Growth measurements of saprotrophic fungi and bacteria reveal differences between canopy and forest floor soils

Canopy-held organic matter develops into a distinct soil system separate from the forest floor in wet temperate coniferous forests, creating a natural microcosm. We distinguished between fungal and bacterial components of the decomposer community in one site with Maple (Acer macrophyllum) and one site with Alder (Alnus rubra) by using direct measurements of growth; acetate incorporation into ergosterol, and leucine incorporation for fungi and bacteria, respectively. The higher organic matter content of the canopy soils correlated with higher fungal growth. The relative importance of fungi, indicated by fungal:bacterial growth ratio, was higher in the canopy soil of the Maple site, while there was no difference in the Alder site. The high C:N ratio of the Maple canopy soil likely contributed to this difference. These results demonstrate a divergence between canopy and forest floor that should be explored to gain insights in decomposer ecology using the natural microcosms that the canopy soils provide.     

Bacterial pH-optima for growth track soil pH, but are higher than expected at low pH

One of the most influential factors determining the growth and composition of soil bacterial communities is pH. However, soil pH is often correlated with many other factors, including nutrient availability and plant community, and causality among factors is not easily determined. If soil pH is directly influencing the bacterial community, this must lead to a bacterial community growth optimised for the in situ pH. Using one set of Iberian soils (46 soils covering pH 4.2-7.3) and one set of UK grassland soils (16 soils covering pH 3.3-7.5) we measured the pH-optima for the growth of bacterial communities. Bacterial growth was estimated by the leucine incorporation method. The pH-optima for bacterial growth were positively correlated with soil pH, demonstrating its direct influence on the soil bacterial community. We found that the pH from a water extraction better matched the bacterial growth optimum compared with salt extractions of soil. Furthermore, we also showed a more subtle pattern between bacterial pH growth optima and soil pH. While closely matched at neutral pHs, pH-optima became higher than the in situ pH in more acid soils, resulting in a difference of about one pH-unit at the low-pH end. We propose that an explanation for the pattern is an interaction between increasing overall bacterial growth with higher pHs and the unimodal pH-response for growth of bacterial communities.     

Induced N-limitation of bacterial growth in soil: Effect of carbon loading and N status in soil

Application of C-rich plant residues can change the soil system from C-limitation for microbial growth to limitation by other nutrients. However, the initial nutrient status of the soil may interact with the added amount of residues in determining limitation. We studied this interactive effect in soils from the Harvard Forest LTER, where annual addition of N since 1988 has resulted in soils with different N-status: No N (Unfertilized), 50 (Low N) and 150 (High N) kg N ha⁻¹. We hypothesized that adding C-rich substrate would change the soil from being C- to being N-limited for bacterial growth and that the extent of N-limitation would be higher with increasing substrate additions, while becoming less evident in soil with increasing N-status. We compared the effect of adding two C-rich substrates, starch (0, 10, 20, 40 mg g⁻¹ soil) and straw (0, 20, 40, 80 mg g⁻¹), incubating the soils for up to 3 and 4 weeks for starch and straw, respectively. Nutrient limitations were studied by measuring bacterial growth 3 days after adding C as glucose and N as NH₄NO₃ in a full factorial design. Initially bacterial growth in all soils was C-limited. As hypothesized, adding C-rich substrates removed the C-limitation, with lower amounts of starch and straw needed in the unfertilized and Low N soils than in the High N soil. Combinations of different N-status of the soil and amendment levels of starch and straw could be found, where bacterial growth appeared close to co-limited both by available C and N. However, at even higher amendment levels, presumable resulting in N-limitation, bacterial growth still responded less by adding N then C-limited soils by adding C. Thus, in a C-limited soil there appeared to be N available immediate for growth, while in an N-limited soil, easily available C was not immediately available.     

Threshold concentration of glucose for bacterial growth in soil

The activity of heterotrophic soil microorganisms is usually limited by the availability and quality of carbon (C). Adding organic substances will thus trigger a microbial response. We studied the response in bacterial growth and respiration after the addition of low amounts of glucose. First we determined if additions of glucose, at concentrations which did not result in an exponential increase in respiration after the lag phase, still stimulated bacterial growth. The second aim was to determine the threshold concentration of glucose needed to induce bacterial growth. Adding glucose-C at 1000 μg g⁻¹ soil resulted in an increased respiration rate, which was stable during 12 h, and then decreased without showing any exponential increase in respiration. Bacterial growth, determined as leucine incorporation, did not change compared to an unamended control during the first 12 h, but then increased to levels 5 times higher than in the control. Thus, after the lag phase, a period with increasing bacterial growth, but at the same time decreasing respiration rates, was found. Similar results, but with a more modest increase in bacterial growth, were found using 500 μg glucose-C g⁻¹ soil. Adding 50–700 μg glucose-C g⁻¹ resulted in increased respiration during 24 h correlating with the addition rate. In contrast, bacterial growth after 24 h was only stimulated by glucose additions >200 μg C g⁻¹ soil. Thus, there was a threshold concentration of added substrate for inducing bacterial growth. Below the threshold concentration growth and respiration appear to be uncoupled.     

Fungal and bacterial recolonisation of acid and alkaline forest soils following artificial heat treatments

The direct response and the short-term recolonisation of soil by fungi and bacteria were studied after heat treatments of a humus soil with high carbon content and low pH, and a calcareous soil with lower carbon content and high pH. Heating was administered using a muffle furnace or an autoclave, with different temperatures and times of heat exposure, after which fresh soil (1%) was added as inoculum. Autoclaved soil showed more marked increases in bacterial growth during the recovery phase than oven-heated soil, and the bacterial growth response was more rapid in calcareous than in humus soil. Fungal growth recovered more rapid and reached values higher than the control in humus soil, while it remained low until the end of the study in calcareous soil. Respiration rate showed similar patterns in both soils. Fungal biomass (ergosterol and PLFA 18:2ω6,9) indicated that fungi benefited by autoclaving in humus soil, while they were disfavoured by this treatment in calcareous soil. The sum of bacterial PLFAs did not change due to heating, but some bacterial PLFAs (e.g. cy17:0) increased in both soils. We propose that the community assembly of the microbial communities after heating were mainly driven by pH, in that the high pH soil selected primarily for bacteria and the low pH soil for fungi.     

Comparison of soil fungal/bacterial ratios in a pH gradient using physiological and PLFA-based techniques

We have compared the total microbial biomass and the fungal/bacterial ratio estimated using substrate-induced respiration (SIR) in combination with the selective inhibition technique and using the phospholipid fatty acid (PLFA) technique in a pH gradient (3.0-7.2) consisting of 53 mature broad-leaved forest soils. A fungal/bacterial biomass index using the PLFA technique was calculated using the PLFA 18:2ω6,9 as an indicator of fungal biomass and the sum of 13 bacterial specific PLFAs as indicator of the bacterial biomass. Good linear correlation (p<0.001) was found between the total microbial biomass estimated with SIR and total PLFAs (totPLFA), indicating that 1 mg biomass-C was equivalent to 130 nmol totPLFA. Both biomass estimates were positively correlated to soil pH. The fungal/bacterial ratio measured using the selective inhibition technique decreased significantly with increasing pH from about 9 at pH 3 to approximately 2 at pH 7, while the fungal/bacterial biomass index using PLFA measurements tended to increase slightly with increasing soil pH. Good correlation between the soil content of ergosterol and of the PLFA 18:2ω6,9 indicated that the lack of congruency between the two methods in estimating fungal/bacterial ratios was not due to PLFA 18:2ω6,9-related non-fungal structures to any significant degree. Several PLFAs were strongly correlated to soil pH (R² values >0.8); for example the PLFAs 16:1ω5 and 16:1ω7c increased with increasing soil pH, while i16:0 and cy19:0 decreased. A principal component analysis of the total PLFA pattern gave a first component that was strongly correlated to soil pH (R²=0.85, p<0.001) indicating that the microbial community composition in these beech/beech-oak forest soils was to a large extent determined by soil pH.     

The use of phospholipid fatty acid analysis to estimate bacterial and fungal biomass in soil

The cell content of 12 bacterial phospholipid fatty acids (PLFA) was determined in bacteria extracted from soil by homogenization/centrifugation. The bacteria were enumerated using acridine orange direct counts. An average of 1.40×10^-17 mol bacterial PLFA cell^-1 was found in bacteria extracted from 15 soils covering a wide range of pH and organic matter contents. With this factor, the bacterial biomass based on PLFA analyses of whole soil samples was calculated as 1.0–4.8 mg bacterial C g^-1 soil C. The corresponding range based on microscopical counts was 0.3–3.0 mg bacterial C g^-1 soil C. The recovery of bacteria from the soils using homogenization/centrifugation was 2.6–16% (mean 8.7%) measured by PLFA analysis, and 12–61% (mean 26%) measured as microscopical counts. The soil content of the PLFA 18:26 was correlated with the ergosterol content (r=0.92), which supports the use of this PLFA as an indicator of fungal biomass. The ratio 18:26 to bacterial PLFA is therefore suggested as an index of the fungal:bacterial biomass ratio in soil. An advantage with the method based on PLFA analyses is that the same technique and even the same sample is used to determine both fungi and bacteria. The fungal:bacterial biomass ratio calculated in this way was positively correlated with the organic matter content of the soils (r=0.94).     

Common and distinguishing features of the bacterial and fungal communities in biological soil crusts and shrub root zone soils

Soil microbial communities in dryland ecosystems play important roles as root associates of the widely spaced plants and as the dominant members of biological soil crusts (biocrusts) colonizing the plant interspaces. We employed rRNA gene sequencing (bacterial 16S/fungal large subunit) and shotgun metagenomic sequencing to compare the microbial communities inhabiting the root zones of the dominant shrub, Larrea tridentata (creosote bush), and the interspace biocrusts in a Mojave desert shrubland within the Nevada Free Air CO₂ Enrichment (FACE) experiment. Most of the numerically abundant bacteria and fungi were present in both the biocrusts and root zones, although the proportional abundance of those members differed significantly between habitats. Biocrust bacteria were predominantly Cyanobacteria while root zones harbored significantly more Actinobacteria and Proteobacteria. Pezizomycetes fungi dominated the biocrusts while Dothideomycetes were highest in root zones. Functional gene abundances in metagenome sequence datasets reflected the taxonomic differences noted in the 16S rRNA datasets. For example, functional categories related to photosynthesis, circadian clock proteins, and heterocyst-associated genes were enriched in the biocrusts, where populations of Cyanobacteria were larger. Genes related to potassium metabolism were also more abundant in the biocrusts, suggesting differences in nutrient cycling between biocrusts and root zones. Finally, ten years of elevated atmospheric CO₂ did not result in large shifts in taxonomic composition of the bacterial or fungal communities or the functional gene inventories in the shotgun metagenomes.     

Hitchhikers on the fungal highway: The helper effect for bacterial migration via fungal hyphae

Previous work in our laboratory showed that several bacterial strains, either singly or in association with other bacteria (community migration), were capable of migrating together with the saprotrophic fungus Lyophyllum sp. strain Karsten through soil microcosms. A possible involvement of the type III secretion system (TTSS) in migration was indicated. In this study, we addressed the basis of the community migration, which might lie in a migration helper effect exerted by particular single-strain migrators on other members of the community. Different culturing (plating) as well as culture-independent (PCR-DGGE) methods were applied to assess the effects of putative bacterial helpers in the migration. We used, as a model, the migration-proficient Burkholderia terrae BS001 as the canonical helper strain. PCR-DGGE analysis of the soil system with or without added strain BS001, revealed that the latter consistently stimulated the migration of different bacterial species through the soil. This was observed both following introduction of the organism to a bacterial community from soil and on the basis of a similar organism that was naturally present. One strain, Dyella japonica BS003, was identified as an avid comigrator with B. terrae BS001, although it appeared to lag behind the latter strain in its migration speed. Further examination of the B. terrae BS001/D. japonica BS003 interaction at Lyophyllum sp. strain Karsten hyphae showed that the presence of the D. japonica strain did not negatively affect the growth and migration of B. terrae BS001 with the fungus. A biofilm of B. terrae BS001 was formed on the fungal hyphal front, and we postulate a role for this biofilm in the migration helper effect.     

Effects of soil frost on growth, composition and respiration of the soil microbial decomposer community

Most climate change scenarios predict that the variability of weather conditions will increase in coming decades. Hence, the frequency and intensity of freeze-thaw cycles in high-latitude regions are likely to increase, with concomitant effect on soil carbon biogeochemistry and associated microbial processes. To address this issue we sampled riparian soil from a Swedish boreal forest and applied treatments with variations in four factors related to soil freezing (temperature, treatment duration, soil water content and frequency of freeze-thaw cycles), at three levels in a laboratory experiment, using a Central Composite Face-centred (CCF) experimental design. We then measured bacterial (leucine incorporation) and fungal (acetate in ergosterol incorporation) growth, basal respiration, soil microbial phospholipid fatty acid (PLFA) composition, and concentration of dissolved organic carbon (DOC). Fungal growth was higher in soil exposed to freeze-thawing perturbations and freezing temperatures of −6 °C and −12 °C, than under more constant conditions (steady 0 °C). The opposite pattern was found for bacteria, resulting in an increasing fungal-to-bacterial growth ratio following more intensive winter conditions. Soil respiration increased with water content, decreased with treatment duration and appeared to mainly be driven by treatment-induced changes in the DOC concentration. There was a clear shift in the PLFA composition at 0 °C, compared with the two lower temperatures, with PLFA markers associated with fungi as well as a number of unsaturated PLFAs being relatively more common at 0 °C. Shifts in the PLFA pattern were consistent with those expected for phenotypic plasticity of the cell membrane to low temperatures. There were small declines in PLFA concentrations after freeze-thawing and with longer durations. However, the number of freeze-thaw events had no effect on the microbiological variables. The findings suggest that the higher frequency of freeze-thaw events predicted to follow the global warming will likely have a limited impact on soil microorganisms.     

Do growth yield efficiencies differ between soil microbial communities differing in fungal:bacterial ratios? Reality check and methodological issues

Soil communities dominated by fungi such as those of no-tillage (NT) agroecosystems are often associated with greater soil organic matter (SOM) storage. This has been attributed in part to fungi having a higher growth yield efficiency (GYE) compared to bacteria. That is, for each unit of substrate C utilized, fungi invest a greater proportion into biomass and metabolite production than do bacteria. The assumption of higher fungal efficiency may be unfounded because results from studies in which fungal and bacterial efficiencies have been characterized are equivocal and because few studies have measured microbial GYE directly. In this study, we measured microbial GYE in agricultural soils by following ¹³C-labeled glucose loss, total CO₂-C, and ¹³CO₂-C evolution at 2 h intervals for 20 h in two experiments (differing in N amendment levels) in which the fungal:bacterial biomass ratios (F:B) were manipulated. No differences in efficiency were observed for communities with high versus low F:B in soils with or without added inorganic N. When calculated using ¹³CO₂-C (in contrast to total CO₂-C) evolution, growth yield efficiencies of soils having high and low F:B were 0.69±0.01 and 0.70±0.01, respectively. When soils were amended with N, soils with high and low F:B had growth yield efficiencies of 0.78±0.01 and 0.76±0.01, respectively. Our experiments do not support the widely held assumption that soil fungi have greater growth efficiency than soil bacteria. Thus, claims of greater fungal efficiency may be unsubstantiated and should be evoked cautiously when explaining the mechanisms underlying greater C storage and slower C turnover in fungal-dominated soils.     

Novel antibiotics as inhibitors for the selective respiratory inhibition method of measuring fungal:bacterial ratios in soil

The use of the selective inhibition (SI) method for measuring fungal:bacterial ratios may be limited due to biocide selectivity and the overlap of antibiotic activity. This study evaluated novel pairs of antibiotics for their specificity in soils of different origins and their potential reduction in inhibition of non-target organisms. Four soils selected for this study were from a semi-arid shrub-steppe, a loblolly pine forest and two grassland sites (restored and farmed prairie plots). Three bactericides were tested: oxytetracycline hydrochloride, streptomycin sulphate, and bronopol. Three fungicides were tested: captan, ketoconazole, and nystatin. The inhibitor additivity ratio and fungal:bacterial ratios were calculated from control and treated soils where inhibition was measured as CO₂ respiration reduction with biocides. We were able to minimize non-target inhibition by the antibiotics to <5% and thus calculate reliable fungal:bacterial ratios using captan to inhibit fungi in all four soils, and bronopol to inhibit bacteria in three of the four soils. The most successful bactericide in the restored prairie was oxytetracycline-HCl. Our results demonstrate that application of novel antibiotics is not uniformly successful in soils of different origin and that the SI technique requires more than just optimization of antibiotic concentration; it also requires optimization of antibiotic selection.     

Particle size fractionation of fungal and bacterial biomass in subalpine grassland and forest soils

Characterization of soil aggregates according to particle size fractions is a useful tool in process-oriented research into soil organic matter and biological properties. Substrate-induced respiration (SIR) inhibition was used to quantify microbial, fungal and bacterial biomass in particle size fractions of soils ranging from forest to grassland in a subalpine region of central Taiwan. In addition, ergosterol content was determined in the same samples to verify fungal biomass measured by SIR inhibition technique. Surface soil (0–10 cm) was fractionated into four particle size fractions: coarse sand (250–2000 μm), fine sand (53–250 μm), silt (2–53 μm) and clay (0.2–2 μm). The larger sized fractions (>250 μm and 53–250 μm) contained higher levels of fungal ergosterol than the smaller sized ones (2–53 μm and 0.2–2 μm). The largest particle size fraction (250–2000 μm) from all studied habitats showed the highest level of microbial biomass, with no clear trend in microbial biomass level among the other size fractions. SIR-calculated fungal biomass level and ergosterol converted fungal biomass content were positively correlated (r=0.71, p<0.05), and such correlation decreased as biomass levels were high. Ratios of fungi to bacteria ranged between 0.6 and 1.3 in fractions obtained in this study. This study indicates a high variability of microbial (fungal and bacterial) biomass level among particle size fractions in soil, and that the large-sized fractions tend to contain a high level of microbial biomass in a given ecosystem.     

Changes in soil pH due to the storage of soils

Abstract. The pH of soil samples was remeasured after storage for 20 years in the laboratory. The pH decreases were minor in acid to neutral soils (-0.3), but greater in alkaline soils (-0.63). The pH differences were statistically significant only for alkaline soils. The decreases of pH with time are probably mainly due to the decomposition of organic matter, the CO₂ produced, the hydroscopic water and the presence of CaCO₃.     

Soil carbon content drives the biogeographical distribution of fungal communities in the black soil zone of northeast China

Black soils (Mollisols) are one of the most important soil resources for maintaining food security in China, and they are mainly distributed in northeast China. A previous comprehensive study revealed the biogeographical distribution patterns of bacterial communities in the black soil zone. In this study, we used the same soil samples and analyzed the 454 pyrosequencing data for the nuclear ribosomal internal transcribed spacer (ITS) region to examine the fungal communities in these black soils. A total of 220,812 fungal ITS sequences were obtained from 26 soil samples that were collected across the black soil zone. These sequences were classified into at least 5 phyla, 20 classes, greater than 70 orders and over 350 genera, suggesting a high fungal diversity across the black soils. The diversity of fungal communities and distribution of several abundant fungal taxa were significantly related to the soil carbon content. Non-metric multidimensional scaling and canonical correspondence analysis plots indicated that the fungal community composition was most strongly affected by the soil carbon content followed by soil pH. This finding differs from the bacterial community results, which indicated that soil pH was the most important edaphic factor in determining the bacterial community composition of these black soils. A variance partitioning analysis indicated that the geographic distance contributed 20% of the fungal community variation and soil environmental factors that were characterized explained approximately 35%. A pairwise analysis revealed that the diversity of the fungal community was relatively higher at lower latitudes, which is similar to the findings for the bacterial communities in the same region and suggests that a latitudinal gradient of microbial community diversity might occur in the black soil zone. By incorporating our previous findings on the bacterial communities, we can conclude that contemporary factors of soil characteristics are more important than historical factor of geographic distance in shaping the microbial community in the black soil zone of northeast China.     

Soil fertility is associated with fungal and bacterial richness,whereas pH is associated with community composition in polar soil microbial communities

Microbial activities in Arctic and Antarctic soils are of particular interest due to uncertainty surrounding the fate of the enormous polar soil organic matter (SOM) pools and the potential to lose unique and vulnerable micro-organisms from these ecosystems. We quantified richness, evenness and taxonomic composition of both fungi and bacteria in 223 Arctic and Antarctic soil samples across 8 locations to test the global applicability of hypotheses concerning edaphic drivers of soil microbial communities that have been primarily developed from studies of bacteria in temperate and tropical systems. We externally validated our model's conclusions with an independent dataset comprising 33 Arctic heath samples. We also explored if our system was responding to large scale climatic or biogeographical processes that we had not measured by evaluating model stability for one location, Mitchell Pennisula, that had been extensively sampled. Soil Fertility (defined as organic matter, nitrogen and chloride content) was the most important edaphic property associated with measures of α-diversity such as microbial richness and evenness (especially for fungi), whereas pH was primarily associated with measures of β-diversity such as phylogenetic structure and diversity (especially for bacteria). Surprisingly, phosphorus emerged as consistently the second most important driver of all facets of microbial community structure for both fungi and bacteria. Despite the clear importance of edaphic factors in controlling microbial communities, our analyses also indicated that fungal/bacterial interactions play a major, but causally unclear, role in structuring the soil microbial communities of which they are a part.     

Antagonistic and synergistic effects of fungal and bacterial growth in soil after adding different carbon and nitrogen sources

The effect of adding easily available and more complex carbon sources, with and without nitrogen, on fungal and bacterial growth and activity in soil were studied in the laboratory. Total microbial activity was estimated by measuring respiration, fungal growth with the acetate-in-ergosterol incorporation technique and bacterial growth with the thymidine and leucine incorporation techniques. The substrate additions consisted of glucose and cellulose, with and without nitrogen (as ammonium nitrate), and gelatine. The microbial development was followed over a 2-month period. The respiration rate increased within a few days after adding glucose, with and without nitrogen, and gelatine, initially by more than 10 times, but after 2 months no differences were seen compared with the control. Bacterial growth estimated with the thymidine and leucine incorporation techniques gave similar results. Adding glucose with nitrogen, or gelatine, increased bacterial growth within a few days up to 10 times, but even after 2 months of incubation bacterial growth rates were still about 5 times higher than in the control. Adding only glucose increased bacterial growth rates by about twice over the whole incubation period. Fungal growth rates especially increased after adding cellulose and nitrogen, although a minor increase was found after adding cellulose alone. Fungal growth rates started to increase after 10 days of incubation with cellulose. There were indications of synergistic effects in that bacterial growth increased after the fungi had started to grow after adding cellulose. Treatments resulting in high bacterial growth rates (adding easily available carbon sources) led to decreased fungal growth rates compared with the control, indicating antagonistic effects of bacteria.     

Predictive relationships between pH and sodicity in soils of tropical Queensland

Abstract

Interrelations between soil pH and exchangeable sodium percentage (ESP) were examined using soils from the Burdekin River area in tropical Queensland. Highly significant correlations were found but the goodness of fit differed between groups of soil profile classes. In general, Typic Natrustalfs of the flood plains had better relationships (r² = 0.85) between these soil properties than did the Chromusterts (r² = 0.50). The regression ESP = 1.935 × 10^‐5 _pH6.205 (_r2 = 0.61; n= 288) for all soils and depths underestimated ESP in Typic Natrustalfs groupings and overestimated this soil property in the Chromusterts.

By using the appropriate regression, pH levels associated with non‐sodic, sodic and strongly sodic horizons have been defined. Either laboratory or field determined estimates of pH may be used but the laboratory determined value is preferred. It is expected the predictive models will remain valid until soil ESP or pH levels are significantly modified as a consequence of agricultural development.     

Mineralization and carbon turnover in subarctic heath soil as affected by warming and additional litter

Arctic soil carbon (C) stocks are threatened by the rapidly advancing global warming. In addition to temperature, increasing amounts of leaf litter fall following from the expansion of deciduous shrubs and trees in northern ecosystems may alter biogeochemical cycling of C and nutrients. Our aim was to assess how factorial warming and litter addition in a long-term field experiment on a subarctic heath affect resource limitation of soil microbial communities (measured by thymidine and leucine incorporation techniques), net growing-season mineralization of nitrogen (N) and phosphorus (P), and carbon turnover (measured as changes in the pools during a growing-season-long field incubation of soil cores in situ). The mainly N limited bacterial communities had shifted slightly towards limitation by C and P in response to seven growing seasons of warming. This and the significantly increased bacterial growth rate under warming may partly explain the observed higher C loss from the warmed soil. This is furthermore consistent with the less dramatic increase in the contents of dissolved organic carbon (DOC) and dissolved organic N (DON) in the warmed soil than in the soil from ambient temperature during the field incubation. The added litter did not affect the carbon content, but it was a source of nutrients to the soil, and it also tended to increase bacterial growth rate and net mineralization of P. The inorganic N pool decreased during the field incubation of soil cores, especially in the separate warming and litter addition treatments, while gross mineralized N was immobilized in the biomass of microbes and plants transplanted into the incubates soil cores, but without any significant effect of the treatments. The effects of warming plus litter addition on bacterial growth rates and of warming on C and N transformations during field incubation suggest that microbial activity is an important control on the carbon balance of arctic soils under climate change.