Changes in soil fungal:bacterial biomass ratios following reductions in the intensity of management of an upland grassland

In this study we examined the effect on soil fungal:bacterial biomass ratios of withholding fertiliser, lime, and sheep-grazing from reseeded upland grassland. The cessation of fertiliser applications on limed and grazed grassland resulted in a reduction in soil pH from 5.4 to 5.1. The cessation of fertiliser applications and liming on grazed grassland resulted in a fall in pH from 5.4 to 4.7, whereas withholding fertiliser and lime and the removal of grazing resulted in a further reduction to pH 4.5. Substrate-induced respiration was reduced in the unfertilised grazed (21%; P<0.01) and unfertilised ungrazed (36%; P<0.001) treatments. Bacterial substrate-induced respiration and bacterial fatty acids were unaffected by the treatments. The relative abundance of the fungal fatty acid 18:26 increased by 39 and 72% (P<0.05) in the limed grazed and unfertilised grazed treatments, respectively. Fungal substrate-induced respiration increased in the limed grazed (18%) and unfertilised grazed (65%; P<0.05) treatments. The ratio of 18:26: bacterial fatty acids was correlated with the ratio of fungal:bacterial substrate-induced respiration (r=0.69; P<0.001).     

The diet and food preferences of Onychiurus procampatus (Collembola) from upland grassland soils

Specimens of Onychiurus procampatus (Collembola) observed grazing on fungal mycelia were collected from the surface soil of three differently manged upland grasslands. A general trend of an increasing proportion of Collembola with a full gut was found along a gradient of reduced sheep management intensity, which was correlated with increased fungal biomass. In the laboratory, this collembolan showed a consistent order of preference for the mycelium of seven common fungal species isolated from the field sites.     

Temporal variability in plant and soil nitrogen pools in a high-Arctic ecosystem

This study determined temporal variability in N pools, both aboveground and belowground, across two contrasting plant communities in high-Arctic Spitsbergen, Svalbard (78°N). We measured N pools in plant material, soil microbial biomass and soil organic matter in moist (Alopecurus borealis dominated) and dry (Dryas octopetala dominated) meadow communities at four times during the growing season. We found that plant, microbial and dissolved inorganic and organic N pools were subject to significant, but surprisingly low, temporal variation that was controlled primarily by changes in temperature and moisture availability over the short growing season. This temporal variability is much less than that experienced in other seasonally cold ecosystems such as alpine tundra where strong seasonal partitioning of N occurs between plant and soil microbial pools. While only a small proportion of the total ecosystem N, the microbial biomass represented the single largest of the dynamic N pools in both moist and dry meadow communities (3.4% and 4.6% of the total ecosystem N pool, respectively). This points to the importance of soil microbial community dynamics for N cycling in high-Arctic ecosystems. Microbial N was strongly and positively related to soil temperature in the dry meadow, but this relationship did not hold true in the wet meadow where other factors such as wetter soil conditions might constrain biological activity. Vascular live belowground plant parts represented the single largest plant N pool in both dry and moist meadow, constituting an average of 1.6% of the total N pool in both systems; this value did not vary across the growing season or between plant communities. Overall, our data illustrate a surprisingly low growing season variability in labile N pools in high-Arctic ecosystems, which we propose is controlled primarily by temperature and moisture.     

Browsing by red deer negatively impacts on soil nitrogen availability in regenerating native forest

Herbivores can have important indirect effects on belowground properties and processes that govern ecosystem form and productivity. A major way that herbivores affect ecosystem productivity is by modifying feedbacks that occur between dominant plants and belowground properties, and especially by changing the rates of nutrient mineralisation. In this study, we examine the effects of browsing by red deer, relative to landscape factors such as variations in microclimate and topography, on soil biological properties and N cycling in regenerating forest in an upland region of the Scottish Highlands. The site at Creag Meagaidh in the Scottish Highlands was sampled three times over the growing season of 2001 and soil cores were taken from four exclosures. Measures of soil biological properties and nitrogen availability were made. The results presented here suggest that although spatial and temporal variation were the dominant factors affecting soil biological properties and nitrogen dynamics in this ecosystem, herbivory also had consistent and significant effects on these measures. Browsing significantly reduced measures of DOC (27%), NO₃⁻ (48%), NH₄⁺ (49%) and N-mineralisation (53%). The microbial C:N ratio was also significantly greater (89%) in browsed than un-browsed areas, suggesting that the removal of deer reduced the extent that the microbial community was N-limited. These significant negative effects of browsing on soil nitrogen cycling have the potential to reduce ecosystem productivity. We conclude that removal of browsing over 14 years has accelerated native woodland regeneration, leading to subsequent increases in soil C- and N-mineralisation, further increasing plant N supply and tree growth in this ecosystem.     

Continuous defoliation of perennial ryegrass (Lolium perenne) and white clover (Trifolium repens) and associated changes in the composition and activity of the microbial population of an upland grassland soil

A microcosm study was conducted to investigate the effect of continuons plant defoliation on the composition and activity of microbial populations in the rhizosphere of perennial ryegrass (Lolium perenne) and white clover (Trifolium repens). Continuons defoliation of ryegrass and clover resulted in sigmficant (P <0.01) increases in soil microbial biomass, although whilst increases were measured from day 2 in soil sown with clover significant increases were only seen from day 21 in soil sown with ryegrass. These increases were paralleled, from day 10 onwards, by increases in the numbers of culturable bacteria. Numbers ofPsendomonas spp. also increased in the later stages of the study. No influence on culturable fungal populations was detected. Whilst shifts in the composition of the microbial populations were measured in response to defoliation there was little effect on microbial activity. No changes in either dehydrogenase activity or microbial respiration in the rhizosphere of ryegrass or clover were measured in response to defoliation, but both dehydrogenase activity and microbial respiration were greater in ryegrass than clover when values over the whole study were combined. Continuous defoliation resulted in significant (P <0.001) reductions in the root dry weight of ryegrass and clover, of the order 19% and 16%, respectively.     

Impact of pasture contamination by copper,chromium, and arsenic timber preservative on soil microbial properties and nematodes

Microbial properties and nematode abundance were measured along a gradient of increasing Cu, Cr, and As concentrations (50–1300 mg Cr kg^-1) in the top 5 cm of a pasture soil contaminated by runoff of preserving liquor from an adjacent timber-treatment plant. Microbial biomass C and N were significantly (P<0.05) lower in contaminated than uncontaminated soils. The amount of microbial biomass C as a percentage of total organic C declined significantly (r ² value with Cr 0.726^*) with increasing contamination, and the ratio of respired C to biomass C was significantly (P<0.05) higher with contamination. Substrate-induced respiration, microbial biomass P, and denitrification declined (r² value with Cr 0.601, 0.833^*, and 0.709^*, respectively) with increasing contamination. Increasing contamination had no effect on prokaryote substrate-induced respiration but eukaryote: eukaryote substrate-induced respiration declined significantly (r ² value with Cr 0.722^*). Accordingly, the ratio of prokaryote substrate-induced respiration increased significantly (r ² value with Cr 0.799^*) with contamination. There was a significant (r ² value with Cr 0.872^*) hyperbolic relationship between sulphatase activity and contamination, with activity declining by approximately 80% at >1000 mg Cr kg^-1. Increasing contamination had no effect on basal respiration, dimethyl sulphoxide reduction, and phosphatase, urease, and invertase activities. Numbers of plant-associated nematodes declined significantly (r ² value with Cr 0.780^*) with contamination. On a percentage basis, plant-feeding nematodes predominated in less contaminated soils, whereas bacterial-feeding and predatory nematodes predominated in heavily contaminated soils. The use of the fumigation—incubation procedure for measurement of microbial biomass C in heavy-metal contaminated soils is discussed.     

Direct and indirect effects of nitrogen deposition on litter decomposition

Elevated nitrogen (N) deposition can affect litter decomposition directly, by raising soil N availability and the quantity and quality of litter inputs, and indirectly by altering plant community composition. We investigated the importance of these controls on litter decomposition using litter bags placed in annual herb based microcosm ecosystems that had been subject to two rates of N deposition (which raised soil inorganic N availability and stimulated litter inputs) and two planting regimes, namely the plant species compositions of low and high N deposition environments. In each microcosm, we harvested litter bags of 10 annual plant species, over an 8-week period, to determine mass loss from decomposition. Our data showed that species differed greatly in their decomposability, but that these differences were unlikely to affect decomposition at the ecosystem level because there was no correlation between a species’ decomposability and its response to N deposition (measured as population seed production under high N, relative to low N, deposition). Litter mass loss was ～2% greater in high N deposition microcosms. Using a comprehensive set of measurements of the microcosm soil environments, we found that the most statistically likely explanation for this effect was increased soil enzyme activity (cellobiosidase, β-glucosidase and β-xylosidase), which appears to have occurred in response to a combination of raised soil inorganic N availability and stimulated litter inputs. Our data indicate that direct effects of N deposition on litter input and soil N availability significantly affected decomposition but indirect effects did not. We argue that indirect effects of changes to plant species composition could be stronger in natural ecosystems, which often contain a greater diversity of plant functional types than those considered here.     

Structure and function of the soil microbial community in microhabitats of a heavy metal polluted soil

 Particle-size fractionation of a heavy metal polluted soil was performed to study the influence of environmental pollution on microbial community structure, microbial biomass, microbial residues and enzyme activities in microhabitats of a Calcaric Phaeocem. In 1987, the soil was experimentally contaminated with four heavy metal loads: (1) uncontaminated controls; (2) light (300 ppm Zn, 100 ppm Cu, 50 ppm Ni, 50 ppm V and 3 ppm Cd); (3) medium; and (4) heavy pollution (two- and threefold the light load, respectively). After 10 years of exposure, the highest concentrations of microbial ninhydrin-reactive nitrogen were found in the clay (2–0.1 μm) and silt fractions (63–2 μm), and the lowest were found in the coarse sand fraction (2,000–250 μm). The phospholipid fatty acid analyses (PLFA) and denaturing gradient gel electrophoresis (DGGE) separation of 16S rRNA gene fragments revealed that the microbial biomass within the clay fraction was predominantly due to soil bacteria. In contrast, a high percentage of fungal-derived PLFA 18 : 2ω6 was found in the coarse sand fraction. Bacterial residues such as muramic acid accumulated in the finer fractions in relation to fungal residues. The fractions also differed with respect to substrate utilization: Urease was located mainly in the <2 μm fraction, alkaline phosphatase and arylsulfatase in the 2–63 μm fraction, and xylanase activity was equally distributed in all fractions. Heavy metal pollution significantly decreased the concentration of ninhydrin-reactive nitrogen of soil microorganisms in the silt and clay fraction and thus in the bulk soil. Soil enzyme activity was reduced significantly in all fractions subjected to heavy metal pollution in the order arylsulfatase >phosphatase >urease >xylanase. Heavy metal pollution did not markedly change the similarity pattern of the DGGE profiles and amino sugar concentrations. Therefore, microbial biomass and enzyme activities seem to be more sensitive than 16S rRNA gene fragments and microbial amino-sugar-N to heavy metal treatment. Received: 21 January 2000     

Functional aspects of soil animal diversity in agricultural grasslands

There has been recent interest in the characterization of soil biodiversity and its function in agricultural grasslands. Much of the interest has come from the need to develop grassland management strategies directed at manipulating the soil biota to encourage a greater reliance on ecosystem self-regulation. This review summarises information on selected groups of soil animals in grasslands, the factors influencing their abundance, diversity and community structure and their relationships to the functioning and stability of grassland ecosystems. Observations on the impacts of agricultural managements on populations and communities of soil fauna and their interactions confirm that high input, intensively managed systems tend to promote low diversity while lower input systems conserve diversity. It is also evident that high input systems favour bacterial-pathways of decomposition, dominated by labile substrates and opportunistic, bacterial-feeding fauna. In contrast, low-input systems favour fungal-pathways with a more heterogeneous habitat and resource leading to domination by more persistent fungal-feeding fauna. In view of this, we suggest that low input grassland farming systems are optimal for increasing soil biotic diversity and hence self-regulation of ecosystem function. Research is needed to test the hypothesis that soil biodiversity is positively associated with stability, and to elucidate relationships between productivity, community integrity and functioning of soil biotic communities.     

The measurement of soil fungal:bacterial biomass ratios as an indicator of ecosystem self-regulation in temperate meadow grasslands

 There is much interest in the development of agricultural land management strategies aimed at enhancing reliance on ecosystem self-regulation rather than on artificial inputs such as fertilisers and pesticides. This study tested the usefulness of measures of soil microbial biomass and fungal:bacterial biomass ratios as indicators of effective conversion from an intensive grassland system, reliant mainly on fertilisers for crop nutrition, to a low-input system reliant mainly on self-regulation through soil biological pathways of nutrient turnover. Analysis of soils from a wide range of meadow grassland sites in northern England, along a gradient of long-term management intensity, showed that fungal:bacterial biomass ratios (measured by phospholipid fatty acid analysis; PLFA) were consistently and significantly higher in the unfertilised than the fertilised grasslands. There was also some evidence that microbial biomass, measured by chloroform fumigation and total PLFA, was higher in the unfertilised than in the fertilised grasslands. It was also found that levels of inorganic nitrogen (N), in particular nitrate-N, were significantly higher in the fertilised than in the unfertilised grasslands. However, microbial activity, measured as basal respiration, did not differ between the sites. A field manipulation trial was conducted to determine whether the reinstatement of traditional management on an improved mesotrophic grassland, for 6 years, resulted in similar changes in the soil microbial community. It was found that neither the cessation of fertiliser applications nor changes in cutting and grazing management significantly affected soil microbial biomass or the fungal:bacterial biomass ratio. It is suggested that the lack of effects on the soil microbial community may be related to high residual fertility caused by retention of fertiliser N in the soil. On the basis of these results it is recommended that following the reinstatement of low-input management, the measurement of a significant increase in the soil fungal:bacterial biomass ratio, and perhaps total microbial biomass, may be an indicator of successful conversion to a grassland system reliant of self-regulation. Received: 4 May 1998