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.     

Soil temperature affects carbon allocation within arbuscular mycorrhizal networks and carbon transport from plant to fungus

How soil carbon balance will be affected by plant–mycorrhizal interactions under future climate scenarios remains a significant unknown in our ability to forecast ecosystem carbon storage and fluxes. We examined the effects of soil temperature (14, 20, 26 °C) on the structure and extent of a multispecies community of arbuscular mycorrhizal (AM) fungi associated with Plantago lanceolata. To isolate fungi from roots, we used a mesh‐divided pot system with separate hyphal compartments near and away from the plant. A ¹³C pulse label was then used to trace the flow of recently fixed photosynthate from plants into belowground pools and respiration. Temperature significantly altered the structure and allocation of the AM hyphal network, with a switch from more vesicles (storage) in cooled soils to more extensive extraradical hyphal networks (growth) in warmed soils. As soil temperature increased, we also observed an increase in the speed at which plant photosynthate was transferred to and respired by roots and AM fungi coupled with an increase in the amount of carbon respired per unit hyphal length. These differences were largely independent of plant size and rates of photosynthesis. In a warmer world, we would therefore expect more carbon losses to the atmosphere from AM fungal respiration, which are unlikely to be balanced by increased growth of AM fungal hyphae.     

Mounding alters environmental filters that drive plant community development in a novel grassland

Earthen mounds are commonly used in ecological restoration to increase environmental heterogeneity, create favorable microclimates and retain soil resources that promote plant establishment. Although mounding is commonly employed in restoration, few microtopography studies focus on the long-term effects of mounding on restored plant community development. We assessed the vegetation and physical environment of earthen mounds installed at a novel grassland ten years after restoration to look for patterns in plant community development. We used permutational multiple analysis of variance (PERMANOVA) to identify differences in plant community composition and the associated mound-driven environmental variables, summer soil moisture and height above peak soil inundation, in relation to mound position. We used indicator species analysis (ISA) to classify the species that defined mound top and intermound space plant communities. We found that mound position drove plot height above flooding and soil moisture while plant community composition was driven by plot height above flooding, summer soil moisture, and mound position. ISA showed that species colonized mound microsites differently: most wetland species occurred between mounds and xeric stress tolerators largely occupied dry mound tops. We visualized these differences with non-metric multidimensional scaling (NMDS) ordination, finding that species sorted out in multivariate space based on mound position. We conclude that mounding can have relatively long-term effects on plant community development, even in highly disturbed, minimally maintained restoration projects.     

Soil heterogeneity and community composition jointly influence grassland biomass

Question: Does the spatial pattern of nutrient supply modify community biomass responses to changes in both species composition and richness? Location: Duke University Phytotron (Durham, North Carolina, USA). Methods: We conducted a microcosm experiment to evaluate individual plant and whole community responses to species richness, species composition and soil nutrient heterogeneity. The experiment consisted of seven levels of species composition (all possible combinations of Lolium perenne, Poa pratensis and Plantago lanceolata) crossed with three levels of soil nutrient distribution (homogeneous, heterogeneous‐up, and heterogeneous‐down, where up and down indicates the location of a nutrient patch in either the upper or the lower half of the soil column, respectively). Results: Communities containing Plantago and Lolium responded to nutrient heterogeneity by increasing above‐ and below‐ground biomass. Nutrient heterogeneity also increased size inequalities among individuals of these species. Significant species composition X nutrient heterogeneity interactions on community biomass and individual size inequality were observed when nutrient patches were located in the upper 10 cm of the soil columns. However, root proliferation in nutrient patches was equivalent regardless of the vertical placement of the patch. Conclusions: Our results suggest that nutrient heterogeneity may interact with plant species composition to determine community biomass, and that small‐scale vertical differences in the location of nutrient patches affect individual and community responses to this heterogeneity.     

Small-scale spatial heterogeneity of arbuscular mycorrhizal fungal abundance and community composition in a wetland plant community

Although it has become increasingly clear that arbuscular mycorrhizal fungi (AMF) play important roles in population, community, and ecosystem ecology, there is limited information on the spatial structure of the community composition of AMF in the field. We assessed small-scale spatial variation in the abundance and molecular diversity of AMF in a calcareous fen, where strong underlying environmental gradients such as depth to water table may influence AMF. Throughout an intensively sampled 2 × 2 m plot, we assessed AMF inoculum potential at a depth of 0–6 and 6–12 cm and molecular diversity of the AMF community using terminal restriction fragment length polymorphism of 18S rDNA. Inoculum potential was only significantly spatially autocorrelated at a depth of 6–12 cm and was significantly positively correlated with depth to water table at both depths. Molecular diversity of the AMF community was highly variable within the plot, ranging from 2–14 terminal restriction fragments (T-RFs) per core, but the number of T-RFs did not relate to water table or plant species richness. Plant community composition was spatially autocorrelated at small scales, but AMF community composition showed no significant spatial autocorrelation. Saturated soils of calcareous fens contain many infective AMF propagules and the abundance and diversity of AMF inoculum is patchy over small spatial scales. An erratum to this article can be found at     

Soil surface disturbance alters cyanobacterial biocrusts and soil properties in dry grassland and shrubland ecosystems

Plant and Soil - Biological soil crusts (biocrusts) dominate soil surfaces in drylands, providing services that include soil stabilization and carbon uptake. In this study, we investigated the...     

Global change alters peatland carbon cycling through plant biomass allocation

Plant and Soil - Global change is shown to significantly affect the C storage function of peatlands; however, a majority of previous research is focused on a single environmental stressor such as...     

Soil community composition and the regulation of grazed temperate grassland

The effect of the community composition of soil microbes on ecosystem processes has received relatively little attention. Here we examined the variation in soil microbial composition in a Yellowstone National Park grassland and the effect of that variation on the growth, in a greenhouse, of the dominant grass in the community. Plants and their rhizospheric soil were collected from paired, Poa pratensis-dominated grassland plots located inside and outside a 40-year-old exclosure. P. pratensis aboveground, belowground, and whole plant growth were greater in pots with soil communities from grazed grassland compared to fenced grassland, indicating (1) soil microbial communities differed, and (2) this difference influenced the growth of the plant that dominated both grasslands. Treating pots with fungicide (benomyl) suppressed the soil community influence, indicating that different fungal communities caused the soil microbe effect. In addition, two lines of evidence are consistent with the hypothesis that arbuscular mycorrhizal fungal (AMF) species composition affected P. pratensis: (1) a divergence in AMF spore communities in the two field soils, and (2) little evidence of pathogenic fungi. These findings emphasize the need to examine the role that the composition of the soil microbial community plays in controlling terrestrial ecosystems.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Exotic plant species in a C4-dominated grassland: invasibility, disturbance, and community structure

We used data from a 15-year experiment in a C₄-dominated grassland to address the effects of community structure (i.e., plant species richness, dominance) and disturbance on invasibility, as measured by abundance and richness of exotic species. Our specific objectives were to assess the temporal and spatial patterns of exotic plant species in a native grassland in Kansas (USA) and to determine the factors that control exotic species abundance and richness (i.e., invasibility). Exotic species (90% C₃ plants) comprised approximately 10% of the flora, and their turnover was relatively high (30%) over the 15-year period. We found that disturbances significantly affected the abundance and richness of exotic species. In particular, long-term annually burned watersheds had lower cover of exotic species than unburned watersheds, and fire reduced exotic species richness by 80–90%. Exotic and native species richness were positively correlated across sites subjected to different fire (r = 0.72) and grazing (r = 0.67) treatments, and the number of exotic species was lowest on sites with the highest productivity of C₄ grasses (i.e., high dominance). These results provide strong evidence for the role of community structure, as affected by disturbance, in determining invasibility of this grassland. Moreover, a significant positive relationship between exotic and native species richness was observed within a disturbance regime (annually burned sites, r = 0.51; unburned sites, r = 0.59). Thus, invasibility of this C₄-dominated grassland can also be directly related to community structure independent of disturbance. Received: 9 February 1999 / Accepted: 12 May 1999     

Effects of earthworms and arbuscular mycorrhizal fungi depend on the successional stage of a grassland plant community

Earthworms and arbuscular mycorrhizal fungi (AMF) have profound impacts on plant performance. However, it is largely unknown if and how earthworms and AMF may affect plant succession. We planted mesocosms with an early-mid successional and a mid-late successional grassland plant community and added endogeic earthworms and commercial AMF in a full-factorial way to natural background soil. Earthworms had a positive effect on the total root and shoot biomass of both plant communities, with the effect on the shoots being slightly enhanced by co-inoculation with AMF. Surprisingly, the earthworm effect on the mid successional plant species depended on the successional stage of the plant community. Earthworms had a positive effect on the mid successional plant species when they were growing in the mid-late successional plant community, but no effect when the same plant species were growing in the early-mid successional plant community. Addition of AMF alone tended to reduce the shoot biomass of the early successional plant species, while the addition of earthworms in the presence or absence of AMF increased their shoot biomass. We conclude that the impacts of earthworms on plant species may depend on the successional stage of the plant community, while the effect of AMF addition depends on the successional stage of the plant community and may be changed by the presence of earthworms. Earthworms and AMF addition affect plants and plant communities of different successional stages differently with potential effects on plant succession.     

Arbuscular mycorrhizal fungi alter plant community composition along a grazing gradient in Inner Mongolia Steppe

Arbuscular mycorrhizal fungi (AMF) have a significant influence on plant productivity and diversity in non-grazing grassland. However, the interactive effects between grazing intensity and AMF on plant community composition in natural grassland communities are not well known. We conducted a field experiment that manipulated AMF colonization and grazing intensity to study the impact of AMF suppression on plant community composition and nutrient status over 2 years (2015–2016) with contrasting rainfall levels. We found that AMF root colonization was significantly reduced by the application of the fungicide benomyl as a soil drench. Grazing intensity regulated plant community composition and aboveground biomass mainly by reducing the growth of Leymus chinensis over 2 years. AMF suppression increased the growth of Chenopodium glaucum, but it did not alter other plant species across all grazing intensities. The effects of AMF suppression on plant community composition changed along a grazing gradient considerably between years: AMF suppression increased the biomass of C. glaucum across all grazing intensities in 2015, but slightly increased it in 2016. Interactions between AMF suppression and grazing intensity altered the phosphorus concentration of Stipa grandis and Cleistogenes squarrosa in 2015 but not in 2016. AMF suppression decreased the shoot phosphorus content of L. chinensis but increased that of C. glaucum across all grazing intensities. Our results indicate that grazing intensity substantially alters aboveground community biomass and affects growth of dominant species; AMF by itself have limited effects on plant communities along a grazing gradient in typical steppe.     

Absence of snow cover reduces understory plant cover and alters plant community composition in boreal forests

Snow regimes affect biogeochemistry of boreal ecosystems and are altered by climate change. The effects on plant communities, however, are largely unexplored despite their influence on relevant processes. Here, the impact of snow cover on understory community composition and below-ground production in a boreal Picea abies forest was investigated using a long-term (8-year) snow cover manipulation experiment consisting of the treatments: snow removal, increased insulation (styrofoam pellets), and control. The snow removal treatment caused longer (118 vs. 57 days) and deeper soil frost (mean minimum temperature −5.5 vs. −2.2°C) at 10 cm soil depth in comparison to control. Understory species composition was strongly altered by the snow cover manipulations; vegetation cover declined by more than 50% in the snow removal treatment. In particular, the dominant dwarf shrub Vaccinium myrtillus (−82%) and the most abundant mosses Pleurozium schreberi (−74%) and Dicranum scoparium (−60%) declined strongly. The C:N ratio in V. myrtillus leaves and plant available N in the soil indicated no altered nitrogen nutrition. Fine-root biomass in summer, however, was negatively affected by the reduced snow cover (−50%). Observed effects are attributed to direct frost damage of roots and/ or shoots. Besides the obvious relevance of winter processes on plant ecology and distribution, we propose that shifts in the vegetation caused by frost damage may be an important driver of the reported alterations in biogeochemistry in response to altered snow cover. Understory plant performance clearly needs to be considered in the biogeochemistry of boreal systems in the face of climate change.     

Large herbivores facilitate an insect herbivore by modifying plant community composition in a temperate grassland

Large herbivores often co‐occur and share plant resources with herbivorous insects in grassland ecosystems; yet, how they interact with each other remains poorly understood. We conducted a series of field experiments to investigate whether and how large domestic herbivores (sheep; Ovis aries) may affect the abundance of a common herbivorous insect (aphid; Hyalopterus pruni) in a temperate grassland of northeast China. Our exclosure experiment showed that 3 years (2010–2012) of sheep grazing had led to 86% higher aphid abundance compared with ungrazed sites. Mechanistically, this facilitative effect was driven by grazing altering the plant community, rather than by changes in food availability and predator abundance for aphids. Sheep significantly altered plant community by reducing the abundance of unpalatable forbs for the aphids. Our small‐scale forb removal experiment revealed an “associational plant defense” by forbs which protect the grass Phragmites australis from being attacked by the aphids. However, selective grazing on forbs by sheep indirectly disrupted such associational plant defense, making P. australis more susceptible to aphids, consequentially increasing the density of aphids. These findings provide a novel mechanistic explanation for the effects of large herbivores on herbivorous insects by linking selective grazing to plant community composition and the responses of insect populations in grassland ecosystems.     

Seasonal carbon isotope discrimination in a grassland community

Summary Grassland communities of arid western North America are often characterized by a seasonal increase in ambient temperature and evaporative demand and a corresponding decline in soil moisture availability. As the environment changes, particular species could respond differently, which should be reflected in a number of physiological processes. Carbon isotope discrimination varies during photosynthetic activity as a function of both stomatal aperture and the biochemistry of the fixation process, and provides an integrated measure of plant response to seasonal changes in the environment. We measured the seasonal course of carbon isotope discrimination in 42 grassland species to evaluate changes in gas exchange processes in response to these varying environmental factors. The seasonal courses were then used to identify community-wide patterns associated with life form, with phenology and with differences between grasses and forbs. Significant differences were detected in the following comparisons: (1) Carbon isotope discrimination decreased throughout the growing season; (2) perennial species discriminated less than annual species; (3) grasses discriminated less than forbs; and (4) early flowering species discriminated more than the later flowering ones. These comparisons suggested that (1) species active only during the initial, less stressful months of the growing season used water less efficiently, and (2) that physiological responses increasing the ratio of carbon fixed to water lost were common in these grassland species, and were correlated with the increase in evaporative demand and the decrease in soil moisture.     

Carbon allocation mediated by arbuscular mycorrhizal fungi alters the soil microbial community under various phosphorus levels

Studies have shown that arbuscular mycorrhizal fungi (AMF) can shape the rhizosphere microbial community of the host plant, but the underlying mechanisms are unclear. Here, we tested the hypotheses that AMF may affect the rhizosphere microbial community by mediating carbon (C) allocation of the host plant, and that this mediation may be modulated by the soil phosphorus (P) level. Using a split-root system, we conducted a microcosm experiment with three main effects (soil P level: 5 or 25 mg kg⁻¹; AMF: with or without inoculation; and spatial niche (i.e., rhizosphere, hyphosphere, and mycorrhizosphere). Host plant growth benefited from AMF under low soil-P conditions. ¹³CO₂ isotope labeling showed that AMF increased C allocation to the colonized root and AMF mycelia under low-P conditions, which promoted AMF growth. ¹³C-DNA-SIP and 16S rRNA sequencing further indicated that the enhanced C allocation from the host altered the soil microbial community. Our results suggest that AMF enhances the C allocation of the host plant below ground, which can shape microbial community composition. These AMF effects were greater with a low than with a high level of soil P.