Differentiating sources of CO2 from organic soil under bioenergy crop cultivation: A field-based approach using 14C

We tested here a plant-soil system to separate recent, plant-derived and native, soil-derived carbon in soil respiration. The approach uses a perennial crop cultivated on an organic soil where upper soil layers have been removed as a result of peat extraction. There, the ¹⁴C signal from native organic matter is highly depleted compared to that in vegetation established at the site after peat extraction ceased. Radiocarbon was analyzed in carbon dioxide respired from soil over one growing season, and a two-pool isotope mixing model was applied to calculate the relative contribution of old vs. new carbon sources. The analysis showed that the approach is reliable for source partitioning with isotopes. After six years of cultivation, old peat decomposition contributed less to total soil respiration than respiration of recent plant material (30% vs. 70% on average, respectively), but the relative proportions were highly variable over the growing season. The approach offers a new possibility to follow the fate of old, native soil organic matter in highly organic soils.     

Species richness of ectomycorrhizal hyphal necromass increases soil CO2 efflux under laboratory conditions

The ectomycorrhizal mycelium is a large component of boreal and temperate forest soil microbial biomass and the resulting necromass is likely to be an important source of nutrients for saprotrophic microorganisms. Here we test the effects of species richness of ectomycorrhizal mycelial biomass on short-term CO₂ efflux by amending forest soil with necromass from 8 fungal species added separately and in mixtures of 2, 4 and 8 species. All additions of necromass rapidly increased soil CO₂ efflux compared to unamended controls but CO₂ efflux increased significantly with species richness. Efflux of CO₂ did not correlate with the carbon (C) or nitrogen (N) contents or the C:N ratio of the added necromass. The study demonstrates that species diversity of dead ectomycorrhizal fungal hyphae can have important consequences for soil CO₂ efflux, and suggests decomposition of hyphae is regulated by specific constituents of the nutrient pools in the necromass rather than the total quantities added.     

Isotope (C and C) analysis of deep peat CO2 using a passive sampling technique

We developed and tested a new method to collect CO₂ from the surface to deep layers of a peatland for radiocarbon analysis. The method comprises two components: i) a probe equipped with a hydrophobic filter that allows entry of peat gases by diffusion, whilst simultaneously excluding water, and, ii) a cartridge containing zeolite molecular sieve that traps CO₂ passively. We field tested the method by sampling at depths of between 0.25 and 4 m at duplicate sites within a temperate raised peat bog. CO₂ was trapped at a depth-dependent rate of between ∼0.2 and 0.8 ml d⁻¹, enabling sufficient CO₂ for routine ¹⁴C analysis to be collected when left in place for several weeks. The age of peatland CO₂ increased with depth from modern to ∼170 BP for samples collected from 0.25 m, to ∼4000 BP at 4 m. The CO₂ was younger, but followed a similar trend to the age profile of bulk peat previously reported for the site (Langdon and Barber, 2005). δ¹³C values of recovered CO₂ increased with depth. CO₂ collected from the deepest sampling probes was considerably ¹³C-enriched (up to ∼+9‰) and agreed well with results reported for other peatlands where this phenomenon has been attributed to fermentation processes. CO₂ collected from plant-free static chambers at the surface of the mire was slightly ¹⁴C-enriched compared to the contemporary atmosphere, suggesting that surface CO₂ emissions were predominantly derived from carbon fixed during the post-bomb era. However, consistent trends of enriched ¹³C and depleted ¹⁴C in chamber CO₂ between autumn and winter samples were most likely explained by an increased contribution of deep peat CO₂ to the surface efflux in winter. The passive sampling technique is readily portable, easy to install and operate, causes minimal site disturbance, and can be reliably used to collect peatland CO₂ from a wide range of depths.     

长期施肥下红壤旱地土壤CO2排放及碳平衡特征

在国家肥力网红壤旱地长期定位试验地上,采用静态箱/气相色谱法测定土壤CO2排放速率,同时利用根去除法区分根系对土壤呼吸的贡献,通过计算净生态系统生产力（NEP）,判断长期不同施肥下红壤旱地农田碳汇强度。结果表明,小麦、玉米生长季各处理的土壤和土体呼吸速率随着作物生长、温度升高均呈现明显的季节变化规律;玉米生长季土壤和土体累积呼吸量大于小麦生长季,小麦、玉米生长季均以NPKM处理土壤和土体呼吸累积呼吸量最大,且显著高于其它处理（P0.05）,NP和NPK处理次之,CK和NK处理最小（P0.05）;小麦、玉米生长季各处理根际呼吸占土壤呼吸的比例分别为7.6 %～17.4 %、4.7%～16.6 %,均以NPKM处理根际呼吸贡献率最大;小麦季NPKM处理、玉米季CK和NPKM处理的NEP值为负,是大气CO2的汇,且NPKM处理的净初级生产力与土壤呼吸的比值（NPP/Rs）最大,其它处理NEP值均为正,是大气CO2的源。有机无机肥配施（NPKM）相比其它处理具有较强的碳汇功能,是红壤旱地比较合理的施肥措施。     

CO2 emission from soils under different uses and flooding conditions

The emission of CO₂ from Galician (NW Spain) forest, grassland and cropped soils was studied in a laboratory experiment, at different temperatures (10-35 °C) and at moisture contents of 100% and 160% of the field capacity (FC) of each soil (the latter value corresponds to saturated conditions, and represents between 120% and 140% of the water holding capacity, depending on the soil). In the forest soil, respiration in the flooded samples at all temperatures was lower than that at 100% field capacity. In the agricultural (grassland and cropped) soils the emission was higher (particularly at the highest incubation temperatures) in the soils wetted to 160% of the field capacity than in those wetted to 100% of the field capacity. In all cases the emission followed first order kinetics and the mineralization constants increased exponentially with temperature. In the forest soil, the Q₁₀ values were almost the same in the soils incubated at the two moisture contents. The grassland and cropped soils displayed different responses, as the Q₁₀ values were higher in the soils at 160% than in those at 100% of field capacity. In addition, and particularly at the highest temperatures, the rate of respiration increased sharply 9 and 17 days after the start of the incubation in the grassland and in the cropped soil, respectively. The above-mentioned anomalous response of the grassland and cropped soils under flooding conditions may be related to the agricultural use of the soils and possibly to the intense use of organic fertilizers in these soils (more than 150 kg N ha⁻¹ year⁻¹ added as cattle slurry or manure, respectively, in the grassland and cropped soils). The observed increase in respiration may either be related to the development of thermophilic facultative anaerobic microbes or to the formation during the incubation period of a readily metabolizable substrate, possibly originating from the remains of organic fertilizers, made accessible by physicochemical processes that occurred during incubation under conditions of high moisture.     

Soil CO2 efflux following rotary tillage of a tropical soil

Stopping the increase of atmospheric CO₂ level is an important task and information on how to implement adjustments on tillage practices could help lower soil CO₂ emissions would be helpful. We describe how rotary tiller use on a red latosol affected soil CO₂ efflux. The impact of changing blade rotation speed and rear shield position on soil CO₂ efflux was investigated. Significant differences among treatments were observed up to 10 days after tillage. Cumulative CO₂ efflux was as much as 40% greater when blade rotation of 216 rpm and a lowered rear shield was compared to blade rotation of 122 rpm and raised shield. This preliminary work suggests that adjusting rotary tiller settings could help reduce CO₂ efflux close to that of undisturbed soil, thereby helping to conserve soil carbon in tropical environments.     

Measurement of soil CO2 efflux using soda lime absorption: both quantitative and reliable

Measurement of soil CO₂ efflux using a non-flow-through steady-state (NFT-SS) chamber with alkali absorption of CO₂ by soda lime was tested and compared with a flow-through non-steady-state (FT-NSS) IRGA method to assess suitability of using soda lime for field monitoring over large spatial scales and integrated over a day. Potential errors and artifacts associated with the soda lime chamber method were investigated and improvements made. The following issues relating to quantification and reliable measurement of soil CO₂ efflux were evaluated: (i) absorption capacity of the soda lime, (ii) additional and thus artifactual absorption of CO₂ by soda lime during the experimental procedure, (iii) variation in the CO₂ concentration inside the chamber headspace, and (iv) effects of chamber closure on soil CO₂ efflux. Soil CO₂ efflux, as measured using soda lime (with a range of quantities: 50, 100, and 200 g per 0.082 m² ground area enclosed in chamber), was compared with transient IRGA measurements as a reference method that is based on well-established physical principles, using several forms of spatial and temporal comparisons. Natural variation in efflux rates ranged from 2 to 5.5 g C m⁻² day⁻¹ between different chambers and over different days. A comparison of the IRGA-based assay with measurement based on soda lime yielded an overall correlation coefficient of 0.82. The slope of the regression line was not significantly different from the 1:1 line, and the intercept was not significantly different from the origin. This result indicated that measurement of CO₂ efflux by soda lime absorption was quantitatively similar and unbiased in relation to the reference method. The soda lime method can be a highly practical method for field measurements if implemented with due care (in terms of drying and weighing soda lime, and in minimizing leakages), and validated for specific field conditions. A detailed protocol is presented for use of the soda lime method for measurement of CO₂ efflux from field soils.     

Spatial and temporal variability of soil CO2 emission in a sugarcane area under green and slash-and-burn managements

Soil management causes changes in physical, chemical, and biological properties that consequently affect soil CO₂ emission (FCO2). Here, we studied the soil carbon dynamics in areas with sugarcane production in southern Brazil under two different sugarcane management systems: green (G), consisting of mechanized harvesting that produces a large amount of crop residues left on the soil surface, and slash-and-burn (SB), in which the residues are burned before manual harvest, leaving no residues on the soil surface. The study was conducted during the period after harvest in two side-by-side grids installed in adjacent areas, having 60 points each. The aim was to characterize the temporal and spatial variability of FCO2, and its relation to soil temperature and soil moisture, in a red latosol (Oxisol) where G and SB management systems have been recently used. Mean FCO2 emission was 39% higher in the SB plot (2.87 μmol m⁻² s⁻¹) when compared to the G plot (2.06 μmol m⁻² s⁻¹) throughout the 70-day period after harvest. A quadratic equation of emissions versus soil moisture was able to explain 73% and 50% of temporal variability of FCO2 in SB and G, respectively. This seems to relate to the sensitivity of FCO2 to precipitation events, which caused a significant increase in SB emissions but not in G-managed area emissions. FCO2 semivariogram models were mostly exponential in both areas, ranging from 72.6 to 73.8 m and 63.0 to 64.7 m for G and SB, respectively. These results indicate that the G management system results in more homogeneous FCO2 when spatial and temporal variability are considered. The spatial variability analysis of soil temperature and soil moisture indicates that those parameters do not adequately explain the changes in spatial variability of FCO2, but emission maps are clearly more homogeneous after a drought period when no rain has occurred, in both sites.     

Net fluxes of CO2, but not N2O or CH4, are affected following agronomic-scale additions of urea to prairie and arable soils

While experimental addition of nitrogen (N) tends to enhance soil fluxes of carbon dioxide (CO₂), methane (CH₄), and nitrous oxide (N₂O), it is not known if lower and agronomic-scale additions of urea-N applied also enhance trace gas fluxes, particularly for semi-arid agricultural lands in the northern plains. We aimed to test if this were true at agronomic rates [low (11 kg N ha⁻¹), moderate (56 kg N ha⁻¹), and high (112 kg N ha⁻¹)] for central North Dakota arable and prairie soils using intact soil cores to minimize disturbance and simulate field conditions. Additions of urea to cores incubated at 21 °C and 57% water-filled pore space enhanced fluxes of CO₂ but not CH₄ and N₂O. At low, moderate, and high urea-N, CO₂ fluxes were significantly greater than control but not fluxes of CH₄ and N₂O. The increases in CO₂ emission with rate of urea-N application indicate that agronomic-scale N inputs may stimulate microbial carbon cycling in these soils, and that the contribution of CO₂ to net greenhouse gas source strength following fertilization of semi-arid agroecosystems may at times be greater than contributions by N₂O and CH₄.     

间伐对杉木林土壤CO2通量的影响

Forest management is expected to influence soil CO₂ efflux (FCO₂) as a result of changes in microenvironmental conditions, soil microclimate, and root dynamics. Soil FCO₂ rate was measured during the growing season of 2006 in both thinning and non-thinning locations within stands ranging from 0 to 8 years after the most recent thinning in Chinese fir (Cunninghamia lanceolata (Lamb.) Hook) plantations in Huitong Ecosystem Research Station, Hunan, China. Soil temperature and moisture were also measured to examine relationships between FCO₂ and soil properties. Forest thinning resulted in huge changes in FCO₂ that varied with time since cutting. Immediately following harvest (year 0) FCO₂ in thinning area increased by about 30%, declined to 20%-27% below pre-cutting levels during years 4-6, and recovered to pre-cutting levels at 8 years post-cutting. A similar temporal pattern, but with smaller changes, was found in non-thinning locations. The initial increase in FCO₂ could be attributed to a combination of root decay, soil disturbance, and increased soil temperature in gaps, while the subsequent decrease and recovery to the death and gradual regrowth of active roots. Strong effects of soil temperature and soil water content on FCO₂ were found. Forest thinning mainly influenced FCO₂ through changes in tree root respiration, and the net result was a decrease in integrated FCO₂ flux through the entire felling cycle.     

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.     

Measuring forest floor CO2 fluxes in a Douglas-fir forest

CO₂ exchange was measured on the forest floor of a coastal temperate Douglas-fir forest located near Campbell River, British Columbia, Canada. Continuous measurements were obtained at six locations using an automated chamber system between April and December, 2000. Fluxes were measured every half hour by circulating chamber headspace air through a sampling manifold assembly and a closed-path infrared gas analyzer. Maximum CO₂ fluxes measured varied by a factor of almost 3 between the chamber locations, while the highest daily average fluxes observed at two chamber locations occasionally reached values near 15 μmol C m⁻² s⁻¹. Generally, fluxes ranged between 2 and 10 μmol C m⁻² s⁻¹ during the measurement period. CO₂ flux from the forest floor was strongly related to soil temperature with the highest correlation found with 5 cm depth temperature. A simple temperature dependent exponential model fit to the nighttime fluxes revealed Q₁₀ values in the normal range of 2–3 during the warmer parts of the year, but values of 4–5 during cooler periods. Moss photosynthesis was negligible in four of the six chambers, while at the other locations, it reduced daytime half-hourly net CO₂ flux by about 25%. Soil moisture had very little effect on forest floor CO₂ flux. Hysteresis in the annual relationship between chamber fluxes and soil temperatures was observed. Net exchange from the six chambers was estimated to be 1920±530 g C m⁻² per year, the higher estimates exceeding measurement of ecosystem respiration using year-round eddy correlation above the canopy at this site. This discrepancy is attributed to the inadequate number of chambers to obtain a reliable estimate of the spatial average soil CO₂ flux at the site and uncertainty in the eddy covariance respiration measurements.     

大气CO2浓度升高对植物 土壤系统地下过程影响的研究

综述了大气CO2浓度升高对根系、根际、根系分泌物、土壤呼吸和土壤物质转化和C、N循环影响的研究进展,阐述了有关实验的研究情况,以及它们在整个生态系统响应大气CO2浓度升高中的重要作用、目前研究中存在的争论、以及还需要研究的领域和方向及其研究的重要性。     

Dependence of soil respiration on soil moisture, clay content, soil organic matter, and CO2 uptake in dry grasslands

The effects of abiotic and biotic drivers on soil respiration (R_s) were studied in four grassland and one forest sites in Hungary in field measurement campaigns (duration of studies by sites 2-7 years) between 2000 and 2008. The sites are within a 100 km distance of each other, with nearly the same climate, but with different soils and vegetation. Soil respiration model with soil temperature (T_s) and soil water content (SWC) as independent variables explained larger part of variance (range 0.47-0.81) than the Lloyd and Taylor model (explained variance: 0.31-0.76). Direct effect of SWC on R_s at much smaller temporal and spatial scale (1.5 h, and a few meters, respectively) was verified.Soil water content optimal for R_s (SWC_opt) was shown to significantly (positively) depend on soil clay content, while parameter related to activation energy (E₀) was significantly (negatively) correlated to the total organic carbon content (TOC) in the upper 10 cm soil layer. Dependence of model parameters on soil properties could easily be utilized in models of soil respiration. The effect of current (a few hours earlier) assimilation rates on soil respiration after removing the effect of abiotic covariates (i.e. temperature and water supply) is shown. The correlation maximum between the R_s residuals (R_{s_res}, from the R_s (SWC, T_s) model) and net ecosystem exchange (NEE) was found at 13.5 h time lag at the sandy grassland. Incorporating the time-lagged effect of NEE on R_s into the model of soil respiration improved the agreement between the simulated vs. measured R_s data. Use of SWC_opt and E₀ parameters and consideration of current assimilation in soil respiration models are proposed.     

Quantification of priming and CO2 emission sources following the application of different slurry particle size fractions to a grassland soil

The highest emissions of CO₂ from soils and most pronounced priming effect (PE) from soils generally occur immediately after slurry application. However, the influence of different particle size slurry fractions on net soil C respiration dynamics and PE has not been studied. Therefore, a slurry separation technique based on particle sizes was used in the present study. Six distinct fractions (>2000, 425-2000, 250-425, 150-250, 45-150, <45 μm) were generated from two dairy slurries (one from cows fed a predominantly maize silage diet and the other from cows fed a grass silage diet) were applied to soil. During the first days of the 332 days experiment, all slurry fraction amendments significantly increased soil CO₂ effluxes (by 2-8 times) compared to the non-amended control. The increased CO₂ emission rates had a negative relationship with slurry particle size, but its duration was positively correlated with slurry particle size. The percentage of the cumulative CO₂ emitted was only higher in the first 8 days in the finest slurry particle sizes (<150 μm). The proportion of slurry-derived C emitted as CO₂ 2 h after addition to soil varied between 29% and 100% of total emitted CO₂-C. Generally, the proportion of slurry-derived C emitted initially decreased rapidly in the <250 μm fractions, but decreased more slowly or even increased in the >250 μm fractions. The overall contribution of slurry C to total CO₂ emissions was higher in smaller slurry particle size treatments in the first days after application. The addition of the various slurry fractions to soil caused both significant positive and negative PEs on the soil organic matter mineralization. The timing and type (positive or negative) of PE depended on the slurry particle size. Clearly, farm based separation pre-treatment leading to two or more fractions with different particle sizes has also the potential to reduce or modify short-term CO₂ emissions immediately after slurry application to soil.     

Linking N2O emission to soil mineral N as estimated by CO2 emission and soil C/N ratio

Based on the N₂O and CO₂ emission data concomitantly measured from agricultural upland fields around the world, we developed an empirical model as follows: cumulative N₂O emission = aexp[b*(E_CO2/S_cn + F_n)] (R²_adj = 0.85∼0.87), where E_CO2 is the rate of heterotrophic respiration from soils, S_cn is the soil C/N ratio, and F_n is the chemical fertilizer N rate. The model parameters derived from the data from the soils without receiving chemical fertilizers were significantly different from the ones from the fertilized soils. This model indicates that CO₂ emission and soil C/N ratio can be used as scaling parameters to produce regional or global inventories of N₂O emission from agricultural soils.     

Uncertainties in static closed chamber measurements of the carbon isotopic ratio of soil-respired CO2

The δ¹³C of soil-respired CO₂ (δ_r) is frequently determined using static closed chamber methods. δ_r is obtained as the intercept of the least squares linear regression of δ vs 1/C*, where measured δ¹³C-CO₂ (δ) and volume fraction of CO₂ (C*) values of chamber headspace samples are used. Theoretically, we show that the variance of the estimate of δ_r can be reduced by extending the 1/C* interval of the regression towards (i) higher or (ii) lower values, or (iii) distributing the 1/C* values optimally within the pre-selected headspace CO₂ sampling time period. Experimental applications of these approaches indicated that: (1) lowering the initial CO₂ level, thereby increasing 1/C*, yielded a positive bias to the δ_r result. (2) It was feasible to obtain lower variance in the δ_r estimate by lowering 1/C* values through extended CO₂ sampling time. We also recommend that each chamber is sampled only once, mainly because this allows freedom to select the sampling times, in order to optimize the distribution of 1/C* values.     

Tillage and wind effects on soil CO2 concentrations in muck soils

Rising atmospheric carbon dioxide (CO₂) concentrations from agricultural activities prompted the need to quantify greenhouse gas emissions to better understand carbon (C) cycling and its role in environmental quality. The specific objective of this work was to determine the effect of no-tillage, deep plowing and wind speeds on the soil CO₂ concentration in muck (organic) soils of the Florida Everglades. Miniature infrared gas analyzers were installed at 30 cm and recorded every 15 min in muck soil plowed with the Harrell Switch Plow (HSP) to 41 cm and in soil Not Tilled (NT), i.e., not plowed in last 9 months. The soil CO₂ concentration exhibited temporal dynamics independent of barometric pressure fluctuations. Loosening the soil resulted in a very rapid decline in CO₂ concentration as a result of “wind-induced” gas exchange from the soil surface. Higher wind speeds during mid-day resulted in a more rapid loss of CO₂ from the HSP than from the NT plots. The subtle trend in the NT plots was similar, but lower in magnitude. Tillage-induced change in soil air porosity enabled wind speed to affect the gas exchange and soil CO₂ concentration at 30 cm, literally drawing the CO₂ out of the soil resulting in a rapid decline in the CO₂ concentration, indicating more rapid soil carbon loss with tillage. At the end of the study, CO₂ concentrations in the NT plots averaged about 3.3% while that in the plowed plots was about 1.4%. Wind and associated aerodynamic pressure fluctuations affect gas exchange from soils, especially tilled muck soils with low bulk densities and high soil air porosity following tillage.     

Effects of elevated CO2 and O3 on soil respiration under ponderosa pine

Soil respiration represents the integrated response of plant roots and soil organisms to environmental conditions and the availability of C in the soil. A multi-year study was conducted in outdoor sun-lit controlled-environment chambers containing a reconstructed ponderosa pine/soil-litter system. The study used a 2×2 factorial design with two levels of CO₂ and two levels of O₃ and three replicates of each treatment. The objectives of our study were to assess the effects of long-term exposure to elevated CO₂ and O₃, singly and in combination, on soil respiration, fine root growth and soil organisms. Fine root growth and soil organisms were included in the study as indicators of the autotrophic and heterotrophic components of soil respiration. The study evaluated three hypotheses: (1) elevated CO₂ will increase C assimilation and allocation belowground increasing soil respiration; (2) elevated O₃ will decrease C assimilation and allocation belowground decreasing soil respiration and (3) as elevated CO₂ and O₃ have opposing effects on C assimilation and allocation, elevated CO₂ will eliminate or reduce the negative effects of elevated O₃ on soil respiration. A mixed-model covariance analysis was used to remove the influences of soil temperature, soil moisture and days from planting when testing for the effects of CO₂ and O₃ on soil respiration. The covariance analysis showed that elevated CO₂ significantly reduced the soil respiration while elevated O₃ had no significant effect. Despite the lack of a direct CO₂ stimulation of soil respiration, there were significant interactions between CO₂ and soil temperature, soil moisture and days from planting indicating that elevated CO₂ altered soil respiration indirectly. In elevated CO₂, soil respiration was more sensitive to soil temperature changes and less sensitive to soil moisture changes than in ambient CO₂. Soil respiration increased more with days from planting in elevated than in ambient CO₂. Elevated CO₂ had no effect on fine root biomass but increased abundance of culturable bacteria and fungi suggesting that these increases were associated with increased C allocation belowground. Elevated CO₂ had no significant effect on microarthropod and nematode abundance. Elevated O₃ had no significant effects on any parameter except it reduced the sensitivity of soil respiration to changes in temperature.