Role of plant relatedness in plant-soil feedback dynamics of sympatric Asclepias species.

Plants affect associated biotic and abiotic edaphic factors, with reciprocal feedbacks from soil characteristics affecting plants. These two-way interactions between plants and soils are collectively known as plant-soil feedbacks (PSFs). The role of phylogenetic relatedness and evolutionary histories have recently emerged as a potential driver of PSFs, although the strength and direction of feedbacks among sympatric congeners are not well-understood. We examined plant-soil feedback responses of Asclepias syriaca, a common clonal milkweed species, with several sympatric congeners across a gradient of increasing phylogenetic distances (A. tuberosa, A. viridis, A. sullivantii, and A. verticillata, respectively). Plant-soil feedbacks were measured through productivity and colonization by arbuscular mycorrhizal (AM) fungi. Asclepias syriaca produced less biomass in soils conditioned by the most phylogenetically distant species (A. verticillata), relative to conspecific-conditioned soils. Similarly, arbuscular mycorrhizal (AM) fungal colonization of A. syriaca roots was reduced when grown in soils conditioned by A. verticillata, compared with colonization in plants grown in soil conditioned by any of the other three Asclepias species, indicating mycorrhizal associations are a potential mechanism of observed positive PSFs. This display of differences between the most phylogenetically distant, but not close or intermediate, paring(s) suggests a potential phylogenetic threshold, although other exogenous factors cannot be ruled out. Overall, these results highlight the potential role of phylogenetic distance in influencing positive PSFs through mutualists. The role of phylogenetic relatedness and evolutionary histories have recently emerged as a potential driver of plant-soil feedbacks (PSFs), although the strength and direction of feedbacks among sympatric congeners are not well-understood. Congeneric, sympatric milkweeds typically generated positive PSFs in terms of productivity and AM fungal colonization, suggesting the low likelihood of coexistence among tested pairs, with a strength of feedback increasing as the phylogenetic distance increases.

Arbuscular mycorrhiza can be disadvantageous for weedy annuals in competition with paired perennial plants.

Arbuscular mycorrhizal (AM) fungi can support the establishment of mycotrophic plants in new environments. However, the role of mycorrhizal symbiosis in interactions between perennial and weedy annual plants is not well understood. In our current study, we examine how widespread generalist AM fungi and soil disturbance, including disturbance of AM fungal networks (CMNs), affect the performance of two late-successional perennial plants of Central Europe, Senecio jacobaea and Crepis biennis, co-occurring with weedy annual forbs, Conyza canadensis and Erigeron annuus. Although presence of weedy annual E. annuus or C. canadensis did not affect the performance of the paired perennials, AM fungi supported perennial C. biennis in competition with weedy annual E. annuus. However, this AM-aided underpinning was independent of disturbance of CMNs. Conversely, although AM fungi benefited perennial S. jacobaea, this did not affect its competitive abilities when grown with weedy annual C. canadensis. Similarly, soil disturbance, independent of AM fungal presence, improved plant tissue P and biomass production of S. jacobaea, but not its competitive abilities. Our results show AM fungi may be advantageous for perennial plants growing in competition with weedy annual plants. Therefore, maintaining healthy soils containing an abundance of AM fungi, may encourage late successional perennial plants, potentially limiting establishment of weedy annual plant species.

Assessing soil microbes that drive fairy ring patterns in temperate semiarid grasslands.

BACKGROUND: Fairy rings occur in diverse global biomes; however, there is a critical knowledge gap regarding drivers of fairy rings in grassland ecosystems. Grassland fairy rings are characterized belowground by an expanding mycelial front and aboveground by vigorous vegetation rings that develop concentrically with each growing season. We evaluated fairy ring dynamics in a field study conducted in semiarid grasslands to elucidate above- and belowground interactions driving distinct vegetation patterns. We followed this initial field investigation with a complementary greenhouse experiment, using soils collected from specific fairy ring zones (inside, ring-edge, outside) to examine plant-soil-microbial interactions under controlled conditions. We selected Leymus chinensis (a dominant grass) as our model plant species to assess the role of diverse fairy ring microbial communities on plant growth and nutrition. RESULTS: In our field study, plants on the ring-edge produced greater shoot biomass with higher concentrations of N and P, compared to plants inside the ring or adjacent (outside) controls. Soil microbial community biomarkers indicate shifts in relative microbial biomass as fairy rings expand. Inside the ring, plant roots showed greater damage from pathogenic fungi, compared to outside or ring-edge. Our greenhouse experiment confirmed that inoculation with live ring-edge soil generally promoted plant growth but decreased shoot P concentration. Inoculation with soil collected from inside the ring increased root pathogen infection and reduced shoot biomass. CONCLUSION: We propose that soil microbial activity within ring-edges promotes plant growth via mobilization of plant-available P or directed stimulation. However, as the ring expands, L. chinensis at the leading edge may increase pathogen accumulation, resulting in reduced growth at the center of the ring in subsequent growing seasons. Our results provide new insights into the plant-soil-microbial dynamics of fairy rings in grasslands, helping to elucidate these mysterious vegetation patterns.

Effects of Commercial Arbuscular Mycorrhizal Inoculants on Plant Productivity and Intra-Radical Colonization in Native Grassland: Unintentional De-Coupling of a Symbiosis?

There has been a surge in industries built on the production of arbuscular mycorrhizal (AM) fungal-based inoculants in the past few decades. This is not surprising, given the positive effects of AM fungi on plant growth and nutritional status. However, there is growing concern regarding the quality and efficacy of commercial inoculants. To assess the potential benefits and negative consequences of commercial AM fungal inoculants in grasslands, we conducted a controlled growth chamber study assessing the productivity and AM fungal root colonization of nine grassland plant species grown in grassland soil with or without one of six commercial AM fungal products. Our research showed no evidence of benefit; commercial inoculants never increased native plant biomass, although several inoculants decreased the growth of native species and increased the growth of invasive plant species. In addition, two commercial products contained excessive levels of phosphorus or nitrogen and consistently reduced AM fungal root colonization, indicating an unintentional de-coupling of the symbiosis. As there is little knowledge of the ecological consequences of inoculation with commercial AM fungal products, it is critical for restoration practitioners, scientists, and native plant growers to assess the presence of local AM fungal communities before investing in unnecessary, or possibly detrimental, AM fungal products.

Arbuscular Mycorrhizal Fungi Taxa Show Variable Patterns of Micro-Scale Dispersal in Prairie Restorations.

Human land use disturbance is a major contributor to the loss of natural plant communities, and this is particularly true in areas used for agriculture, such as the Midwestern tallgrass prairies of the United States. Previous work has shown that arbuscular mycorrhizal fungi (AMF) additions can increase native plant survival and success in plant community restorations, but the dispersal of AMF in these systems is poorly understood. In this study, we examined the dispersal of AMF taxa inoculated into four tallgrass prairie restorations. At each site, we inoculated native plant species with greenhouse-cultured native AMF taxa or whole soil collected from a nearby unplowed prairie. We monitored AMF dispersal, AMF biomass, plant growth, and plant community composition, at different distances from inoculation. In two sites, we assessed the role of plant hosts in dispersal, by placing known AMF hosts in a "bridge" and "island" pattern on either side of the inoculation points. We found that AMF taxa differ in their dispersal ability, with some taxa spreading to 2-m in the first year and others remaining closer to the inoculation point. We also found evidence that AMF spread altered non-inoculated neighboring plant growth and community composition in certain sites. These results represent the most comprehensive attempt to date to evaluate AMF spread.

The root-associated arbuscular mycorrhizal fungal assemblages of exotic alien plants are simplified in invaded distribution ranges, but dominant species are retained: A trans-continental perspective.

Arbuscular mycorrhizal fungi (AMF) provide crucial support for the establishment of plants in novel environments. We hypothesized that the OTU/genus richness and diversity of soil- and root-associated AMF associated with alien plant species in their exotic ranges are lower than those in their native ranges. We examined the root-associated and soil-dwelling AMF of 11 invasive plant species in their native and exotic ranges in the United States and Europe by DNA sequencing of the ITS2 locus. Examined root-associated AMF assemblages were simplified, which manifested as the loss of several AMF genera in the exotic ranges of the plants. These fungal assemblages were also characterized by greater dominance and simplification of the fungal assemblages. The dominant fungal genera were present regardless of whether their host plants were in their native or exotic ranges. Interestingly, both the native and invaded soils hosted diverse local AMF assemblages. Therefore, alien plant invasions were not limited to soils with low AMF diversity. Some AMF taxa could be context-dependent passengers rather than drivers of alien plant invasions. Further studies should identify functions of AMF missing or less abundant in roots of plants growing in exotic ranges.

Mycorrhizal and rhizobial interactions influence model grassland plant community structure and productivity.

Arbuscular mycorrhizal (AM) fungi and rhizobium are likely important drivers of plant coexistence and grassland productivity due to complementary roles in supplying limiting nutrients. However, the interactive effects of mycorrhizal and rhizobial associations on plant community productivity and competitive dynamics remain unclear. To address this, we conducted a greenhouse experiment to determine the influences of these key microbial functional groups on communities comprising three plant species by comparing plant communities grown with or without each symbiont. We also utilized N-fertilization and clipping treatments to explore potential shifts in mycorrhizal and rhizobial benefits across abiotic and biotic conditions. Our research suggests AM fungi and rhizobium co-inoculation was strongly facilitative for plant community productivity and legume (Medicago sativa) growth and nodulation. Plant competitiveness shifted in the presence of AM fungi and rhizobium, favoring M. sativa over a neighboring C4 grass (Andropogon gerardii) and C3 forb (Ratibida pinnata). This may be due to rhizobial symbiosis as well as the relatively greater mycorrhizal growth response of M. sativa, compared to the other model plants. Clipping and N-fertilization altered relative costs and benefits of both symbioses, presumably by altering host-plant nitrogen and carbon dynamics, leading to a relative decrease in mycorrhizal responsiveness and proportional biomass of M. sativa relative to the total biomass of the entire plant community, with a concomitant relative increase in A. gerardii and R. pinnata proportional biomass. Our results demonstrate a strong influence of both microbial symbioses on host-plant competitiveness and community dynamics across clipping and N-fertilization treatments, suggesting the symbiotic rhizosphere community is critical for legume establishment in grasslands.

Arbuscular mycorrhizal fungi favor invasive Echinops sphaerocephalus when grown in competition with native Inula conyzae.

In a globalized world, plant invasions are common challenges for native ecosystems. Although a considerable number of invasive plants form arbuscular mycorrhizae, interactions between arbuscular mycorrhizal (AM) fungi and invasive and native plants are not well understood. In this study, we conducted a greenhouse experiment examining how AM fungi affect interactions of co-occurring plant species in the family Asteracea, invasive Echinops sphaerocephalus and native forb of central Europe Inula conyzae. The effects of initial soil disturbance, including the effect of intact or disturbed arbuscular mycorrhizal networks (CMNs), were examined. AM fungi supported the success of invasive E. sphaerocephalus in competition with native I. conyzae, regardless of the initial disturbance of CMNs. The presence of invasive E. sphaerocephalus decreased mycorrhizal colonization in I. conyzae, with a concomitant loss in mycorrhizal benefits. Our results confirm AM fungi represent one important mechanism of plant invasion for E. sphaerocephalus in semi-natural European grasslands.

Arbuscular mycorrhizal fungi in roots and soil respond differently to biotic and abiotic factors in the Serengeti.

This study explores the relationships of AM fungal abundance and diversity with biotic (host plant, ungulate grazing) and abiotic (soil properties, precipitation) factors in the Serengeti National Park, Tanzania. Soil and root samples were collected from grazed and ungrazed plots at seven sites across steep soil fertility and precipitation gradients. AM fungal abundance in the soil was estimated from the density of spores and the concentration of a fatty acid biomarker. Diversity of AM fungi in roots and soils was measured using DNA sequencing and spore identification. AM fungal abundance in soil decreased with grazing and precipitation and increased with soil phosphorus. The community composition of AM fungal DNA in roots and soils differed. Root samples had more AM fungal indicator species associated with biotic factors (host plant species and grazing), and soil samples had more indicator species associated with particular sample sites. These findings suggest that regional edaphic conditions shape the site-level species pool from which plant species actively select root-colonizing fungal assemblages modified by grazing. Combining multiple measurements of AM fungal abundance and community composition provides the most informed assessment of the structure of mycorrhizal fungal communities in natural ecosystems.

Plant Diversity and Fertilizer Management Shape the Belowground Microbiome of Native Grass Bioenergy Feedstocks.

Plants may actively cultivate microorganisms in their roots and rhizosphere that enhance their nutrition. To develop cropping strategies that substitute mineral fertilizers for beneficial root symbioses, we must first understand how microbial communities associated with plant roots differ among plant taxa and how they respond to fertilization. Arbuscular mycorrhizal (AM) fungi and rhizobacteria are of particular interest because they enhance nutrient availability to plants and perform a suite of nutrient cycling functions. The purpose of this experiment is to examine the root and soil microbiome in a long-term switchgrass (Panicum virgatum) biofuel feedstock experiment and determine how AM fungi and rhizobacteria respond to plant diversity and soil fertility. We hypothesize that intra- and interspecific plant diversity, nitrogen fertilization (+N), and their interaction will influence the biomass and community composition of AM fungi and rhizobacteria. We further hypothesize that +N will reduce the abundance of nitrogenase-encoding nifH genes on the rhizoplane. Roots and soils were sampled from three switchgrass cultivars (Cave-in-Rock, Kanlow, Southlow) grown in monoculture, intraspecific mixture, and interspecific planting mixtures with either Andropogon gerardii or diverse native tallgrass prairie species. Molecular sequencing was performed on root and soil samples, fatty acid extractions were assessed to determine microbial biomass, and quantitative polymerase chain reaction (qPCR) was performed on nifH genes from the rhizoplane. Sequence data determined core AM fungal and bacterial microbiomes and indicator taxa for plant diversity and +N treatments. We found that plant diversity and +N influenced AM fungal biomass and community structure. Across all plant diversity treatments, +N reduced the biomass of AM fungi and nifH gene abundance by more than 40%. The AM fungal genus Scutellospora was an indicator for +N, with relative abundance significantly greater under +N and in monoculture treatments. Community composition of rhizobacteria was influenced by plant diversity but not by +N. Verrucomicrobia and Proteobacteria were the dominant bacterial phyla in both roots and soils. Our findings provide evidence that soil fertility and plant diversity structure the root and soil microbiome. Optimization of soil communities for switchgrass production must take into account differences among cultivars and their unique responses to shifts in soil fertility.

Determining landscape-level drivers of variability for over fifty soil chemical elements.

Syntheses of large datasets have allowed increased clarity of distribution patterns and variation in soil major and trace elements. However, the drivers of variation in topsoil elements across biogeographical scales are not well understood. Our aim was to (1) identify how landscape-scale climate, geographical features, and edaphic factors influence soil elements, and (2) determine key environmental thresholds for shifts in soil element concentration. We analyzed patterns of variation in topsoil elements using 9830 samples collected across 39,000km2 in subtropical land in southeast China. Canonical correlations and multiple linear regressions were used to model variations of each element across mean annual temperature (MAT), mean annual precipitation (MAP), land use, spatial topography, and soil pH. Element concentrations show significant latitudinal and longitudinal trends, and are significantly influenced by climate, land use, spatial topography, and soil pH. Longitude, pH, MAT, and MAP were the environmental factors most tightly correlated with element concentrations. Climate and soil pH drove positive or negative alterations in soil elements, with threshold indicators of MAP=1000mm/1500mm, MAT=17.8°C/18.0°C, and pH=5.8/5.0, respectively. Our results indicate topsoil elements have structural and functional thresholds of climate and soil pH in relatively wet and acidic environments. Our findings can facilitate holistic soil element concentration predictions and help elucidate the specific influences of climate and soil pH, enabling development of more complete biogeochemical models.

Erratum: Author Correction: Evolutionary history of plant hosts and fungal symbionts predicts the strength of mycorrhizal mutualism.

[This corrects the article DOI: 10.1038/s42003-018-0120-9.].

Evolutionary history of plant hosts and fungal symbionts predicts the strength of mycorrhizal mutualism.

Most plants engage in symbioses with mycorrhizal fungi in soils and net consequences for plants vary widely from mutualism to parasitism. However, we lack a synthetic understanding of the evolutionary and ecological forces driving such variation for this or any other nutritional symbiosis. We used meta-analysis across 646 combinations of plants and fungi to show that evolutionary history explains substantially more variation in plant responses to mycorrhizal fungi than the ecological factors included in this study, such as nutrient fertilization and additional microbes. Evolutionary history also has a different influence on outcomes of ectomycorrhizal versus arbuscular mycorrhizal symbioses; the former are best explained by the multiple evolutionary origins of ectomycorrhizal lifestyle in plants, while the latter are best explained by recent diversification in plants; both are also explained by evolution of specificity between plants and fungi. These results provide the foundation for a synthetic framework to predict the outcomes of nutritional mutualisms.

Influence of alternative soil amendments on mycorrhizal fungi and cowpea production.

Alternative soil amendments (worm compost, pyrolyzed carbon [biochar]) and crop symbioses with arbuscular mycorrhizal (AM) fungi have the potential to reduce food production costs while promoting sustainable agriculture by improving soil quality and reducing commercial (N and P) fertilizer use. Our greenhouse studies investigated the influence of alternative soil amendments on AM fungi associated with cowpea (Vigna unguiculata [L.] Walp.) and common bean (Phaseolus vulgaris L.) by examining productivity and plant nutrition. We conducted an experiment to select a cowpea or common bean genotype based on AM fungal colonization, seed production, and seed nutritional content. We then grew the selected cowpea genotype (Resina) in low-fertility soil with 10 different soil amendments (combinations of biochar, worm compost, and/or commercial fertilizers) plus a non-amended control. There were no significant differences in AM fungal colonization of cowpea plants grow with different soil amendments. However, an amendment blend containing worm compost, biochar, and 50% of the typically recommended commercial fertilizer rate produced plants with similar aboveground biomass, protein concentration, and total protein production, with increased tissue K, P, and Zn concentration and total content, compared to plants receiving only the recommended (100%) rate of commercial fertilizer. As previous research links uptake of P and Zn with plant-mycorrhizal symbioses, our results indicate cowpea nutritional benefits may be derived from AM partnership and alternative soil amendments. These synergies between alternative soil amendments and AM fungi may help reduce farm costs while maintaining or improving crop yield and nutrition, thus increasing global food and nutrition security.

Trichoderma Biofertilizer Links to Altered Soil Chemistry, Altered Microbial Communities, and Improved Grassland Biomass.

In grasslands, forage and livestock production results in soil nutrient deficits as grasslands typically receive no nutrient inputs, leading to a loss of grassland biomass. The application of mature compost has been shown to effectively increase grassland nutrient availability. However, research on fertilization regime influence and potential microbial ecological regulation mechanisms are rarely conducted in grassland soil. We conducted a two-year experiment in meadow steppe grasslands, focusing on above- and belowground consequences of organic or Trichoderma biofertilizer applications and potential soil microbial ecological mechanisms underlying soil chemistry and microbial community responses. Grassland biomass significantly (p = 0.019) increased following amendment with 9,000 kg ha-1 of Trichoderma biofertilizer (composted cattle manure + inoculum) compared with other assessed organic or biofertilizer rates, except for BOF3000 (fertilized with 3,000 kg ha-1 biofertilizer). This rate of Trichoderma biofertilizer treatment increased soil antifungal compounds that may suppress pathogenic fungi, potentially partially responsible for improved grassland biomass. Nonmetric multidimensional scaling (NMDS) revealed soil chemistry and fungal communities were all separated by different fertilization regime. Trichoderma biofertilizer (9,000 kg ha-1) increased relative abundances of Archaeorhizomyces and Trichoderma while decreasing Ophiosphaerella. Trichoderma can improve grassland biomass, while Ophiosphaerella has the opposite effect as it may secrete metabolites causing grass necrosis. Correlations between soil properties and microbial genera showed plant-available phosphorus may influence grassland biomass by increasing Archaeorhizomyces and Trichoderma while reducing Ophiosphaerella. According to our structural equation modeling (SEM), Trichoderma abundance was the primary contributor to aboveground grassland biomass. Our results suggest Trichoderma biofertilizer could be an important tool for management of soils and ultimately grassland plant biomass.

Defoliation and arbuscular mycorrhizal fungi shape plant communities in overgrazed semiarid grasslands.

Overgrazing substantially contributes to global grassland degradation by decreasing plant community productivity and diversity through trampling, defoliation, and removal of nutrients. Arbuscular mycorrhizal (AM) fungi also play a critical role in plant community diversity, composition, and primary productivity, maintaining ecosystem functions. However, interactions between grazing disturbances, such as trampling and defoliation, and AM fungi in grassland communities are not well known. We examined influences of trampling, defoliation, and AM fungi on semiarid grassland plant community composition for 3 yr, by comparing all combinations of these factors. Benomyl fungicide was applied to reduce AM fungal abundance. Overgrazing typically resulted in reduced dominance of Stipa Krylovii, contributing to degradation of typical steppe grasslands. Our results indicated trampling generally had little effect on plant community composition, unless combined with defoliation or AM fungal suppression. Defoliation was the main component of grazing that promoted dominance of Potentilla acaulis over Stipa krylovii and Artemisia frigida, presumably by alleviating light limitation. In non-defoliated plots, AM fungi promoted A. frigida, with a concomitant reduction in S. krylovii growth compared to corresponding AM suppressed plots. Our results indicate AM fungi and defoliation jointly suppress S. krylovii biomass; however, prolonged defoliation weakens mycorrhizal influence on plant community composition. These findings give new insight into dominant plant species shifts in degraded semiarid grasslands.

Long-term effects of grazing and topography on extra-radical hyphae of arbuscular mycorrhizal fungi in semi-arid grasslands.

Grazing and topography have drastic effects on plant communities and soil properties. These effects are thought to influence arbuscular mycorrhizal (AM) fungi. However, the simultaneous impacts of grazing pressure (sheep ha-1) and topography on plant and soil factors and their relationship to the production of extra-radical AM hyphae are not well understood. Our 10-year study assessed relationships between grazing, plant species richness, aboveground plant productivity, soil nutrients, edaphic properties, and AM hyphal length density (HLD) in different topographic areas (flat or sloped). We found HLD linearly declined with increasing grazing pressure (1.5-9.0 sheep ha-1) in sloped areas, but HLD was greatest at moderate grazing pressure (4.5 sheep ha-1) in flat areas. Structural equation modeling indicates grazing reduces HLD by altering soil nutrient dynamics in sloped areas, but non-linearly influences HLD through plant community and edaphic changes in flat areas. Our findings highlight how topography influences key plant and soil factors, thus regulating the effects of grazing pressure on extra-radical hyphal production of AM fungi in grasslands. Understanding how grazing and topography influence AM fungi in semi-arid grasslands is vital, as globally, severe human population pressure and increasing demand for food aggravate the grazing intensity in grasslands.

MycoDB, a global database of plant response to mycorrhizal fungi.

Plants form belowground associations with mycorrhizal fungi in one of the most common symbioses on Earth. However, few large-scale generalizations exist for the structure and function of mycorrhizal symbioses, as the nature of this relationship varies from mutualistic to parasitic and is largely context-dependent. We announce the public release of MycoDB, a database of 4,010 studies (from 438 unique publications) to aid in multi-factor meta-analyses elucidating the ecological and evolutionary context in which mycorrhizal fungi alter plant productivity. Over 10 years with nearly 80 collaborators, we compiled data on the response of plant biomass to mycorrhizal fungal inoculation, including meta-analysis metrics and 24 additional explanatory variables that describe the biotic and abiotic context of each study. We also include phylogenetic trees for all plants and fungi in the database. To our knowledge, MycoDB is the largest ecological meta-analysis database. We aim to share these data to highlight significant gaps in mycorrhizal research and encourage synthesis to explore the ecological and evolutionary generalities that govern mycorrhizal functioning in ecosystems.

Belowground interactions with aboveground consequences: Invasive earthworms and arbuscular mycorrhizal fungi.

A mounting body of research suggests that invasive nonnative earthworms substantially alter microbial communities, including arbuscular mycorrhizal fungi (AMF). These changes to AMF can cascade to affect plant communities and vertebrate populations. Despite these research advances, relatively little is known about (1) the mechanisms behind earthworms' effects on AMF and (2) the factors that determine the outcomes of earthworm-AMF interactions (i.e., whether AMF abundance is increased or decreased and subsequent effects on plants). We predict that AMF-mediated effects of nonnative earthworms on ecosystems are nearly universal because (1) AMF are important components of most terrestrial ecosystems, (2) nonnative earthworms have become established in nearly every type of terrestrial ecosystem, and (3) nonnative earthworms, due to their burrowing and feeding behavior, greatly affect AMF with potentially profound concomitant effects on plant communities. We highlight the multiple direct and indirect effects of nonnative earthworms on plants and review what is currently known about the interaction between earthworms and AMF. We also illustrate how the effects of nonnative earthworms on plant-AMF mutualisms can alter the structure and stability of aboveground plant communities, as well as the vertebrate communities relying on these habitats. Integrative studies that assess the interactive effects of earthworms and AMF can provide new insights into the role that belowground ecosystem engineers play in altering aboveground ecological processes. Understanding these processes may improve our ability to predict the structure of plant and animal communities in earthworm-invaded regions and to develop management strategies that limit the numerous undesired impacts of earthworms.

Mycorrhizal phenotypes and the Law of the Minimum.

Mycorrhizal phenotypes arise from interactions among plant and fungal genotypes and the environment. Differences in the stoichiometry and uptake capacity of fungi and plants make arbuscular mycorrhizal (AM) fungi inherently more nitrogen (N) limited and less phosphorus (P) limited than their host plants. Mutualistic phenotypes are most likely in P-limited systems and commensal or parasitic phenotypes in N-limited systems. Carbon (C) limitation is expected to cause phenotypes to shift from mutualism to commensalism and even parasitism. Two experiments compared the influence of fertilizer and shade on mycorrhizas in Andropogon gerardii across three naturally N-limited or P-limited grasslands. A third experiment examined the interactive effects of N and P enrichment and shade on A. gerardii mycorrhizas. Our experiments generated the full spectrum of mycorrhizal phenotypes. These findings support the hypothesis that mutualism is likely in P-limited systems and commensalism or parasitism is likely in N-limited systems. Furthermore, shade decreased C-assimilation and generated less mutualistic mycorrhizal phenotypes with reduced plant and fungal biomass. Soil fertility is a key controller of mycorrhizal costs and benefits and the Law of the Minimum is a useful predictor of mycorrhizal phenotype. In our experimental grasslands arbuscular mycorrhizas can ameliorate P-limitation but not N-limitation.