Minimal direct contribution of arbuscular mycorrhizal fungi to DOC leaching in grassland through losses of glomalin-related soil protein

Arbuscular mycorrhizal fungi (AMF) have multiple influences on ecosystem C cycling, but most research has focused on ecosystem C gains. We explore here the possibility of direct contributions of AMF to ecosystem C losses, namely via leaching of glomalin-related soil protein (GRSP). We tested the hypothesis that GRSP, an operationally defined SOM pool to which AMF contribute (especially as evidenced with monoclonal antibody MAb32B11-based detection), is mobile in soils and can be lost in leachate. For two New Zealand soils, we showed that only insignificant amounts of GRSP were lost: a maximum of 0.03% of MAb32B11-immunoreactive GRSP present in soils was lost during the week-long experiment, representing a minute fraction of total leachate dissolved organic carbon (0.06%). Our data showed that this pathway of C loss may be relatively unimportant in many soils. However, other indirect contributions of AMF to soil C losses remain yet to be explored.     

Relating microarthropod community structure and diversity to soil fertility manipulations in temperate grassland

We aimed to identify patterns of diversity in a below-ground community of microarthropods (mites and Collembola) after 15 months of a nutrient (calcium and nitrogen) manipulation experiment, located at the Natural Environment Research Council (NERC) Soil Biodiversity Site in Scotland, UK. We found that microarthropod densities increased with elevated soil fertility, but we detected no concurrent change in the diversity of soil microarthropods (mites and Collembola combined). That microarthropod density increased concurrently with improvements in soil fertility and plant productivity suggests that soil microarthropod communities are predominately regulated by bottom-up forces, driven by increased energy transfer via plant inputs to soil, providing increased food resources for fauna. However, that we found no concurrent change in the diversity of soil microarthropods provides little support for the idea that the diversity of soil fauna is positively related to their population density, primary productivity or improvements in soil conditions resulting from nutrient manipulations. However, we did find that microarthropod communities of more fertile sites contained a greater proportion of predators suggesting that more energy was transferred to higher trophic levels under elevated soil fertility. Our findings suggest that unlike plant communities, soil faunal diversity may not be strongly regulated by competition in productive situations, since competitive exclusion might not occur due to increased predation. Whilst we conclude that soil microarthropod diversity at our study site has not been affected by the nutrient additions to date, in the longer term we predict that changes in community composition and diversity could arise, most likely through top-down regulation of the soil food web.     

Consumption of mycorrhizal and saprophytic fungi by Collembola in grassland soils

Although soil-dwelling Collembola can influence plant growth and nutrient cycling, their specific role in soil food webs is poorly understood. Soil-free microcosm studies suggest that Collembola are primarily fungivores where they feed preferentially on saprophytic fungi (SF) over other fungal types. We directly assessed collembolan consumption of arbuscular mycorrhizal fungi (AMF) and SF using plant-soil mesocosms and natural abundance stable carbon isotope techniques. Mycorrhizal Andropogon gerardii (C₄ grass) seedlings were placed in pots containing Collembola and soil from a C₃ plant dominated site, while mycorrhizal Pascopyrum smithii (C₃ grass) seedlings were placed in pots with Collembola and soil collected at a C₄ plant dominated site. After 6 weeks, collembolans assimilated carbon derived from C₃ and C₄ sources in both A. gerardii and P. smithii treatments. Comparing Collembola isotope values in AMF vs. AMF-suppressed treatments, our data show that both AMF and SF were consumed in these experimental soil environments.     

Mycorrhizal fungi mitigate nitrogen losses of an experimental grassland by facilitating plant uptake and soil microbial immobilization

Nitrogen (N) is one of the most limited nutrients of terrestrial ecosystems, whose losses are prevented in tightly coupled cycles in finely tuned systems. Global change-induced N enrichment through atmospheric deposition and application of vast amounts of fertilizer are now challenging the terrestrial N cycle. Arbuscular mycorrhizal fungi (AMF) are known drivers of plant-soil nutrient fluxes, but a comprehensive assessment of AMF involvement in N cycling under global change is still lacking. Here, we simulated N enrichment by fertilization (low/high) in experimental grassland microcosms under greenhouse conditions in the presence or absence of AMF and continuously monitored different N pathways over nine months. We found that high N enrichment by fertilization decreased the relative abundance of legumes and the plant species dominating the plant community changed from grasses to forbs in the presence of AMF, based on aboveground biomass. The presence of AMF always maintained plant N:phosphorus (P) ratios between 14 and 16, no matter how the soil N availability changed. Shifts in plant N:P ratios due to the increased plant N and P uptake might thus be a primary pathway of AMF altering plant community composition. Furthermore, we constructed a comprehensive picture of AMF’s role in N cycling, highlighting that AMF reduced N losses primarily by mitigating N leaching, while N₂O emissions played a marginal role. Arbuscular mycorrhizal fungi reduced N₂O emissions directly through the promotion of N₂O-consuming denitrifiers. The underlying mechanism for reducing N leaching is mainly the AMF-mediated improved nutrient uptake and AMF-associated microbial immobilization. Our results indicate that synergies between AMF and other soil microorganisms cannot be ignored in N cycling and that the integral role of AMF in N cycling terrestrial ecosystems can buffer the upcoming global changes.     

Specific response of fungal and bacterial residues to one-season tillage and repeated slurry application in a permanent grassland soil

The dynamics of fungal and bacterial residues to a one-season tillage event in combination with manure application in a grassland soil are unknown. The objectives of this study were (1) to assess the effects of one-season tillage event in two field trials on the stocks of microbial biomass, fungal biomass, microbial residues, soil organic C (SOC) and total N in comparison with permanent grassland; (2) to determine the effects of repeated manure application to restore negative tillage effects on soil microbial biomass and residues. One trial was started 2 years before sampling and the other 5 years before sampling. Mouldboard ploughing decreased the stocks of SOC, total N, microbial biomass C, and microbial residues (muramic acid and glucosamine), but increased those of the fungal biomarker ergosterol in both trials. Slurry application increased stocks of SOC and total N only in the short-term, whereas the stocks of microbial biomass C, ergosterol and microbial residues were generally increased in both trials, especially in combination with tillage. The ergosterol to microbial biomass C ratio was increased by tillage, and decreased by slurry application in both trials. The fungal C to bacterial C ratio was generally decreased by these two treatments. The metabolic quotient qCO₂ showed a significant negative linear relationship with the microbial biomass C to SOC ratio and a significant positive relationship with the soil C/N ratio. The ergosterol to microbial biomass C ratio revealed a significant positive linear relationship with the fungal C to bacterial C ratio, but a negative one with the SOC content. Our results suggest that slurry application in grassland soil may promote SOC storage without increasing the role of saprotrophic fungi in soil organic matter dynamics relative to that of bacteria.     

Microbial community PLFA and PHB responses to ecosystem restoration in tallgrass prairie soils

Native North American prairie grasslands are renowned for the richness of their soils, having excellent soil structure and very high organic content and microbial biomass. In this study, surface soils from three prairie restorations of varying ages and plant community compositions were compared with a nearby undisturbed native prairie remnant and a cropped agricultural field in terms of soil physical, chemical and microbial properties. Soil moisture, organic matter, total carbon, total nitrogen, total sulfur, C:N, water-holding capacity and microbial biomass (total PLFA) were significantly greater (p<0.05) in the virgin prairie remnant as well as the two long-term (21 and 24 year) prairie restorations, compared with the agricultural field and the restoration that was begun more recently (7 years prior to sampling). Soil bulk density was significantly greater (p<0.05) in the agricultural and recently restored sites. In most cases, the soil quality indicators and microbial community structures in the restoration sites were intermediate between those of the virgin prairie and the agricultural sites. Levels of poly-β-hydroxybutyrate (PHB) and PLFA indicators of nutritional stress were significantly greater (p<0.05) in the agricultural and recent restoration sites than in the long-term restorations or the native prairie. Samples could be assigned to the correct site by discriminant analysis of the PLFA data, with the exception that the two long-term restoration sites overlapped. Redundancy analysis showed that prairie age (p<0.005) was the most important environmental factor in determining the PLFA microbial community composition, with C:N (p<0.015) also being significant. These findings demonstrate that prairie restorations can lead to improved quality of surface soils. We predict that the conversion of farmland into prairie will shift the soil quality, microbial community biomass and microbial community composition in the direction of native prairies, but with the restoration methods tested it may take many decades to approach the levels found in a virgin prairie throughout the soil profile.     

Strong bottom-up and weak top-down effects in soil: nematode-parasitized insects and nematode-trapping fungi

The soils of the Bodega Marine Reserve (BMR, Sonoma County, California) contain many nematode-trapping fungi and many ghost moth larvae parasitized by entomopathogenic nematodes. The current study determined whether these nematode-parasitized moth larvae, which can produce very large numbers of nematodes, enhanced the population densities of nematode-trapping fungi and whether the fungi trapped substantial numbers of nematodes emerging and dispersing from moths. Wax moths were used in place of ghost moths because the former are easier to obtain. When nematode-parasitized moth larvae were added to laboratory microcosms containing BMR field soil, the population densities of four nematode-trapping fungi increased substantially. The greatest increase in population density was by Arthrobotrys oligospora, which uses adhesive networks to capture nematodes. A. oligospora population density increased about 10 times when the added moth larvae were parasitized by the nematode Heterorhabditis marelatus and about 100 times when added moth larvae were parasitized by the nematode Steinernema glaseri. Other trapping fungi endemic to the soil and enhanced by nematode-parasitized moth larvae included Myzocytium glutinosporum, Drechslerella brochopaga, and Gamsylella gephyropaga, which produce adhesive spores, constricting rings, and adhesive branches, respectively. The data suggest that the previously documented abundance and diversity of nematode-trapping fungi in BMR soil can be explained, at least in part, by nematode-parasitized insects, although that inference requires further studies with ghost moths. The strong bottom-up enhancement of nematode-trapping fungi was not matched by a strong top-down suppression of nematodes, i.e. the fungi trapped fewer than 30% of dispersing nematodes.