Changes in soil microbial biomass carbon and enzyme activities under elevated CO2 affect fine root decomposition processes in a Mongolian oak ecosystem

The relationships between soil microbial properties and fine root decomposition processes under elevated CO₂ are poorly understood. To address this question, we determined soil microbial biomass carbon (SMB-C) and nitrogen (SMB-N), enzymes related to soil carbon (C) and nitrogen (N) cycling, the abundance of cultivable N-fixing bacteria and cellulolytic fungi, fine root organic matter, lignin and holocellulose decomposition, and N mineralization from 2006 to 2007 in a Mongolian oak (Quercus mongolica Fischer ex Ledebour) ecosystem in northeastern China. The experiment consisted of three treatments: elevated CO₂ chambers, ambient CO₂ chambers, and chamberless plots. Fine roots had significantly greater organic matter decomposition rates under elevated CO₂. This corresponded with significantly greater SMB-C. Changes in the activities of protease and phenol oxidase under elevated CO₂ could not explain the changes in fine root N release and lignin decomposition rates, respectively, while holocellulose decomposition rate had the same response to experimental treatments as did cellulase activity. Changes in cultivable N-fixing bacterial and cellulolytic fungal abundances in response to experimental treatments were identical to those of N mineralization and lignin decomposition rates, respectively, suggesting that the two indices were closely related to fine root N mineralization and lignin decomposition. Our results showed that the increased fine root organic matter, lignin and holocellulose decomposition, and N mineralization rates under elevated CO₂ could be explained by shifts in SMB-C and the abundance of cellulolytic fungi and N-fixing bacteria. Enzyme activities are not reliable for the assessment of fine root decomposition and more attention should be given to the measurement of specific bacterial and fungal communities.     

Elevated CO2 and O3t concentrations differentially affect selected groups of the fauna in temperate forest soils

Rising atmospheric CO₂ concentrations may change soil fauna abundance. How increase of tropospheric ozone (O_3t) concentration will modify these responses is still unknown. We have assessed independent and interactive effects of elevated [CO₂] and [O_3t] on selected groups of soil fauna. The experimental design is a factorial arrangement of elevated [CO₂] and [O_3t] treatments, applied using Free-Air CO₂ Enrichment technology to 30 m diameter rings, with all treatments replicated three times. Within each ring, three communities were established consisting of: (1) trembling aspen (Populus tremuloides) and paper birch (Betula papyrifera) (2) trembling aspen and sugar maple (Acer saccharum) and (3) trembling aspen. After 4 yr of stand development, soil fauna were extracted in each ring. Compared to the control, abundance of total soil fauna, Collembola and Acari decreased significantly under elevated [CO₂] (−69, −79 and −70%, respectively). Abundance of Acari decreased significantly under elevated [O_3t] (−47%). Soil fauna abundance was similar to the control under the combination of elevated [CO₂+O_3t]. The individual negative effects of elevated [CO₂] and elevated [O_3t] are negated upon exposure to both gases. We conclude that soil fauna communities will change under elevated [CO₂] and elevated [O_3t] in ways that cannot be predicted or explained from the exposure of ecosystems to each gas individually.     

Elevated CO2 differentially alters belowground plant and soil microbial community structure in reed canary grass-invaded experimental wetlands

Several recent studies have indicated that an enriched atmosphere of carbon dioxide (CO₂) could exacerbate the intensity of plant invasions within natural ecosystems, but little is known of how rising CO₂ impacts the belowground characteristics of these invaded systems. In this study, we examined the effects of elevated CO₂ and nitrogen (N) inputs on plant and soil microbial community characteristics of plant communities invaded by reed canary grass, Phalaris arundinacea L. We grew the invasive grass under two levels of invasion: the invader was either dominant (high invasion) at >90% plant cover or sub-dominant (low invasion) at <50% plant cover. Experimental wetland communities were grown for four months in greenhouses that received either 600 or 365 μl l⁻¹ (ambient) CO₂. Within each of three replicate rooms per CO₂ treatment, the plant communities were grown under high (30 mg l⁻¹) or low (5 mg l⁻¹) N. In contrast to what is often predicted under N limitation, we found that elevated CO₂ increased native graminoid biomass at low N, but not at high N. The aboveground biomass of reed canary grass did not respond to elevated CO₂, despite it being a fast-growing C3 species. Although elevated CO₂ had no impact on the plant biomass of heavily invaded communities, the relative abundance of several soil microbial indicators increased. In contrast, the moderately invaded plant communities displayed increased total root biomass under elevated CO_2, while little impact occurred on the relative abundance of soil microbial indicators. Principal components analysis indicated that overall soil microbial community structure was distinct by CO₂ level for the varying N and invasion treatments. This study demonstrates that even when elevated CO₂ does not have visible effects on aboveground plant biomass, it can have large impacts belowground.     

Changes in the fungal-to-bacterial respiratory ratio and microbial biomass in agriculturally managed soils under free-air CO2 enrichment (FACE) - A six-year survey of a field study

In soil ecology, microbial parameters have been identified as sensitive indicators of changes in the soil environment. The Braunschweig FACE project provided the opportunity to study the effects of elevated CO₂ (550 μmol mol⁻¹) as compared to ambient CO₂ (370 μmol mol⁻¹) on total microbial biomass (C_mic), C_mic-to-C_org ratio and the fungal-to-bacterial respiratory ratio together with total C_org, N_t, C:N ratio and pH over a six-year period. Field management followed a typical crop rotation system of this region with either a crop-related full nitrogen supply (N100) or 50% reduced N supply (N50). The soil microbial parameters responded to the elevated CO₂ treatment in varying intensities and time spans. The fungal-to-bacterial respiratory ratio was the most sensitive parameter in responding to an elevated CO₂ treatment with highly significant differences to ambient CO₂-treated control plots in the third year of CO₂ fumigation. After six years bacterial respiratory activity had increased in ascending order to 34% in FACE-treated plots (N50 and N100) as compared to control plots. Soil microbial biomass (C_mic) responded more slowly to the FACE treatment with highly significant increases of >12% after the fourth year of CO₂ fumigation. The C_mic-to-C_org ratio responded very late in the last two years of the CO₂ treatment with a significant increase of >7.0% only in the N100 variant. Total C_org and N_t were slightly but significantly increased under FACE around 10.0% with ascending tendency over time starting with the second year of CO₂ treatment. No significant FACE effects could be recorded for the C:N ratio or pH.These results suggest that under FACE treatment changes in the soil microbial community will occur. In our study the fungal-to-bacterial respiratory ratio was superior to total C_mic as microbial bioindicators in reflecting changes in the soil organic matter composition.     

Abundance and composition response of wheat field soil bacterial and fungal communities to elevated CO<Subscript>2</Subscript> and increased air temperature

The abundance and diversity of soil bacterial and fungal communities in a wheat field under elevated atmospheric CO₂ concentrations and increased air temperatures were investigated using qPCR and pyrosequencing. Elevated CO₂ concentrations significantly increased the abundances of bacteria and fungi, and an increase of air temperatures significantly reduced fungal abundance. We found that Proteobacteria, Bacteroidetes, Chloroflexi, and Ascomycota were the most abundant bacterial and fungal phyla in the wheat field soil. Elevated CO₂ concentrations and increased air temperatures had no significant effect on the bacterial alpha diversity, whereas fungal richness was reduced under warming treatments. Moreover, we note that certain bacterial and fungal groups responded differentially to elevated CO₂ concentrations and increased air temperatures, and fungal species were highly sensitive to climatic changes.     

Soil CO2 flux in six monospecific forest plantations in Southern Rwanda

Forest soils contain the largest carbon stock of all terrestrial biomes and are probably the most important source of carbon dioxide (CO₂) to atmosphere. Soil CO₂ fluxes from 54 to 72-year-old monospecific stands in Rwanda were quantified from March 2006 to December 2007. The influences of soil temperature, soil water content, soil carbon (C) and nitrogen (N) stocks, soil pH, and stand characteristics on soil CO₂ flux were investigated. The mean annual soil CO₂ flux was highest under Eucalyptus saligna (3.92 μmol m⁻² s⁻¹) and lowest under Entandrophragma excelsum (3.13 μmol m⁻² s⁻¹). The seasonal variation in soil CO₂ flux from all stands followed the same trend and was highest in rainy seasons and lowest in dry seasons. Soil CO₂ flux was mainly correlated to soil water content (R² = 0.36-0.77), stand age (R² = 0.45), soil C stock (R² = 0.33), basal area (R² = 0.21), and soil temperature (R² = 0.06-0.17). The results contribute to the understanding of factors that influence soil CO₂ flux in monocultural plantations grown under the same microclimatic and soil conditions. The results can be used to construct models that predict soil CO₂ emissions in the tropics.     

Structural and functional diversity of soil microbes is affected by elevated [CO<Subscript>2</Subscript>] and N addition in a poplar plantation

Background, Aims, and Scope  The genetic structure and the functionality of soil microbes are both important when studying the role of soil in the C cycle in elevated CO₂ scenarios. The aim of this work was to investigate the genetic composition of the fungal community by means of PCR-DGGE and the functional diversity of soil micro-organisms in general with MicroResp-based community level physiological profiling (CLPP) in a poplar plantation (POPFACE) grown under elevated [CO₂] with and without nitrogen fertilization. Materials and Methods  The POPFACE experimental plantation and FACE facility are located in central Italy, Tuscania (VT). Clones of Populus alba, Populus nigra and Populus x euramericana were grown, from 1999 to 2004, in six 314 m² plots treated either with atmospheric (control) or enriched (550 μmol mol⁻¹) CO₂ with FACE (Free Air CO₂ Enrichment) technology in each growing season. Each plot is divided into six triangular sectors, with two sectors per poplar genotype: three species × two nitrogen levels. After removal of the litter layer one soil core per genotype (10 cm wide, 20 cm depth) was taken inside each of the three sectors in each plot, for a total of 36 soil cores (3 replicates × 2 [CO₂] × 2 fertilization × 3 species) in October 2004 and in July 2005. DNA was extracted with a bead beating procedure. 18S rDNA gene fragments were amplified with PCR using fungal primers (FR1 GC and FF390). Analysis of CLPP was performed using the MicroResp method. Carbon substrates were selected depending on their ecological relevance to soil and their solubility in water. In particular rhizospheric C sources (carboxylic acids and carbohydrates) were chosen considering the importance of root inputs for microbial metabolism. Results  The fertilization treatment differentiated the fungal community composition regardless of elevated [CO₂] or the poplar species; moreover the number of fungal species was lower in fertilized soil. The effect of elevated [CO₂] on the fungal community composition was evident only as interaction with the fertilization treatment as, in N-sufficient soils, the elevated [CO₂] selected a different microbial community. For CLPP, the differ ent poplar species were the main factors of variation. The FACE treatment, on average, resulted in lower C utilization rates in un-fertilized soils and higher in fertilized soils. Discussion  Fungal biomass and fungal composition depend on different factors: from previous studies we know that the greater quantity and the higher C/N ratio of organic inputs under elevated [CO₂] influenced positively the fungal biomass both in fertilized and in un-fertilized soil, whereas nitrogen availability resulted to be the main determinant of fungal community composition in this work. Whole active microbial community was directly influenced by the soil nutrient availability and the poplar species. Under elevated CO₂ the competition for N with plants strongly affected the microbial communities, which were not able to benefit from added rhizospheric substrates. Under Nsufficient conditions, the increase of microbial activity due to [CO₂] enrichment was related to a more active microbial community, favoured by the current availability of C and N. Conclusions  Different factors influenced the microbial community at different levels: poplar species and root exudates affected the functional properties of the microbial community, while the fungal specific composition (as seen with DGGE) remained unaffected. On the other hand, factors such as N and C availability had a strong impact on the community functionality and composition. Fungal community structure reflected the availability of N in soils and the effect of elevated [CO₂] on community structure and function was evident only in N-sufficient soils. The simultaneous availability of C and N was therefore the main driving force for microbial structure and function in this plantation. Recommendations and Perspectives  Using the soil instead of soil extracts for CLPP determination provides a direct measurement of substrate catabolism by microbial communities and reflects activity rather than growth because more immediate responses to substrates are measured. Further applications of this approach could include selective inhibition of different microbial functional groups to investigate specific CLPPs. To combine the structural analysis and the catabolic responses of specific microbial communities (i.e. fungi or bacteria) could provide new outlooks on the role of microbes on SOM decomposition. ESS-Submission Editor: Dr. Kirk Semple (k.semple@lancaster.ac.uk)     

Effects of past, present and future atmospheric CO2 concentrations on soil organic matter dynamics in a chaparral ecosystem

Owing to the continuously increasing concentration of atmospheric CO₂, it has become a priority to understand if soil organic matter (SOM) will behave as a sink or a source of CO₂ under future environmental changes. Although many studies have addressed this question, a clear understanding is still missing, particularly with respect to long-term responses. In this study, we quantified soil C stores and dynamics in relationship to soil aggregation and pool composition in a Californian chaparral ecosystem exposed for 6 years to a gradient of atmospheric CO₂ concentrations, ranging from pre-industrial levels 250 to 750 μl l⁻¹ CO₂. Fossil fuel-derived CO₂ depleted in ¹³C was used for the fumigation, thus providing a tracer of C input from the vegetation to the soil.Long-term CO₂ exposure invariably affected soil aggregation, with a significant decrease in the macroaggregate fraction at highest CO₂ levels relative to the other two size fractions (i.e. microaggregates and silt and clay). This soil structural change most likely reduced the stability and protection of SOM, and C content generally decreased in most fractions over the CO₂ treatments, and induced faster turnover of recently fixed C at high CO₂ levels. The strongest response was found in the C content of the microaggregates, which decreased significantly (P<0.05) with rising levels of CO₂. We conclude that increasing atmospheric CO₂ concentrations will decrease soil C in chaparral ecosystems, and that the microaggregate fraction is the most responsive to increasing concentrations of atmospheric CO₂.     

Soil CO2 efflux vs. soil respiration: Implications for flux models

In the long term, all CO₂ produced in the soil must be emitted by the surface and soil CO₂ efflux (FCO₂) must correspond to soil respiration (R_soil). In the short term, however, the efflux can deviate from the instantaneous soil respiration, if the amount of CO₂ stored in the soil pore-space (SCO₂) is changing. We measured FCO₂ continuously for one year using an automated chamber system. Simultaneously, vertical soil profiles of CO₂ concentration, moisture, and temperature were measured in order to assess the changes in the amount of CO₂ stored in the soil. R_soil was calculated as the sum of the rate of change of the CO₂ storage over time and FCO₂. The experiment was split into a warm and a cold season. The dependency of soil respiration and soil efflux on soil temperature and on soil moisture was analyzed separately. Only the moisture-driven model of the warm season was significantly different for FCO₂ and R_soil. At our site, a moisture-driven soil-respiration model derived from CO₂ efflux data would underestimate the importance of soil moisture. This effect can be attributed to a temporary storage of CO₂ in the soil pore-space after rainfalls where up to 40% of the respired CO₂ were stored.     

Effects of root and litter exclusion on soil CO2 efflux and microbial biomass in wet tropical forests

We examined the effects of root and litter exclusion on the rate of soil CO₂ efflux and microbial biomass at a soil depth of 25 cm in a secondary forest (dominated by Tabebuia heterophylla) and a pine (Pinus caribaea) plantation in the Luquillo Experimental Forest in Puerto Rico. The experimental plots were initially established in 1990, when root, forest floor mass and new litterfall were excluded for 7 y since then. Soil respiration was significantly reduced in the litter and root exclusion plots in both the secondary forest and the pine plantation compared with the control. Root exclusion had a greater effect on soil CO₂ efflux than the litter exclusion in the plantation, whereas a reversed pattern was observed in the secondary forest. The reduction of microbial biomass in the root exclusion plot was greater in the secondary forest (59%) than in the plantation (31%), while there was no difference of the reduction in the litter exclusion plots between these forests. Our results suggest that above-ground input and roots (root litter and exudates) differentially affect soil CO₂ efflux under different vegetation types.     

Decomposition of C-labeled roots in a pasture soil exposed to 10 years of elevated CO2

The net flux of soil C is determined by the balance between soil C input and microbial decomposition, both of which might be altered under prolonged elevated atmospheric CO₂. In this study, we determined the effect of elevated CO₂ on decomposition of grass root material (Lolium perenne L.). ¹⁴C-labeled root material, produced under ambient (35 Pa pCO₂) or elevated CO₂ (70 Pa pCO₂) was incubated in soil for 64 days. The soils were taken from a pasture ecosystem which had been exposed to ambient (35 Pa pCO₂) or elevated CO₂ (60 Pa pCO₂) under FACE-conditions for 10 years and two fertilizer N rates: 140 and 560 kg N ha⁻¹ year⁻¹. In soil exposed to elevated CO₂, decomposition rates of root material grown at either ambient or elevated CO₂ were always lower than in the control soil exposed to ambient CO₂, demonstrating a change in microbial activity. In the soil that received the high rate of N fertilizer, decomposition of root material grown at elevated CO₂ decreased by approximately 17% after incubation for 64 days compared to root material grown at ambient CO₂. The amount of ¹⁴CO₂ respired per amount of ¹⁴C incorporated in the microbial biomass (q¹⁴CO₂) was significantly lower when roots were grown under high CO₂ compared to roots grown under low CO₂. We hypothesize that this decrease is the result of a shift in the microbial community, causing an increase in metabolic efficiency. Soils exposed to elevated CO₂ tended to respire more native SOC, both with and without the addition of the root material, probably resulting from a higher C supply to the soil during the 10 years of treatment with elevated CO₂. The results show the importance of using soils adapted to elevated CO₂ in studies of decomposition of roots grown under elevated CO₂. Our results further suggest that negative priming effects may obscure CO₂ data in incubation experiments with unlabeled substrates. From the results obtained, we conclude that a slower turnover of root material grown in an ‘elevated-CO₂ world’ may result in a limited net increase in C storage in ryegrass swards.     

大气CO2浓度升高对温带森林土壤微生物活性的影响

Microorganisms play a key role in the response of soil ecosystems to the rising atmospheric carbon dioxide (CO2) as they mineralize organic matter and drive nutrient cycling. To assess the effects of elevated CO2 on soil microbial C and N immobilization and on soil enzyme activities, in years 8 (2006) and 9 (2007) of an open-top chamber experiment that begun in spring of 1999, soil was sampled in summer, and microbial biomass and enzyme activity related to the carbon (C), nitrogen (N) and phosphorus (P) cycling were measured. Although no effects on microbial biomass C were detected, changes in microbial biomass N and metabolic activity involving C, N and P were observed under elevated CO2. Invertase and dehydrogenase activities were significantly enhanced by different degrees of elevated CO2. Nitrifying enzyme activity was significantly (P < 0.01) increased in the August 2006 samples that received the elevated CO2 treatment, as compared to the samples that received the ambient treatment. Denitrifying enzyme activity was significantly (P < 0.04) decreased by elevated CO2 treatments in the August 2006 and June 2007 (P < 0.09) samples. β-N-acetylglucosaminidase activity was increased under elevated CO2 by 7% and 25% in June and August 2006, respectively, compared to those under ambient CO2. The results of June 2006 samples showed that acid phosphatase activity was significantly enhanced under elevated CO2. Overall, these results suggested that elevated CO2 might cause changes in the belowground C, N and P cycling in temperate forest soils.     

Gross fluxes of nitrogen in grassland soil exposed to elevated atmospheric pCO2 for seven years

Plant response to increasing atmospheric CO₂ partial pressure (pCO₂) depends on several factors, one of which is mineral nitrogen availability facilitated by the mineralisation of organic N. Gross rates of N mineralisation were examined in grassland soils exposed to ambient (36 Pa) and elevated (60 Pa) atmospheric pCO₂ for 7 years in the Swiss Free Air Carbon dioxide Enrichment experiment. It was hypothesized that increased below-ground translocation of photoassimilates at elevated pCO₂ would lead to an increase in immobilisation of N due to an excess supply of energy to the roots and rhizosphere. Intact soil cores were sampled from Lolium perenne and Trifolium repens swards in May and September, 2000. The rates of gross N mineralisation (m) and NH₄⁺ consumption (c) were determined using ¹⁵N isotopic dilution during a 51-h period of incubation. The rates of N immobilisation were estimated either as the difference between m and the net N mineralisation rate or as the amount of ¹⁵N released from the microbial biomass after chloroform fumigation. Soil samples from both swards showed that the rates of gross N mineralisation and NH₄⁺ consumption did not change significantly under elevated pCO₂. The lack of a significant effect of elevated pCO₂ on organic N turnover was consistent with the similar size of the microbial biomass and similar immobilisation of applied ¹⁵N in the microbial N pool under ambient and elevated pCO₂. Rates of m and c, and microbial ¹⁵N did not differ significantly between the two sward types although a weak (p<0.1) pCO₂ by sward interaction occurred. A significantly larger amount of NO₃⁻ was recovered at the end of the incubation in soil taken from T. repens swards compared to that from L. perenne swards. Eleven percent of the added ¹⁵N were recovered in the roots in the cores sampled under L. perenne, while only 5% were recovered in roots of T. repens. These results demonstrate that roots remained a considerable sink despite the shoots being cut at ground level prior to incubation and suggest that the calculation of N immobilisation from gross and net rates of mineralisation in soils with a high root biomass does not reflect the actual immobilisation of N in the microbial biomass. The results of this study did not support the initial hypothesis and indicate that below-ground turnover of N, as well as N availability, measured in short-term experiments are not strongly affected by long-term exposure to elevated pCO₂. It is suggested that differences in plant N demand, rather than major changes in soil N mineralisation/immobilisation, are the long-term driving factors for N dynamics in these grassland systems.     

Rice root-derived carbon input and its effect on decomposition of old soil carbon pool under elevated CO2

Rice (Oryza sativa) was grown in sunlit, semi-closed growth chambers (4×3×2 m, L×W×H) at 650 μl l⁻¹ CO₂ (elevated CO₂) to determine: (1) rice root-derived carbon (C) input into the soil under elevated CO₂ in one growing season, and (2) the effect of the newly input C on decomposition of the more recalcitrant native soil organic C. The initial δ¹³C value of the experimental soil was −25.8‰, which was 6‰ less depleted in ¹³C than the plants grown under elevated CO₂. Significant changes in δ¹³C of the soil organic C were detected after one growing season. The amount of new soil C input was estimated to be 0.9 t ha⁻¹ (or 2.1%) at 30 kg N ha⁻¹ and 1.8 t ha⁻¹ (4.1%) at 90 kg N ha⁻¹. Changes in soil δ¹³C suggested that the surface 5 cm of soil received more C input from plants than soils below. Laboratory incubation (25 °C) of soils from different horizons indicated that increased availability of the labile plant-derived C in the soil reduced decomposition of the native soil organic C. Provided the retardant effect of the new C on old soil organic C holds in the field in the longer-term, paddy soils will likely sequester more C from the atmosphere if more plant C enters the soil under elevated atmospheric CO₂.     

Elevated CO2 increases the effect of an arbuscular mycorrhizal fungus and a plant-growth-promoting rhizobacterium on structural stability of a semiarid agricultural soil under drought conditions

In arid and semiarid Mediterranean regions, an increase in the severity of drought events could be caused by rising atmospheric CO₂ concentrations. We studied the effects of the interaction of CO₂, water supply and inoculation with a plant-growth-promoting rhizobacterium (PGPR), Pseudomonas mendocina Palleroni, or inoculation with an arbuscular mycorrhizal (AM) fungus, Glomus intraradices (Schenk & Smith), on aggregate stabilisation of the rhizosphere soil of Lactuca sativa L. cv. Tafalla. The influence of such structural improvements on the growth of lettuce was evaluated. We hypothesised that elevated atmospheric CO₂ concentration would increase the beneficial effects of inoculation with a PGPR or an AM fungus on the aggregate stability of the rhizosphere soil of lettuce plants. Leaf hydration, shoot dry biomass and mycorrhizal colonisation were decreased significantly under water-stress conditions, but this decrease was more pronounced under ambient vs elevated CO₂. The root biomass decreased under elevated CO₂ but only in non-stressed plants. Under elevated CO₂, the microbial biomass C of the rhizosphere of the G. intraradices-colonised plants increased with water stress. Bacterial and mycorrhizal inoculation and CO₂ had no significant effect on the easily-extractable glomalin concentration. Plants grown under elevated CO₂ had a significantly higher percentage of stable aggregates under drought stress than under well-watered conditions, particularly the plants inoculated with either of the assayed microbial inocula (about 20% higher than the control soil). We conclude that the contribution of mycorrhizal fungi and PGPR to soil aggregate stability under elevated atmospheric CO₂ is largely enhanced by soil drying.     

Climate change goes underground: effects of elevated atmospheric CO<Subscript>2</Subscript> on microbial community structure and activities in the rhizosphere

General concern about climate change has led to growing interest in the responses of terrestrial ecosystems to elevated concentrations of CO₂ in the atmosphere. Experimentation during the last two to three decades using a large variety of approaches has provided sufficient information to conclude that enrichment of atmospheric CO₂ may have severe impact on terrestrial ecosystems. This impact is mainly due to the changes in the organic C dynamics as a result of the effects of elevated CO₂ on the primary source of organic C in soil, i.e., plant photosynthesis. As the majority of life in soil is heterotrophic and dependent on the input of plant-derived organic C, the activity and functioning of soil organisms will greatly be influenced by changes in the atmospheric CO₂ concentration. In this review, we examine the current state of the art with respect to effects of elevated atmospheric CO₂ on soil microbial communities, with a focus on microbial community structure. On the basis of the existing information, we conclude that the main effects of elevated atmospheric CO₂ on soil microbiota occur via plant metabolism and root secretion, especially in C3 plants, thereby directly affecting the mycorrhizal, bacterial, and fungal communities in the close vicinity of the root. There is little or no direct effect on the microbial community of the bulk soil. In particular, we have explored the impact of these changes on rhizosphere interactions and ecosystem processes, including food web interactions.     

Soil carbon dioxide fluxes in established switchgrass land managed for biomass production

Switchgrass (Panicum virgatum L.) grown for biomass feedstock production has the potential to increase soil C sequestration, and soil CO₂ flux in grassland is an important component in the global C budget. The objectives of this study were to: (1) determine the effects of N fertilization and harvest frequency on soil CO₂ flux, soil microbial biomass carbon (SMBC), and potentially mineralizable carbon (PMC); and (2) evaluate the relationship of soil CO₂ flux with soil temperature, soil moisture, SMBC, and PMC. Two N rates (0 and 224 kg ha⁻¹) were applied as NH₄NO₃ and cattle (Bos Taurus L.) manure. Switchgrass was harvested every year at anthesis or alternate years at anthesis. The data were collected during growing season (May-October) 2001-2004 on switchgrass-dominated Conservation Reserve Program (CRP) land in east-central South Dakota, USA. Manure application increased soil CO₂ flux, SMBC, and PMC during the early portion of the growing season compared with the control, but NH₄NO₃ application did not affect soil CO₂ flux, SMBC, and PMC. However, seasonal variability of soil CO₂ flux was not related to SMBC and PMC. Estimated average soil CO₂ fluxes during the growing periods were 472, 488, and 706 g CO₂-C m⁻² for control, NH₄NO₃-N, and manure-N plots, respectively. Switchgrass land with manure application emitted more CO₂, and approximately 45% of the C added with manure was respired to the atmosphere. Switchgrass harvested at anthesis decreased soil CO₂ flux during the latter part of the growing season, and flux was lower under every year harvest treatment than under alternate years harvest. Soil temperature was the most significant single variable to explain the variability in soil CO₂ flux. Soil water content was not a limiting factor in controlling seasonal CO₂ flux.     

Atmospheric CO2 enrichment and nutrient additions to planted soil increase mineralisation of soil organic matter, but do not alter microbial utilisation of plant- and soil C-sources

Plants link atmospheric and soil carbon pools through CO₂ fixation, carbon translocation, respiration and rhizodeposition. Within soil, microbial communities both mediate carbon-sequestration and return to the atmosphere through respiration. The balance of microbial use of plant-derived and soil organic matter (SOM) carbon sources and the influence of plant-derived inputs on microbial activity are key determinants of soil carbon-balance, but are difficult to quantify. In this study we applied continuous ¹³C-labelling to soil-grown Lolium perenne, imposing atmospheric CO₂ concentrations and nutrient additions as experimental treatments. The relative use of plant- and SOM-carbon by microbial communities was quantified by compound-specific ¹³C-analysis of phospholipid fatty acids (PLFAs). An isotopic mass-balance approach was applied to partition the substrate sources to soil respiration (i.e. plant- and SOM-derived), allowing direct quantification of SOM-mineralisation. Increased CO₂ concentration and nutrient amendment each increased plant growth and rhizodeposition, but did not greatly alter microbial substrate use in soil. However, the increased root growth and rhizosphere volume with elevated CO₂ and nutrient amendment resulted in increased rates of SOM-mineralisation per experimental unit. As rhizosphere microbial communities utilise both plant- and SOM C-sources, the results demonstrate that plant-induced priming of SOM-mineralisation can be driven by factors increasing plant growth. That the balance of microbial C-use was not affected on a specific basis may suggest that the treatments did not affect soil C-balance in this study.     

Changes in forest soil organic matter pools after a decade of elevated CO2 and O3

The impact of rising atmospheric carbon dioxide (CO₂) may be mitigated, in part, by enhanced rates of net primary production and greater C storage in plant biomass and soil organic matter (SOM). However, C sequestration in forest soils may be offset by other environmental changes such as increasing tropospheric ozone (O₃) or vary based on species-specific growth responses to elevated CO₂. To understand how projected increases in atmospheric CO₂ and O₃ alter SOM formation, we used physical fractionation to characterize soil C and N at the Rhinelander Free Air CO₂-O₃ Enrichment (FACE) experiment. Tracer amounts of ¹⁵NH₄⁺ were applied to the forest floor of Populus tremuloides, P. tremuloides-Betula papyrifera and P. tremuloides-Acer saccharum communities exposed to factorial CO₂ and O₃ treatments. The ¹⁵N tracer and strongly depleted ¹³C-CO₂ were traced into SOM fractions over four years. Over time, C and N increased in coarse particulate organic matter (cPOM) and decreased in mineral-associated organic matter (MAOM) under elevated CO₂ relative to ambient CO₂. As main effects, neither CO₂ nor O₃ significantly altered ¹⁵N recovery in SOM. Elevated CO₂ significantly increased new C in all SOM fractions, and significantly decreased old C in fine POM (fPOM) and MAOM over the duration of our study. Overall, our observations indicate that elevated CO₂ has altered SOM cycling at this site to favor C and N accumulation in less stable pools, with more rapid turnover. Elevated O₃ had the opposite effect, significantly reducing cPOM N by 15% and significantly increasing the C:N ratio by 7%. Our results demonstrate that CO₂ can enhance SOM turnover, potentially limiting long-term C sequestration in terrestrial ecosystems; plant community composition is an important determinant of the magnitude of this response.     

Nitrogen fixation by biological soil crusts and heterotrophic bacteria in an intact Mojave Desert ecosystem with elevated CO2 and added soil carbon

Fixation of N by biological soil crusts and free-living heterotrophic soil microbes provides a significant proportion of ecosystem N in arid lands. To gain a better understanding of how elevated CO₂ may affect N₂-fixation in aridland ecosystems, we measured C₂H₂ reduction as a proxy for nitrogenase activity in biological soil crusts for 2 yr, and in soils either with or without dextrose-C additions for 1 yr, in an intact Mojave Desert ecosystem exposed to elevated CO₂. We also measured crust and soil δ¹⁵N and total N to assess changes in N sources, and δ¹³C of crusts to determine a functional shift in crust species, with elevated CO₂. The mean rate of C₂H₂ reduction by biological soil crusts was 76.9±5.6 μmol C₂H₄ m⁻² h⁻¹. There was no significant CO₂ effect, but crusts from plant interspaces showed high variability in nitrogenase activity with elevated CO₂. Additions of dextrose-C had a positive effect on rates of C₂H₂ reduction in soil. There was no elevated CO₂ effect on soil nitrogenase activity. Plant cover affected soil response to C addition, with the largest response in plant interspaces. The mean rate of C₂H₂ reduction in soils either with or without C additions were 8.5±3.6 μmol C₂H₄ m⁻² h⁻¹ and 4.8±2.1 μmol m⁻² h⁻¹, respectively. Crust and soil δ¹⁵N and δ¹³C values were not affected by CO₂ treatment, but did show an effect of cover type. Crust and soil samples in plant interspaces had the lowest values for both measurements. Analysis of soil and crust [N] and δ¹⁵N data with the Rayleigh distillation model suggests that any plant community changes with elevated CO₂ and concomitant changes in litter composition likely will overwhelm any physiological changes in N₂-fixation.