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)     

Assessment of 10 years of CO2 fumigation on soil microbial communities and function in a sweetgum plantation

Increased vegetative growth and soil carbon (C) storage under elevated carbon dioxide concentration ([CO₂]) has been demonstrated in a number of experiments. However, the ability of ecosystems, either above- or belowground, to maintain increased C storage relies on the response of soil processes, such as those that control nitrogen (N) mineralization, to climatic change. These soil processes are mediated by microbial communities whose activity and structure may also respond to increasing atmospheric [CO₂]. We took advantage of a long-term (ca 10 y) CO₂ enrichment experiment in a sweetgum plantation located in the southeastern United States to test the hypothesis that observed increases in root production in elevated relative to ambient CO₂ plots would alter microbial community structure, increase microbial activity, and increase soil nutrient cycling. We found that elevated [CO₂] had no detectable effect on microbial community structure using 16S rRNA gene clone libraries, on microbial activity measured with extracellular enzyme activity, or on potential soil N mineralization and nitrification rates. These results support findings at other forested Free Air [CO₂] Enrichment (FACE) sites.     

Warming and elevated CO2 combine to increase microbial mineralisation of soil organic matter

The net annual exchange of carbon between the atmosphere and terrestrial ecosystems is of prime importance in determining the concentration of CO₂ ([CO₂]) in the atmosphere and consequently future climate. Carbon loss occurs primarily through soil respiration; it is known that respiration is sensitive to the global changes in [CO₂] and temperature, suggesting that the net carbon balance may change in the future. However, field manipulations of temperature and [CO₂] alter many important environmental factors so it is unclear how much of the observed alterations in soil respiration is due to changes of microbial function itself instead of changes to the physical and chemical environment. Here we focus on resolving the importance of changes in the microbial community in response to warming and elevated [CO₂] on carbon mineralisation, something not possible in field measurements. We took plant material and soil inocula from a long running experiment where native grassland had been exposed to both warming and elevated CO₂ and constructed a reciprocal transplant experiment. We found that the rate of decomposition (heterotrophic respiration) was strongly determined by the origin of the microbial community. The combined warming + elevated CO₂ treatment produced a soil community that gave respiration rates 30% higher when provided with shoot litter and 70% for root litter than elevated CO₂ treatment alone, with the treatment source of the litter being unimportant. Warming, especially in the presence of elevated CO₂, increased the size of the apparent labile carbon pool when either C₃ or C₄ litter was added. Thus, the metabolic activity of the soil community was affected by the combination of warming and elevated CO₂ such that it had an increased ability to mineralise added organic matter, regardless of its source. Therefore, soil C efflux may be substantially increased in a warmer, high CO₂ world. Current ecosystem models mostly drive heterotrophic respiration from plant litter quality, soil moisture and temperature but our findings suggest equal attention will need to be paid to capturing microbial processes if we are to accurately project the future C balance of terrestrial ecosystems and quantify the feedback effect on atmospheric concentrations of CO₂.     

Short and long-term effects of elevated CO2 on Lolium perenne rhizodeposition and its consequences on soil organic matter turnover and plant N yield

It is still unclear whether elevated CO₂ increases plant root exudation and consequently affects the soil microbial biomass. The effects of elevated CO₂ on the fate of the C and nitrogen (N) contained in old soil organic matter pools is also unclear. In this study the short and long-term effects of elevated CO₂ on C and N pools and fluxes were assessed by growing isolated plants of ryegrass (Lolium perenne) in glasshouses at elevated and ambient atmospheric CO₂ and using soil from the New Zealand FACE site that had >4 years exposure to CO₂ enrichment. Using ¹⁴CO₂ pulse labelling, the effects of elevated CO₂ on C allocation within the plant-soil system were studied. Under elevated CO₂ more root derived C was found in the soil and in the microbial biomass 48 h after labelling. The increased availability of substrate significantly stimulated soil microbial growth and acted as priming effect, enhancing native soil organic matter decomposition regardless of the mineral N supply. Despite indications of faster N cycling in soil under elevated CO₂, N availability to plants stayed unchanged. Soil previously exposed to elevated CO₂ exhibited a higher N cycling rate but again there was no effect on plant N uptake. With respect to the difficulties of extrapolating glasshouse experiment results to the field, we concluded that the accumulation of coarse organic matter observed in the field under elevated CO₂ was probably not created by an imbalance between C and N but was likely to be due to more complex phenomena involving soil mesofauna and/or other nutrients limitations.     

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.     

Microbial assimilation of atmospheric CO2 into soil organic matter revealed by the incubation of paddy soils under 14C-CO2 atmosphere

Similar to higher plants, microbial autotrophs possess photosynthetic systems that enable them to fix CO₂. To measure the activity of microbial autotrophs in assimilating atmospheric CO₂, five paddy soils were incubated with ¹⁴C-labeled CO₂ for 45 days to determine the amount of ¹⁴C-labeled organic C being synthesized. The results showed that a significant amount of ¹⁴C-labeled CO₂ incorporated into microbial biomass was soil specific, accounting for 0.37%–1.18% of soil organic carbon (¹⁴C-labeled organic C range: 81.6–156.9 mg C kg^?1 of the soil after 45 days). Consequently, high amounts of C-labeled organic C were synthesized (the synthesis rates ranged from 86 to 166 mg C m^?2 d^?1). The amount of atmospheric ¹⁴CO₂ incorporated into microbial biomass (¹⁴C-labeled microbial biomass) was significantly correlated with organic C components (¹⁴C-labeled organic C) in the soil (r = 0.80, p < 0.0001). Our results indicate that the microbial assimilation of atmospheric CO₂ is an important process for the sequestration and cycling of terrestrial C. Our results showed that microbial assimilation of atmospheric CO₂ has been underestimated by researchers globally, and that it should be accounted for in global terrestrial carbon cycle models.     

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 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.     

A call to investigate drivers of soil organic matter retention vs. mineralization in a high CO2 world

Understanding how elevated atmospheric CO₂ alters the formation and decomposition of soil organic carbon (SOC) is important but challenging. If elevated CO₂ induces even small changes in rates of formation or decay of SOC, there could be substantial feedbacks on the atmosphere's concentration of CO₂. However, the long turnover times of many SOC pools - decades to centuries - make the detection of changes in the soil's pool size difficult. Long-term CO₂ enrichment experiments have offered unprecedented opportunities to explore these issues in intact ecosystems for more than a decade. Increased NPP with elevated CO₂ has prompted the hypothesis that SOC may increase at the same time that increased vegetation nitrogen (N) uptake and accumulation indicates probable declines in SON. Varying investigators thus have hypothesized that SOC will increase and SON will decline to explain increased NPP with elevated CO₂; researchers also invoke biogeochemical theory and stoichiometric constraints to argue for strong limitations on the co-occurrence of these phenomena. We call for researchers to investigate two broad research questions to elucidate the drivers of these processes. First, we ask how elevated CO₂ influences compound structure and stoichiometry of that proportion of NPP retained by soil profiles for relatively long time periods. We also call for investigations of the mechanisms underlying the decomposition of mineralizable organic matter with elevated CO₂. Specifically, we need to understand how elevated CO₂ influences microbial priming (driven by enhanced microbial energy needs associated with increases in biomass or activity) and microbial mining of N (driven by enhanced microbial N demand associated with greater vegetative N uptake), two processes that necessarily will be constrained by the stoichiometry of both substrates and microbial demands. Applying technologies such as nuclear magnetic resonance and the detection of biomarkers that reveal organic matter structure and origins, and studying microbial stoichiometric constraints, will dramatically improve our ability to predict future patterns of ecosystem C and N cycling.     

Repeated CO2 pulse-labelling reveals an additional net gain of soil carbon during growth of spring wheat under free air carbon dioxide enrichment (FACE)

Rising levels of atmospheric CO₂ have often been found to increase above and belowground biomass production of C3 plants. The additional translocation of organic matter into soils by increased root mass and exudates are supposed to possibly increase C pools in terrestrial ecosystems. Corresponding investigations were mostly conducted under more or less artificial indoor conditions with disturbed soils. To overcome these limitations, we conducted a ¹⁴CO₂ pulse-labelling experiment within the German FACE project to elucidate the role of an arable crop system in carbon sequestration under elevated CO₂. We cultivated spring wheat cv. “Minaret” with usual fertilisation and ample water supply in stainless steel cylinders forced into the soil of a control and a FACE plot. Between stem elongation and beginning of ripening the plants were repeatedly pulse-labelled with ¹⁴CO₂ in the field. Soil born total CO₂ and ¹⁴CO₂ was monitored daily till harvest. Thereafter, the distribution of ¹⁴C was analysed in all plant parts, soil, soil mineral fractions and soil microbial biomass. Due to the small number of grown wheat plants (40) in each ring and the inherent low statistical power, no significant above and belowground growth effect of elevated CO₂ was detected at harvest. But in comparison to ambient conditions, 28% more ¹⁴CO₂ and 12% more total CO₂ was evolved from soil under elevated CO₂ (550 μmol CO₂ mol⁻¹). In the root-free soil 27% more residual ¹⁴C was found in the FACE soil than in the soil from the ambient ring. In soil samples from both treatments about 80% of residual ¹⁴C was found in the clay fraction and 7% in the silt fraction. Very low ¹⁴C contents in the CFE extracts of microbial biomass in the soil from both CO₂ treatments did not allow assessing their influence on this parameter. Since the calculated specific radioactivity of soil born ¹⁴CO₂ gave no indication of an accelerated priming effect in the FACE soil, we conclude that wheat plants grown under elevated CO₂ can contribute to an additional net carbon gain in soils.     

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.     

Carbon turnover in the rhizosphere under continuous plant labeling with <Superscript>13</Superscript>CO<Subscript>2</Subscript>: Partitioning of root,microbial, and rhizomicrobial respiration

The input of labeled C into the pool of soil organic matter, the CO₂ fluxes from the soil, and the contribution of root and microbial respiration to the CO₂ emission were studied in a greenhouse experiment with continuous labeling of oat plants with ¹³CO₂ using the method of the natural ¹³C abundance in the air. The carbon of the microbial biomass composed 56 and 39% of the total amounts of ¹³C photoassimilates in the rhizosphere and in the bulk soil, respectively. The contribution of root respiration to the CO₂ emission from the soil reached 61–92%, including 4–23% of the rhizomicrobial respiration. The contribution of the microbial respiration to the total CO₂ emission from the soil varied from 8 to 39%. The soil organic matter served as the major carbon-containing substrate for microorganisms in the bulk soil and in the rhizosphere: 81–91% of the total amount of carbon involved in the microbial metabolism was derived from the soil organic matter.     

Effect of long-term free-air CO2 enrichment on the diversity and activity of soil methanogens in a periodically waterlogged grassland

As atmospheric CO₂ levels continue to rise researchers now identify concomitant changes in plant biomass and diversity, which are postulated to alter the quality and quantity of the organic carbon entering the soil. In anoxic soils, CH₄ is the end product of the degradation of organic carbon and the system's terminal electron sink. Some soils (such as wetlands) are usually waterlogged and therefore constitute permanent CH₄ sources, while others (hydromorphic soils) are only occasionally saturated with water and alternate between acting as net CH₄ sources or sinks. Since methanogenesis is ultimately dependent on soil organic carbon, we hypothesized that a long term alteration of the latter will cause changes in type and magnitude of the former. To test this, we studied the effect of 10 years of atmospheric CO₂ enrichment on the methanogenic potential and community in a hydromorphic temperate grassland soil at the experimental Free Air Carbon dioxide Enrichment (FACE) site in Giessen, Germany. While all soils demonstrated methanogenic potential, we detected no significant changes in CH₄ production rates, lag times, methanogenic pathways, diversity, or population sizes in soils that were exposed to either 20 or 30% elevated ambient CO₂. Our findings suggest that the methanogenic potential of the soil and the methanogenic community might be insensitive to changes in atmospheric CO₂ concentrations, at least not on a decadal timescale. Thus, if our results can be extrapolated to other temporarily flooded or even wetland ecosystems, the often-observed increase in CH₄ emissions under elevated CO₂ may simply be due to an increase in labile-C input via living root and increasing fresh litter deposition, but not due to shifts in the microbial population characteristics. This could make it easier to model and extrapolate the global effect of elevated CO₂ on terrestrial CH₄ emissions.     

The impact of long-term CO2 enrichment and moisture levels on soil microbial community structure and enzyme activities

Soil organic matter is the most important reservoir of terrestrial organic C and minor changes in the balance may have a significant impact on the climate. However, the response of microbial decomposers of soil C to global changes is not fully apprehended. This is particularly the case with regard to the interactive effects of the various climatic changes. Here, we present data from the Giessen Free Air CO₂ Experiment (Gi-FACE, University of Giessen, Germany) in which the CO₂ concentration at a grassland site was increased by 20% relative to atmospheric levels during a period of 10 years. The site included a slope that resulted in differences in average soil moisture. The effects of CO₂ and soil moisture on soil microbial community structure, measured by Denaturing Gradient Gel Electrophoresis (DGGE), PhosphoLipid Fatty Acids (PLFA) and enzyme activity profiles were determined. Total carbon, nitrogen and phosphorous contents were also determined. Soil moisture explained a large part of the variance in the microbial community structure data, by affecting fungi and bacteria. Furthermore, the CO₂ treatment had no significant effect on either overall PLFA or DGGE profiles, despite the fact that the fungal:bacterial PLFA ratio was altered. Overall enzyme activity profiles were also only affected by soil moisture levels, although the CO₂ treatment induced a significant increase of the acid phosphatase activity. Finally, neither soil moisture nor elevated CO₂ induced changes in the soil C stock.     

Priming depletes soil carbon and releases nitrogen in a scrub-oak ecosystem exposed to elevated CO2

Elevated atmospheric CO₂ tends to stimulate plant productivity, which could either stimulate or suppress the processing of soil carbon, thereby feeding back to atmospheric CO₂ concentrations. We employed an acid-hydrolysis-incubation method and a net nitrogen-mineralization assay to assess stability of soil carbon pools and short-term nitrogen dynamics in a Florida scrub-oak ecosystem after six years of exposure to elevated CO₂. We found that soil carbon concentration in the slow pool was 27% lower in elevated than ambient CO₂ plots at 0-10 cm depth. The difference in carbon mass was equivalent to roughly one-third of the increase in plant biomass that occurred in the same experiment. These results concur with previous reports from this ecosystem that elevated CO₂ stimulates microbial degradation of relatively stable soil organic carbon pools. Accordingly, elevated CO₂ increased net N mineralization in the 10-30 cm depth, which may increase N availability, thereby allowing for continued stimulation of plant productivity by elevated CO₂. Our findings suggest that soil texture and climate may explain the differential response of soil carbon among various long-term, field-based CO₂ studies. Increased mineralization of stable soil organic carbon by a CO₂-induced priming effect may diminish the terrestrial carbon sink globally.     

Long term CO2 enrichment stimulates N-mineralisation and enzyme activities in calcareous grassland

Elevated concentration of atmospheric carbon dioxide will affect carbon cycling in terrestrial ecosystems. Possible effects include increased carbon input into the soil through the rhizosphere, altered nutrient concentrations of plant litter and altered soil moisture. Consequently, the ongoing rise in atmospheric carbon dioxide might indirectly influence soil biota, decomposition and nutrient transformations.N-mineralisation and activities of the enzymes invertase, xylanase, urease, protease, arylsulfatase, and alkaline phosphatase were investigated in spring and summer in calcareous grassland, which had been exposed to ambient and elevated CO₂ concentrations (365 and 600 μl l⁻¹) for six growing seasons.In spring, N-mineralisation increased significantly by 30% at elevated CO₂, while there was no significant difference between treatments in summer (+3%). The response of soil enzymes to CO₂ enrichment was also more pronounced in spring, when alkaline phosphatase and urease activities were increased most strongly by 32 and 21%. In summer, differences of activities between CO₂ treatments were greatest in the case of urease and protease (+21 and +17% at elevated CO₂).The stimulation of N-mineralisation and enzyme activities at elevated CO₂ was probably caused by higher soil moisture and/or increased root biomass. We conclude that elevated CO₂ will enhance below-ground C- and N-cycling in grasslands.     

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.     

Impact of Land Use and Land Cover Changes on Organic Carbon Stocks in Mediterranean Soils (1956–2007)

During the last few decades, land use changes have largely affected the global warming process through emissions of CO₂. However, C sequestration in terrestrial ecosystems could contribute to the decrease of atmospheric CO₂ rates. Although Mediterranean areas show a high potential for C sequestration, only a few studies have been carried out in these systems. In this study, we propose a methodology to assess the impact of land use and land cover change dynamics on soil organic C stocks at different depths. Soil C sequestration rates are provided for different land cover changes and soil types in Andalusia (southern Spain). Our research is based on the analysis of detailed soil databases containing data from 1357 soil profiles, the Soil Map of Andalusia and the Land Use and Land Cover Map of Andalusia. Land use and land cover changes between 1956 and 2007 implied soil organic C losses in all soil groups, resulting in a total loss of 16·8 Tg (approximately 0·33 Tg y⁻¹). Afforestation increased soil organic C mostly in the topsoil, and forest contributed to sequestration of 8·62 Mg ha⁻¹ of soil organic C (25·4 per cent). Deforestation processes implied important C losses, particularly in Cambisols, Luvisols and Vertisols. The information generated in this study will be a useful basis for designing management strategies for stabilizing the increasing atmospheric CO₂ concentrations by preservation of C stocks and C sequestration. Copyright © 2012 John Wiley & Sons, Ltd.     

Effect of temperature on below-ground N-dynamics in a weedy model ecosystem at ambient and elevated atmospheric CO2 levels

Model multispecies terrestrial communities composed of four trophic levels (plants, herbivores, parasitoids, decomposers) were established in the Ecotron controlled environment facility. Two experimental runs enabled us to investigate the effects of enhanced temperature on below-ground microbial processes (N-mineralisation, urease, arginine deaminase, protease activity and potential denitrification) in both ambient and elevated (ambient +200 ppm) CO₂ atmospheres.The enzyme activities involved in nitrogen cycling showed weak responses to elevated temperature in both experimental runs. In the Ambient CO₂ Run, protease and arginine deaminase values tended to be lower in elevated temperature; on the other hand, N-mineralisation, urease and denitrification enzyme activity (DEA) were higher. In the Elevated CO₂ Run, all microbial variables showed higher activities at elevated temperature, although only the results for DEA and arginine deaminase were statistically significant. The interaction between higher temperature and elevated CO₂ weakly affected root growth and tissue C:N ratio, limiting feedbacks into the microbial community.Besides temperature and CO₂, substrate availability, water stress and successional development regulated the response of the soil microbes. The supply of organic carbon and nitrogen in the soil allowed plant growth and maintenance of the microbial population. Nitrogen competition between vegetation and microbes restricted net microbial growth. The increase of dissolved organic carbon (DOC) at higher CO₂ and temperature levels significantly favoured DEA. The high water regime in the soil also favoured DEA and inhibited oxidation of organic compounds, as indicated by low levels of enzyme activity. Additionally, water stress decreased rooting density in the soil; this resulted in negative feedback into microbial processes. We conclude that water stress and soil nitrogen deficiency caused an early levelling-off of both microbial population growth and activity rates during the early part of the model ecosystem's development.     

The effect of geocryological conditions and soil properties on the spatial variation in the CO<Subscript>2</Subscript> emission from flat-topped peat mounds in the isolated permafrost zone of Western Siberia

The active layer thickness, CO2 emission, and contents of organic substances (including the total organic carbon, labile carbon, and the carbon of microbial biomass) in the soils of flat-topped peat mounds in the area of the Nadym Experimental Station in the north of Western Siberia (experimental site CALM R1) are characterized by considerable spatial variability. The low values of the CО₂ emission are confined to the microelevations on the peatland surface. The high values of the emission (>200 mg CO₂/(m² h)) are typical of the soils with the highest content of the carbon of microbial biomass and the lowest content of the labile organic carbon. The soils of elevated flat-topped peat mounds statistically differ from the soils of waterlogged mires in the contents of total, labile, and microbial carbon and in the CO₂ emission values. Though the soils of elevated flat-topped peat mounds are characterized by the high content of the carbon of microbial biomass (4260 ± 880 mg С/kg soil), the CO₂ emission from them is low (158 ± 23 mg CO₂/(m² h)), which is explained by the structure of microbial communities in the cryogenic soils and by the effect of specific hydrothermic conditions.