C and N natural abundance of the soil microbial biomass

Stable isotope analysis is a powerful tool in the study of soil organic matter formation. It is often observed that more decomposed soil organic matter is ¹³C, and especially ¹⁵N-enriched relative to fresh litter and recent organic matter. We investigated whether this shift in isotope composition relates to the isotope composition of the microbial biomass, an important source for soil organic matter. We developed a new approach to determine the natural abundance C and N isotope composition of the microbial biomass across a broad range of soil types, vegetation, and climates. We found consistently that the soil microbial biomass was ¹⁵N-enriched relative to the total (3.2 ‰) and extractable N pools (3.7 ‰), and ¹³C-enriched relative to the extractable C pool (2.5 ‰). The microbial biomass was also ¹³C-enriched relative to total C for soils that exhibited a C3-plant signature (1.6 ‰), but ¹³C-depleted for soils with a C4 signature (−1.1 ‰). The latter was probably associated with an increase of annual C3 forbs in C4 grasslands after an extreme drought. These findings are in agreement with the proposed contribution of microbial products to the stabilized soil organic matter and may help explain the shift in isotope composition during soil organic matter formation.     

Diversity of dead wood inhabiting fungi and bryophytes in semi-natural beech forests in Europe

Saproxylic organisms are among the most threatened species in Europe and constitute a major conservation problem because they depend on the most important forestry product - dead wood. Diversity of fungal and bryophyte communities occurring on dead beech trees was analyzed in five European countries (Slovenia, Hungary, The Netherlands, Belgium and Denmark) considering tree level species richness (TLSR), country level species richness (CLSR), frequency distributions of species, occurrence of threatened species and relations between TLSR and decay stage, tree size and countries. Altogether 1009 trees were inventoried in 19 beech dominated forest reserves.The number of fungi in the full dataset was approximately three times larger (456 versus 161 species) and the proportion of low frequent species was higher than among bryophytes. The species richness of bryophytes and fungi was significantly different among countries considering both TLSR and CLSR. In addition the diversity patterns deviated considerably between the two groups of organisms. Slovenian sites appeared to be biodiversity hotspots for bryophytes characterized by high TLSR and CLSR and a high fraction of threatened species. Hungarian sites had somewhat lower bryophyte diversity, while the Atlantic region had deteriorated assemblages. Fungal species richness was very high in Denmark, but the Hungarian and Slovenian sites were richer in threatened and low frequency species. Tree size was better able to explain variation in TLSR in both organism groups than decay stage. TLSR was found to vary significantly between countries but the difference was most considerable in the case of bryophytes.The diversity patterns of both organism groups along the investigated geographical gradient appear to be influenced by both climatic and management related factors (forest history, dead wood availability and continuity, habitat fragmentation). There is no doubt that an increase in the abundance of dead wood in European beech forests will benefit diversity of saproxylic fungi and bryophytes, especially if a continuous presence of large diameter logs are secured within individual stands.     

Distribution and abundance of soil fungi in Antarctica at sites on the Peninsula, Ross Sea Region and McMurdo Dry Valleys

Fungal abundance and diversity were studied from 245 soil samples collected in 18 distinct ice-free locations in Antarctica including areas in the McMurdo Dry Valleys, Ross Sea Region, and the Antarctic Peninsula. Cultivable fungal abundance in soil was found to be most positively correlated with percent carbon and nitrogen based on a Spearman’s rank correlation test of six soil parameters. Soil moisture and C/N ratio were also positively correlated with fungal abundance while pH and conductivity were negatively correlated. These results suggest that nutrient limitations in these highly oligotrophic environments are a primary factor in determining the distribution and abundance of indigenous fungi. Other effects of the extreme Antarctic environment likely affect fungi indirectly by limiting the distribution and abundance of plant-derived sources of carbon.     

Soil profile distribution of total C content and natural abundance of 13C in two volcanic soils subjected to crop residue burning versus crop residue retention

Abstract

Soil organic carbon (SOC) plays a key role in crop productivity and soil quality. Conservation agriculture has a positive effect on SOC accumulation in the surface soil horizons, but little information is available regarding the effect of the removal of crop residues by burning. This study aimed to assess the impact of different types of crop residue management practices on the total C distribution and natural abundance of ¹³C (‰, δ¹³C). Two volcanic soils, located in the Mediterranean temperate zone of Southern Chile, were studied: an Ultisol (Collipulli Series, CPL) and an Andisol (Santa Bárbara Series, SBA). Both soils had been cultivated under direct-drilling and a typical annual crop rotation system for a long period of time. Two different types of crop residue management practices were imposed in both soils: (i) crop residue burning (CPL-B; SBA-B) and (ii) crop residue retention over the soil (CPL-R; SBA-R), corresponding to treatments B and R, respectively. Soil profile distribution of the C content and natural abundance of ¹³C were analysed for bulk soils (down to 100 cm depth) and three particle-size fractions of the soils (down to 20 cm of soil depth): (a) ≤ 53 µm, (b) 53-212 µm and (c) ≥ 212 µm. It was found that the effect of crop residue management can be observed in the variations of C content and δ¹³C in the soil profile in both volcanic soils. Crop residue burning (B treatment) increased the C content in bulk soil and the particle-size fractions. On the other hand, soil organic matter of crop residue retention (R treatment) showed higher natural abundance of ¹³C (δ¹³C) compared with residue burning (B treatment) in the two volcanic soils. R treatment enriched the particle-size fractions (except ≥ 212 µm fraction of CPL soil) with ¹³C. Factors that could account for these findings are also discussed here.     

Compartmentalization of the soil animal food web as indicated by dual analysis of stable isotope ratios (N/N and C/C)

The soil animal food web has become a focus of recent ecological research but trophic relationships still remain enigmatic for many taxa. Analysis of stable isotope ratios of N and C provides a powerful tool for disentangling food web structure. In this study, animals, roots, soil and litter material from a temperate deciduous forest were analysed. The combined measurement of δ¹⁵N and δ¹³C provided insights into the compartmentalization of the soil animal food web. Leaf litter feeders were separated from animals relying mainly on recent belowground carbon resources and from animals feeding on older carbon. The trophic pathway of leaf litter-feeding species appears to be a dead end, presumably because leaf litter feeders (mainly diplopods and oribatid mites) are unavailable to predators due to large size and/or strong sclerotization. Endogeic earthworms that rely on older carbon also appear to exist in predator-free space. The data suggest that the largest trophic compartment constitutes of ectomycorrhizal feeders and their predators. Additionally, there is a smaller trophic compartment consisting of predators likely feeding on enchytraeids and potentially nematodes.     

Carbon fluxes in soil food webs of increasing complexity revealed by C labelling and C natural abundance

Soil food webs are mainly based on three primary carbon (C) sources: root exudates, litter, and recalcitrant soil organic matter (SOM). These C sources vary in their availability and accessibility to soil organisms, which could lead to different pathways in soil food webs. The presence of three C isotopes (¹²C, ¹³C and ¹⁴C) offers an unique opportunity to investigate all three C sources simultaneously. In a microcosm experiment we studied the effect of food web complexity on the utilization of the three carbon sources. We choose an incomplete three factorial design with (i) living plants, (ii) litter and (iii) food web complexity. The most complex food web consisted of autochthonous microorganisms, nematodes, collembola, predatory mites, endogeic and anecic earthworms. We traced C from all three sources in soil, in CO₂ efflux and in individual organism groups by using maize grown on soil developed under C₃ vegetation and application of ¹⁴C labelled ryegrass shoots as a litter layer. The presence of living plants had a much greater effect on C pathways than food web complexity. Litter decomposition, measured as ¹⁴CO₂ efflux, was decreased in the presence of living plants from 71% to 33%. However, living plants increased the incorporation of litter C into microbial biomass and arrested carbon in the litter layer and in the upper soil layer. The only significant effect of food web complexity was on the litter C distribution in the soil layers. In treatments with fungivorous microarthropods (Collembola) the incorporation of litter carbon into mineral soil was reduced. Root exudates as C source were passed through rhizosphere microorganisms to the predator level (at least to the third trophic level). We conclude that living plants strongly affected C flows, directly by being a source of additional C, and indirectly by modifying the existing C flows within the food web including CO₂ efflux from the soil and litter decomposition.     

Spatial variation of soil organic carbon concentrations and stable isotopic composition in 1-ha plots of forest and pasture in Costa Rica: implications for the natural abundance technique

The goal of this study was to examine spatial variation of soil organic C and its stable isotopic composition (δ¹³C) in 1-ha plots of mature rain forest and a cattle pasture dominated by C4 grasses in Costa Rica. Soil samples were taken from 80 mapped locations per plot and analyzed for organic C and δ¹³C. The range of values for soil C concentrations was similar between forest and pasture, although the mean values were higher in the forest. δ¹³C was narrowly constrained in the forest (from −27.96 to −26.09‰) but varied from −15.09 to −28.59‰ in the pasture. Variograms revealed spatial autocorrelation of soil C and δ¹³C in the pasture and organic C concentration in the forest soil. The large range and spatial variability of δ¹³C in the pasture site may be due to varying contributions of C3 and C4 vegetation to the soil C pool, which may limit the usefulness of the natural abundance technique as a precise tracer of soil C dynamics in this pasture.     

Relationships between C and N availability, substrate age, and natural abundance C and N signatures of soil microbial biomass in a semiarid climate

Soil microbial organisms are central to carbon (C) and nitrogen (N) transformations in soils, yet not much is known about the stable isotope composition of these essential regulators of element cycles. We investigated the relationship between C and N availability and stable C and N isotope composition of soil microbial biomass across a three million year old semiarid substrate age gradient in northern Arizona. The δ¹⁵N of soil microbial biomass was on average 7.2‰ higher than that of soil total N for all substrate ages and 1.6‰ higher than that of extractable N, but not significantly different for the youngest and oldest sites. Microbial ¹⁵N enrichment relative to soil extractable and total N was low at the youngest site, increased to a maximum after 55,000 years, and then decreased slightly with age. The degree of ¹⁵N enrichment of microbial biomass correlated negatively with the C:N mass ratio of the soil extractable pool. The δ¹³C signature of soil microbial biomass was 1.4‰ and 4.6‰ enriched relative to that of soil total and extractable pools respectively and showed significant differences between sites. However, microbial ¹³C enrichment was unrelated to measures of C and N availability. Our results confirm that ¹⁵N, but not ¹³C enrichment of soil microbial biomass reflects changes in C and N availability and N processing during long-term ecosystem development.     

Organic matter turnover in soil physical fractions following woody plant invasion of grassland: Evidence from natural C and N

Soil physical structure causes differential accessibility of soil organic carbon (SOC) to decomposer organisms and is an important determinant of SOC storage and turnover. Techniques for physical fractionation of soil organic matter in conjunction with isotopic analyses (δ¹³C, δ¹⁵N) of those soil fractions have been used previously to (a) determine where organic C is stored relative to aggregate structure, (b) identify sources of SOC, (c) quantify turnover rates of SOC in specific soil fractions, and (d) evaluate organic matter quality. We used these two complementary approaches to characterize soil C storage and dynamics in the Rio Grande Plains of southern Texas where C₃ trees/shrubs (δ¹³C=−27‰) have largely replaced C₄ grasslands (δ¹³C=−14‰) over the past 100-200 years. Using a chronosequence approach, soils were collected from remnant grasslands (Time 0) and from woody plant stands ranging in age from 10 to 130 years. We separated soil organic matter into specific size/density fractions and determined their C and N concentrations and natural δ¹³C and δ¹⁵N values. Mean residence times (MRTs) of soil fractions were calculated based on changes in their δ¹³C with time after woody encroachment. The shortest MRTs (average=30 years) were associated with all particulate organic matter (POM) fractions not protected within aggregates. Fine POM (53-250 μm) within macro- and microaggregates was relatively more protected from decay, with an average MRT of 60 years. All silt+clay fractions had the longest MRTs (average=360 years) regardless of whether they were found inside or outside of aggregate structure. δ¹⁵N values of soil physical fractions were positively correlated with MRTs of the same fractions, suggesting that higher δ¹⁵N values reflect an increased degree of humification. Increased soil C and N pools in wooded areas were due to both the retention of older C₄-derived organic matter by protection within microaggregates and association with silt+clay, and the accumulation of new C₃-derived organic matter in macroaggregates and POM fractions.     

Savanna-derived organic matter remaining in arable soils of the South African Highveld long-term mixed cropping: Evidence from C and N natural abundance

Sustainable agriculture requires the formation of new humus from the crops. We utilized ¹³C and ¹⁵N signatures of soil organic matter to assess how rapidly wheat/maize cropping contributed to the humus formation in coarse-textured savanna soils of the South African Highveld. Composite samples were taken from the top 20 cm of soils (Plinthustalfs) cropped for lengths of time varying from 0 to 98 years, after conversion from native grassland savanna (C4). We performed natural ¹³C and ¹⁵N abundance measurements on bulk and particle-size fractions. The bulk soil δ¹³C values steadily decreased from −14.6 in (C4 dominated) grassland to −16.5‰ after 90 years of arable cropping. This δ¹³C shift was attributable to increasing replacement of savanna-derived C by wheat crop (C3) C which dominated over maize (C4) inputs. After calculating the annual C input from the crop yields and the output from literature data, by using a stepwise C replacement model, we were able to correct the soil δ¹³C data for the irregular maize inputs for a period of about one century. Within 90 years of cropping 41-89% of the remaining soil organic matter was crop-derived in the three studied agroecosystems. The surface soil C stocks after 90 years of the wheat/maize crop rotation could accurately be described with the Rothamsted Carbon Model, but modelled C inputs to the soil were very low. The coarse sand fraction reflected temporal fluctuations in ¹³C of the last C₃ or C₄ cropping and the silt fraction evidenced selective erosion loss of old savanna-derived C. Bulk soil ¹⁵N did not change with increasing cropping length. Decreasing δ¹⁵N values caused by fertilizer N inputs with prolonged arable cropping were only detected for the coarse sand fraction. This indicated that the present N fertilization was not retained in stable soil C pool. Clearly, conventional cropping practices on the South African highlands neither contribute to the preservation of old savanna C and N, nor the effective humus reformation by the crops.