SO3-W4) and poplar ( Populus nigra L. × P. maximowiczii cv. A. Henry ‘Max 5’) was conducted on an arable field site in southwestern Québec. The species of AM fungi colonizing the roots were assessed by PCR-RFLP and DNA sequence analyses of SSU rRNA genes. A total of 13 different phylotypes were identified. Phylogenetic analyses demonstrated that these belonged to the genera Glomus (11), Acaulospora (1) and Scutellospora (1). The AM fungal diversity, as expressed by the Shannon–Wiener indices, were 0.82 ± 0.08 for the soybean and 0.70 ± 0.11 for poplar under the TBI system, and 0.53 ± 0.08 for poplar under the FO system. The study also investigated the distribution of AM fungal spores in the soil, and this differed in the two systems. Under the TBI system, spore abundance increased with increasing distance from the poplar trunk, while it remained relatively constant under the FO system. Our results also show that neighbouring trees and soybean plants hosted different AM fungal communities, suggesting that TBI systems may enhance AM fungal richness compared to monocultures.

Shifting the balance from qualitative to quantitative analysis of arbuscular mycorrhizal communities in field soils

Population studies of arbuscular mycorrhizal fungi (AMF) have traditionally been achieved by indirect analyses of soil-borne spore populations. These studies are not necessarily reflective of populations of AMF within the roots. Advances in molecular biology have revolutionized the analysis of fungal populations colonizing roots and forming mycorrhizas. Initially these studies were qualitative and reported presence or absence of particular AMF species in soils or in roots for comparison between different environments. More recently, the methodology has developed for direct quantification of AMF within roots. Quantitative PCR provides the means to study spatial distribution and individual quantification of AMF in mixed communities over time. In this review, we discuss the progress and application of indirect, direct and finally quantitative methodologies for studying arbuscular mycorrhizal communities. We conclude that the molecular tools now exist to quantitatively analyse the effect of environment, management or inoculation of soils on AMF communities within roots.

Influence of commercial inoculation with Glomus intraradices on the structure and functioning of an AM fungal community from an agricultural site

The use of commercial arbuscular mycorrhizal (AM) inoculants is growing. However, we know little about how resident AM communities respond to inoculations under different soil management conditions. The objective of this study was to simulate the application of a commercial AM fungal inoculant of Glomus intraradices to soil to determine whether the structure and functioning of that soil’s resident AM community would be affected. The effects of inoculation were investigated over time under disturbed or undisturbed soil conditions. We predicted that the introduction of an infective AM fungus, such as G. intraradices, would have greater consequences in disturbed soil. Using a combination of molecular (terminal restriction length polymorphism analysis based on the large subunit of the rRNA gene) and classical methods (AM fungal root colonization and P nutrition) we found that, contrary to our prediction, adding inoculant to soil containing a resident AM fungal community does not necessarily have an impact on the structure of that community either under disturbed or undisturbed conditions. However, we found evidence of positive effects of inoculation on plant nutrition under disturbed conditions, suggesting that the inoculant interacted, directly or indirectly, with the resident AM fungi. The inoculant significantly improved the P content of the host but only in presence of the resident AM fungal community. In contrast to inoculation, soil disturbance had a significant negative impact on species richness of AM fungi and influenced the AM fungal community composition as well as its functioning. Thus, we conclude that soil disturbance may under certain conditions have greater consequences for the structure of resident AM fungal communities in agricultural soils than commercial AM fungal inoculations with G. intraradices.

Compatible host/mycorrhizal fungus combinations for micropropagated sea oats: II. Field evaluation

Sea oats (Uniola paniculata L.) are the dominant plant in the pioneer coastal dunes of Florida and are widely used for dune restoration. DNA analysis has revealed significant ecotypic variation among Atlantic and Gulf coast populations of sea oats, but little is known about the diversity of the arbuscular mycorrhizal (AM) communities present in the dune systems. In a prior greenhouse study, we evaluated the functional diversity that exists among the AM fungal communities from divergent Florida dunes and selected effective host/AM fungus combinations for further study. The objective of this study was to evaluate the effect of these compatible combinations on the growth of sea oats planted at Anastasia State Recreation Area (AN) on the Atlantic coast and St. George Island State Park (SG) on the Gulf coast. Micropropagated sea oats from each site were inoculated with AM fungal communities also from AN and SG or a microbial filtrate control. The complete factorial of treatment combinations were grown in the greenhouse for 8 weeks and outplanted to the AN and SG field sites. After 1 year, root colonization was evaluated, and after 2 years, root colonization, shoot and root dry masses, and shoot- and root-P contents were determined. Overall, sea oats planted at AN had greater percent root colonization, shoot dry mass, and shoot-P content than those planted at SG. At AN, the local sea oat ecotype responded more to the fungal community from the same site relative to shoot dry mass and shoot-P content. At SG, the local fungal community produced larger plants with greater P content regardless of the origin of the host. We conclude that sea oat productivity is responsive to AM fungal ecotype as well as host ecotype, and fungal origin should therefore be taken into account when planning sea oat plantings on coastal dunes.

A DGGE-cloning method to characterize arbuscular mycorrhizal community structure in soil

Although arbuscular mycorrhizal fungi (AMF) are crucial for ecosystem functioning, characterizing AMF community structure in soil is challenging. In this study, nested polymerase chain reaction (PCR) and denaturing gradient gel electrophoresis (DGGE) were combined with cloning of fungal 18S ribosomal gene fragments for the rapid comparison of AMF community structure in soil. Reference AMF isolates, representing four major genera of AMF, were used to develop the method. Sequential amplification of 18S rDNA fragments by nested PCR using primer pairs AM1-NS31 and Glo1-NS31GC followed by DGGE analysis yielded a high-resolution band profile. In parallel, 18S rDNA fragment clone libraries were constructed and clones screened by DGGE. Sequence identity was inferred by matching the electrophoretic mobility of the sample fingerprint bands to that of bands from individual clones. The effectiveness of this approach was tested on soil samples from different ecosystems, yielding reproducible, complex DGGE band patterns specific to each site. The coupling of PCR–DGGE with clone library analysis provides a robust, reliable, and precise means to characterize AMF community structure in soils.

Glomalin in Ecosystems

Glomalin is an abundant soil glycoprotein that could sequester substantial amounts of C and N on a global scale. Nevertheless, major controls over the distribution of this compound in ecosystems are little understood. We reviewed shifts in the production, decomposition, and standing stocks of soil glomalin in response to arbuscular mycorrhizal (AM) communities, plant dynamics, inorganic resources (e.g., N, P, and atmospheric CO2), and land use regimes. We also conducted a literature survey of glomalin concentrations from 22 ecosystems to test for relationships between standing stocks of glomalin and net primary productivity (NPP) and AM abundance. Globally, glomalin stocks were positively correlated with NPP but not with AM abundance. Rates of NPP may impose an upper bound for C available to AM fungi for glomalin construction. Availability of C to AM fungi appears to consistently influence glomalin dynamics in individual studies as well. Glomalin stocks are often greater where AM host plants are more abundant or are more effective hosts for the fungi (implying that more C is allocated to the fungi). In addition, elevated CO2 frequently increases glomalin concentrations in soil. Other inorganic resources and land use change produce inconsistent effects on glomalin stocks. Overall, glomalin dynamics appear to be linked to C dynamics, and this relationship may allow us to better predict the contribution of glomalin to ecosystem feedbacks on global change.

Testing nickel tolerance of Sorghastrum nutans and its associated soil microbial community from serpentine and prairie soils

Ecotypes of Sorghastrum nutans from a naturally metalliferous serpentine grassland and the tallgrass prairie were assessed for Ni tolerance and their utility in remediation of Ni-polluted soils. Plants were inoculated with serpentine arbuscular mycorrhizal (AM) root inoculum or whole soil microbial communities, originating from either prairie or serpentine, to test their effects on plant performance in the presence of Ni. Serpentine plants had marginally higher Ni tolerance as indicated by higher survival. Ni reduced plant biomass and AM root colonization for both ecotypes. The serpentine AM fungi and whole microbial community treatments decreased plant biomass relative to uninoculated plants, while the prairie microbial community had no effect. Differences in how the soil communities affect plant performance were not reflected in patterns of root colonization by AM fungi. Thus, serpentine plants may be suited for reclamation of Ni-polluted soils, but AM fungi that occur on serpentine do not improve Ni tolerance.

Response of arbuscular mycorrhizal communities to seasonality and long-term phosphate application

Response of arbuscular mycorrhizal communities to seasonality and long-term phosphate application

Glomus intraradices dominates arbuscular mycorrhizal communities in a heavy textured agricultural soil

Arbuscular mycorrhizal fungal (AMF) spore communities were surveyed in a long-term field fertilization experiment in Switzerland, where different amounts of phosphorus (P) were applied to soil. Plots receiving no P as well as plots systematically fertilized in excess to plant needs for 31 years were used to test the hypothesis that application of P fertilizer changes the composition and diversity of AMF communities. AMF spores were isolated from the field soil, identified, and counted so as to quantify the effect of P fertilization on AMF spore density, composition, and diversity. Trap cultures were established from field soil with four host plants (sunflower, leek, maize, and Crotalaria grahamiana), and the spore communities were then analyzed in substrate samples from the pots. Altogether, nine AMF species were detected in the soil. No evidence has been acquired for effect of P fertilization on spore density, composition, and diversity of AMF in both the field soil and in trap cultures. On the other hand, we observed strong effect of crop plant species on spore densities in the soil, the values being lowest under rapeseed and highest under Phacelia tanacetifolia covercrop. The identity of plant species in trap pots also significantly affected composition and diversity of associated AMF communities, probably due to preferential establishment of symbiosis between certain plant and AMF species. AMF spore communities under mycorrhizal host plants (wheat and Phacelia in the fields and four host plant species in trap pots) were dominated by a single AMF species, Glomus intraradices. This resulted in exceptionally low AMF spore diversity that seems to be linked to high clay content of the soil.

Spatial structure in soil chemical and microbiological properties in an upland grassland

We characterised the spatial structure of soil microbial communities in an unimproved grazed upland grassland in the Scottish Borders. A range of soil chemical parameters, cultivable microbes, protozoa, nematodes, phospholipid fatty acid (PLFA) profiles, community-level physiological profiles (CLPP), intra-radical arbuscular mycorrhizal community structure, and eubacterial, actinomycete, pseudomonad and ammonia-oxidiser 16S rRNA gene profiles, assessed by denaturing gradient gel electrophoresis (DGGE) were quantified. The botanical composition of the vegetation associated with each soil sample was also determined. Geostatistical analysis of the data revealed a gamut of spatial dependency with diverse semivariograms being apparent, ranging from pure nugget, linear and non-linear forms. Spatial autocorrelation generally accounted for 40–60% of the total variance of those properties where such autocorrelation was apparent, but accounted for 97% in the case of nitrate-N. Geostatistical ranges extending from approximately 0.6–6 m were detected, dispersed throughout both chemical and biological properties. CLPP data tended to be associated with ranges greater than 4.5 m. There was no relationship between physical distance in the field and genetic similarity based on DGGE profiles. However, analysis of samples taken as close as 1 cm apart within a subset of cores suggested some spatial dependency in community DNA-DGGE parameters below an 8 cm scale. Spatial correlation between the properties was generally weak, with some exceptions such as between microbial biomass C and total N and C. There was evidence for scale-dependence in the relationships between properties. PLFA and CLPP profiling showed some association with vegetation composition, but DGGE profiling did not. There was considerably stronger association between notional sheep urine patches, denoted by soil nutrient status, and many of the properties. These data demonstrate extreme spatial variation in community-level microbiological properties in upland grasslands, and that despite considerable numeric ranges in the majority of properties, overarching controlling factors were not apparent.

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.

Temporal variation in the arbuscular mycorrhizal communities colonising seedlings in a tropical forest

Abstract In order to investigate temporal variation in the arbuscular mycorrhizal (AM) fungal community in a tropical forest in the Republic of Panama, seedlings of the canopy emergent Tetragastris panamensis were sampled three times over a period of 3 years. We used AM-specific primers to amplify and clone partial small subunit ribosomal RNA gene sequences. Over 550 clones were classified into 18 AM fungal types. As the seedlings matured, the fungal diversity decreased and there was a significant shift so that previously rare types replaced formerly dominant fungal types. Further, seedlings of different ages sampled at the same time point were colonised by significantly different fungal populations. Our results indicate that both time and host age may influence the mycorrhizal population.

Temporal variation in the arbuscular mycorrhizal communities colonising seedlings in a tropical forest

In order to investigate temporal variation in the arbuscular mycorrhizal (AM) fungal community in a tropical forest in the Republic of Panama, seedlings of the canopy emergent Tetragastris panamensis were sampled three times over a period of 3 years. We used AM-specific primers to amplify and clone partial small subunit ribosomal RNA gene sequences. Over 550 clones were classified into 18 AM fungal types. As the seedlings matured, the fungal diversity decreased and there was a significant shift so that previously rare types replaced formerly dominant fungal types. Further, seedlings of different ages sampled at the same time point were colonised by significantly different fungal populations. Our results indicate that both time and host age may influence the mycorrhizal population.

Arbuscular mycorrhizal community composition associated with two plant species in a grassland ecosystem.

Arbuscular mycorrhizal (AM) fungi are biotrophic symbionts colonizing about two-thirds of land plant species and found in all ecosystems. They are of major importance in plant nutrient supply and their diversity is suggested to be an important determinant of plant community composition. The diversity of the AM fungal community composition in the roots of two plant species (Agrostis capillaris and Trifolium repens) that co-occurred in the same grassland ecosystem was characterized using molecular techniques. We analysed the small subunit (SSU) ribosomal RNA gene amplified from a total root DNA extract using AM fungal-specific primers. A total of 2001 cloned fragments from 47 root samples obtained on four dates were analysed by restriction fragment length polymorphism, and 121 of them were sequenced. The diversity found was high: a total of 24 different phylotypes (groups of phylogenetically related sequences) colonized the roots of the two host species. Phylogenetic analyses demonstrate that 19 of these phylotypes belonged to the Glomaceae, three to the Acaulosporaceae and two to the Gigasporaceae. Our study reveals clearly that the AM fungal community colonizing T. repens differed from that colonizing A. capillaris, providing evidence for AM fungal host preference. In addition, our results reveal dynamic changes in the AM fungal community through time.

HETEROGENEITY IN INOCULUM POTENTIAL AND EFFECTIVENESS OF ARBUSCULAR MYCORRHIZAL FUNGI

Arbuscular mycorrhizae are symbiotic associations among glomalean fungi and plant roots that often lead to enhanced water and nutrient uptake and plant growth. We describe experiments to test whether inoculum potential of arbuscular mycorrhizal (AM) fungal communities varies spatially within a broadleaf temperate forest, and also whether there is variability in the effectiveness of AM fungal communities in enhancing seedling growth. Inoculum potential of arbuscular mycorrhizal fungi in a temperate broad-leaved forest did not vary significantly among sites. Inoculum potential, measured as the extent to which the roots of red maple seedlings that had been germinated on sterile sand and then transplanted into the forest, were colonized by AM fungi, was similar in floodplain and higher elevation sites. It was as similar under ectomycorrhizal oaks as it was under red maples and other AM tree species. It was also similar among sites with deciduous understory shrubs with arbuscular mycorrhizae (spicebush, Lindera benzoin) and those with evergreen vegetation with ericoid mycorrhizae (mountain laurel, Kalmia latifolia). Where spicebush was the dominant understory shrub, inoculum potential was greater under gaps in the canopy than within the understory. Survivorship of transplanted red maple seedlings varied significantly over sites but was not strongly correlated with measures of inoculum potential. In a greenhouse growth experiment, arbuscular mycorrhizal fungal communities obtained from tree roots from the forest had different effects on plant growth. Seedlings inoculated with roots of red maple had twice the leaf area after 10 wk of growth compared to the AM community obtained from roots of southern red oaks. Thus, although there appears to be little heterogeneity in inoculum potential in the forest, there are differences in the effectiveness of different inocula. These effects have the potential to affect tree species diversity in forests by modifying patterns of seedling recruitment.

SHIFTS IN ARBUSCULAR MYCORRHIZAL COMMUNITIES ALONG AN ANTHROPOGENIC NITROGEN DEPOSITION GRADIENT

We evaluated arbuscular mycorrhizal (AM) species diversity and abundance in nine locations along an anthropogenic nitrogen deposition gradient in coastal sage scrub (CSS) vegetation in southern California. The primary pollutants were nitrogen oxides derived from vehicular emissions. Extractable soil N on the gradient ranged from 5 to 87 μg/g during the summer months. For comparative purposes, we also assessed AM communities in nitrogen-fertilized (60 kg N·ha−1·yr−1) and unfertilized plots. Nitrogen enrichment induced a shift in AM community composition. In particular, an increasing input of nitrogen was associated with the displacement of the larger-spored species of Scutellospora and Gigaspora (due to a failure to sporulate) with a concomitant proliferation of small-spored Glomus species (e.g., Glomus aggregatum, Glomus leptotichum). A subsequent reduction in species richness and diversity (as measured by Shannon–Wiener index) accompanied eutrophication. Nitrogen enrichment also significantly reduced spo...

Shifts in Arbuscular Mycorrhizal Communities along an Anthropogenic Nitrogen Deposition Gradient

Shifts in Arbuscular Mycorrhizal Communities along an Anthropogenic Nitrogen Deposition Gradient