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.     

Effects of elevated CO2 concentration on rhizodeposition from Lolium perenne grown on soil exposed to 9 years of CO2 enrichment

The effects of enriched CO₂ atmosphere on partitioning of recently assimilated carbon were investigated in a plant-soil-microorganism system in which Lolium perenne seedlings were planted into cores inserted into the resident soil within a sward that had been treated with elevated CO₂ for 9 consecutive years, under two N fertilisation levels (Swiss FACE experiment). The planted cores were excavated from the ambient (35 Pa pCO₂) and enriched (60 Pa pCO₂) rings at two dates, in spring and autumn, during the growing season. The cores were brought back to the laboratory for ¹⁴C labelling of shoots in order to trace the transfer of recently assimilated C both within the plant and to the soil and microbial biomass. At the spring sampling, high N supply stimulated shoot and total dry matter production. Consistently, high N enhanced the allocation of recently fixed C to shoots, and reduced it to belowground compartments. Elevated CO₂ had no consequences for DM or the pattern of C allocation. At the autumn sampling, at high N plot, yield of L. perenne was stimulated by elevated CO₂. Consistently, ¹⁴C was preferentially allocated aboveground and, consequently belowground recent C allocation was depressed and rhizodeposition reduced. At both experimental periods, total soil C content was similar in all treatments, providing no evidence for soil carbon sequestration in the Swiss Free Air CO₂ Enrichment experiment (FACE) after 9 years of enrichment. Recently assimilated C and soil C were mineralised faster in soils from enriched rings, suggesting a CO₂-induced shift in the microbial biomass characteristics (structure, diversity, activity) and/or in the quality of the root-released organic compounds.     

Effects of long-term CO2 fumigation on fungal communities in a temperate forest soil

This study aimed at determining the impact of long-time elevated CO₂ fumigation on fungal communities in a temperate forest soil. In addition to the CO₂ concentration, both time and its interaction with the CO₂ affected the activity of 1,4-β-N-acetylglucosaminidase that is mainly of the fungal origin in the soil. No significant change in Shannon's indexes (from 18S rDNA-PCR-DGGE) was observed between the ambient and elevated CO₂ treatments. Analysis of time-course indicated that the succession of soil fungal community was altered by the elevated CO₂ fumigation, and the variations in the soil samples under Pinus koraiensis Sieb. et Zucc were larger than those under the Pinus sylvestriformis (Takenouchi) T. Wang ex Cheng samples. The results suggest that the increase in atmospheric CO₂ concentrations could alter the temporal patterning of soil fungal communities.     

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.     

Relationship between soil CO2 concentrations and forest-floor CO2 effluxes

To better understand the biotic and abiotic factors that control soil CO₂ efflux, we compared seasonal and diurnal variations in simultaneously measured forest-floor CO₂ effluxes and soil CO₂ concentration profiles in a 54-year-old Douglas fir forest on the east coast of Vancouver Island. We used small solid-state infrared CO₂ sensors for long-term continuous real-time measurement of CO₂ concentrations at different depths, and measured half-hourly soil CO₂ effluxes with an automated non-steady-state chamber. We describe a simple steady-state method to measure CO₂ diffusivity in undisturbed soil cores. The method accounts for the CO₂ production in the soil and uses an analytical solution to the diffusion equation. The diffusivity was related to air-filled porosity by a power law function, which was independent of soil depth. CO₂ concentration at all depths increased with increase in soil temperature, likely due to a rise in CO₂ production, and with increase in soil water content due to decreased diffusivity or increased CO₂ production or both. It also increased with soil depth reaching almost 10 mmol mol⁻¹ at the 50-cm depth. Annually, soil CO₂ efflux was best described by an exponential function of soil temperature at the 5-cm depth, with the reference efflux at 10 °C (F₁₀) of 2.6 μmol m⁻² s⁻¹ and the Q₁₀ of 3.7. No evidence of displacement of CO₂-rich soil air with rain was observed.Effluxes calculated from soil CO₂ concentration gradients near the surface closely agreed with the measured effluxes. Calculations indicated that more than 75% of the soil CO₂ efflux originated in the top 20 cm soil. Calculated CO₂ production varied with soil temperature, soil water content and season, and when scaled to 10 °C also showed some diurnal variation. Soil CO₂ efflux and concentrations as well as soil temperature at the 5-cm depth varied in phase. Changes in CO₂ storage in the 0–50 cm soil layer were an order of magnitude smaller than measured effluxes. Soil CO₂ efflux was proportional to CO₂ concentration at the 50-cm depth with the slope determined by soil water content, which was consistent with a simple steady-state analytical model of diffusive transport of CO₂ in the soil. The latter proved successful in calculating effluxes during 2004.     

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.     

A new technique to measure soil CO2 efflux at constant CO2 concentration

A new principle for measuring soil CO₂ efflux at constant ambient concentration is introduced. The measuring principle relies on the continuous absorption of CO₂ within the system to achieve a constant CO₂ concentration inside the soil chamber at ambient level, thus balancing the amount of CO₂ entering the soil chamber by diffusion from the soil. We report results that show reliable soil CO₂ efflux measurements with the new system. The novel measuring principle does not disturb the natural gradient of CO₂ within the soil, while allowing for continuous capture of the CO₂ released from the soil. It therefore holds great potential for application in simultaneous measurements of soil CO₂ efflux and its δ¹³C, since both variables show sensitivity to a distortion of the soil CO₂ profile commonly found in conventional chamber techniques.     

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.     

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.     

Effect of elevated CO2 on soil N dynamics in a temperate grassland soil

The response of terrestrial ecosystems to elevated atmospheric CO₂ is related to the availability of other nutrients and in particular to nitrogen (N). Here we present results on soil N transformation dynamics from a N-limited temperate grassland that had been under Free Air CO₂ Enrichment (FACE) for six years. A ¹⁵N labelling laboratory study (i.e. in absence of plant N uptake) was carried out to identify the effect of elevated CO₂ on gross soil N transformations. The simultaneous gross N transformation rates in the soil were analyzed with a ¹⁵N tracing model which considered mineralization of two soil organic matter (SOM) pools, included nitrification from NH₄⁺ and from organic-N to NO₃⁻ and analysed the rate of dissimilatory NO₃⁻ reduction to NH₄⁺ (DNRA). Results indicate that the mineralization of labile organic-N became more important under elevated CO₂. At the same time the gross rate of NH₄⁺ immobilization increased by 20%, while NH₄⁺ oxidation to NO₃⁻ was reduced by 25% under elevated CO₂. The NO₃⁻ dynamics under elevated CO₂ were characterized by a 52% increase in NO₃⁻ immobilization and a 141% increase in the DNRA rate, while NO₃⁻ production via heterotrophic nitrification was reduced to almost zero. The increased turnover of the NH₄⁺ pool, combined with the increased DNRA rate provided an indication that the available N in the grassland soil may gradually shift towards NH₄⁺ under elevated CO₂. The advantage of such a shift is that NH₄⁺ is less prone to N losses, which may increase the N retention and N use efficiency in the grassland ecosystem under elevated CO₂.     

Denitrification in grass swards is increased under elevated atmospheric CO2

Emissions of N₂O and N₂ were measured from Lolium perenne L. swards under ambient (36 Pa) and elevated (60 Pa) atmospheric CO₂ at the Swiss free air carbon dioxide enrichment experiment following application of 11.2 g N m⁻² as ¹⁵NH₄¹⁵NO₃ or ¹⁴NH₄¹⁵NO₃ (1 at.% excess ¹⁵N). Total denitrification (N₂O+N₂) was increased under elevated pCO₂ with emissions of 6.2 and 19.5 mg ¹⁵N m⁻² measured over 22 d from ambient and elevated pCO₂ swards, respectively, supporting the hypothesis that increased belowground C allocation under elevated pCO₂ provides the energy for denitrification. Nitrification was the predominant N₂O producing process under ambient pCO₂ whereas denitrification was predominant under elevated pCO₂. The N₂-to-N₂O ratio was often higher under elevated pCO₂ suggesting that previous estimates of gaseous N losses based only on N₂O emissions have greatly underestimated the loss of N by denitrification.     

Impact of atmospheric CO2 and plant life forms on soil microbial activities

From the global change perspective, increase of atmospheric CO₂ and land cover transformation are among the major impacts caused by human activities. In this study, we are addressing the combined issues of the effect of CO₂ concentration increase and plant type on soil microbial activities by asking how annual and perennial plant groups affect soil microbial processes under elevated CO₂. The experimental design used a mix of species of different growth forms for both annuals and perennials. Our objective was: (1) to determine how two years of annual or perennial plant cover and CO₂ enrichment could affect Mediterranean soil microbial processes; (2) to test the resistance and the resilience of these soil functional processes after a natural perturbation. We determined the effects of 2 years atmospheric CO₂ enrichment on soil potential respiration (SIR), denitrification (DEA) and nitrification (NEA) activities. We could not find any significant effect of CO₂ increase on SIR, DEA and NEA. However, we found a strong effect of the plant cover type, i.e. annuals versus perennials, on the potential microbial activity related to N cycling. DEA and NEA were significantly higher in soil under annual plants while SIR was not significantly different. To determine whether these changes would survive a natural perturbation, we carried out a rain event experiment once the experimental treatments (i.e. different plant cover and atmospheric CO₂ concentration) were stopped. The soil potential respiration, as expressed by the SIR, was not affected and remained stable. DEA rates converged rapidly under annuals and perennials after the rain event. Under both annuals and perennials NEA increased significantly after the rain event but remained significantly higher in the soil with annual plants. The relative change of the soil microbial processes induced by annual and perennial plants was inversely related to the density and the diversity of the corresponding microbial functional groups.     

Carbon dynamics in a temperate grassland soil after 9 years exposure to elevated CO2 (Swiss FACE)

Elevated pCO₂ increases the net primary production, C/N ratio, and C input to the soil and hence provides opportunities to sequester CO₂-C in soils to mitigate anthropogenic CO₂. The Swiss 9 y grassland FACE (free air carbon-dioxide enrichment) experiment enabled us to explore the potential of elevated pCO₂ (60 Pa), plant species (Lolium perenne L. and Trifolium repens L.) and nitrogen fertilization (140 and 540 kg ha⁻¹ y⁻¹) on carbon sequestration and mineralization by a temperate grassland soil. Use of ¹³C in combination with respired CO₂ enabled the identification of the origins of active fractions of soil organic carbon. Elevated pCO₂ had no significant effect on total soil carbon, and total soil carbon was also independent of plant species and nitrogen fertilization. However, new (FACE-derived depleted ¹³C) input of carbon into the soil in the elevated pCO₂ treatments was dependent on nitrogen fertilization and plant species. New carbon input into the top 15 cm of soil from L. perennne high nitrogen (LPH), L. perenne low nitrogen (LPL) and T. repens low nitrogen (TRL) treatments during the 9 y elevated pCO₂ experiment was 9.3±2.0, 12.1±1.8 and 6.8±2.7 Mg C ha⁻¹, respectively. Fractions of FACE-derived carbon in less protected soil particles >53 μm in size were higher than in <53 μm particles. In addition, elevated pCO₂ increased CO₂ emission over the 118 d incubation by 55, 61 and 13% from undisturbed soil from LPH, LPL and TRL treatments, respectively; but only by 13, 36, and 18%, respectively, from disturbed soil (without roots). Higher input of new carbon led to increased decomposition of older soil organic matter (priming effect), which was driven by the quantity (mainly roots) of newly input carbon (L. perenne) as well as the quality of old soil carbon (e.g. higher recalcitrance in T. repens). Based on these results, the potential of well managed and established temperate grassland soils to sequester carbon under continued increasing concentrations of atmospheric CO₂ appears to be rather limited.     

不同水分条件下CO2浓度升高对冬小麦碳氮转运的影响

CO2浓度升高对作物的影响日益受到重视,水分是作物生长的必要条件之一。冬小麦是我国的主要粮食作物之一,阐明高CO2浓度和水分条件互作对冬小麦碳氮转运的影响,对客观认识气候变化背景下作物的水分管理及肥料施用具有实际指导意义。本研究利用开放式CO2富集系统(FACE)平台,以冬麦品种‘中麦175’为试验材料,采用盆栽试验方法,研究了不同CO2浓度[正常浓度(391±40)μmol·mol?1和高浓度(550±60)μmol·mol?1]及水分条件(湿润条件和干旱条件,即75%和55%田间土壤最大持水量)的冬小麦花前碳氮积累及花后碳氮转运的规律特征。结果表明:湿润条件下,与正常CO2浓度相比,高CO2浓度促进冬小麦地上部干物质及碳氮积累,开花期增幅分别为18.1%、16.5%、14.9%,成熟期增幅分别为6.6%、1.3%、4.5%,并提高碳氮转运能力及对籽粒贡献率,转运量、转运率及对籽粒贡献率的增幅碳素依次为39.3%、20.0%、30.0%,氮素依次为19.1%、3.8%、10.8%。干旱条件下,与正常CO2浓度相比,高CO2浓度对地上部碳氮积累有一定的促进作用,开花期和成熟期碳积累量分别增加3.0%和10.7%,氮积累量分别增加0和15.8%;但高CO2浓度阻碍了碳氮的转运,转运量、转运率降幅碳素分别为10.2%、12.8%,氮素分别为7.2%、7.1%;碳氮对籽粒贡献率则变化不同,碳降低14.4%,而氮升高31.3%。干旱及高CO2浓度互作与湿润条件正常CO2浓度处理相比,冬小麦碳素转运对籽粒贡献率降低更明显,地上部碳素转运量、转运率及对籽粒贡献率降幅分别为36.2%、16.9%、22.3%,但提高了氮素转运对籽粒贡献率,氮素转运量及转运率分别降低35.7%、15.2%,对籽粒贡献率增加7.0%。综合而言,高CO2浓度可促进冬小麦碳氮积累及其在花后向籽粒的转运,水分不足可能成为主要的物质转运障碍因子,限制CO2促进作用发挥。     

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.     

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.     

Elevated CO2 stimulates N2O emissions in permanent grassland

To evaluate climate forcing under increasing atmospheric CO₂ concentrations, feedback effects on greenhouse gases such as nitrous oxide (N₂O) with a high global warming potential should be taken into account. This requires long-term N₂O flux measurements because responses to elevated CO₂ may vary throughout annual courses. Here, we present an almost 9 year long continuous N₂O flux data set from a free air carbon dioxide enrichment (FACE) study on an old, N-limited temperate grassland. Prior to the FACE start, N₂O emissions were not different between plots that were later under ambient (A) and elevated (E) CO₂ treatments, respectively. However, over the entire experimental period (May 1998–December 2006), N₂O emissions more than doubled under elevated CO₂ (0.90 vs. 2.07 kg N₂O-N ha⁻¹ y⁻¹ under A and E, respectively). The strongest stimulation occurred during vegetative growth periods in the summer when soil mineral N concentrations were low. This was surprising because based on literature we had expected the highest stimulation of N₂O emissions due to elevated CO₂ when mineral N concentrations were above background values (e.g. shortly after N application in spring). N₂O emissions under elevated CO₂ were moderately stimulated during late autumn–winter, including freeze–thaw cycles which occurred in the 8th winter of the experiment. Averaged over the entire experiment, the additional N₂O emissions caused by elevated CO₂ equaled 4738 kg CO₂-equivalents ha⁻¹, corresponding to more than half a ton (546 kg) of CO₂ ha⁻¹ which has to be sequestered annually to balance the CO₂-induced N₂O emissions. Without a concomitant increase in C sequestration under rising atmospheric CO₂ concentrations, temperate grasslands may be converted into greenhouse gas sources by a positive feedback on N₂O emissions. Our results underline the need to include continuous N₂O flux measurements in ecosystem-scale CO₂ enrichment experiments.     

C and N in soil organic matter density fractions under elevated atmospheric CO2: Turnover vs. stabilization

Turnover of C and N in an arable soil under Free Air Carbon Dioxide (FACE) experiment was studied by the use of ¹³C natural abundance and ¹⁵N-labeled fertilizers. Wheat was kept four growing seasons under ambient and elevated CO₂ concentrations and fertilized for three growing seasons. Density fractionation of soil organic matter (SOM) allowed to track ¹³C and ¹⁵N in free particulate organic matter (fPOM; <1.6 g cm⁻³), particulate organic matter occluded within aggregates with two densities (oPOM 1.6, oPOM 1.6-2.0 g cm⁻³), and in mineral-associated organic matter (>2.0 g cm⁻³) fractions. Elevated CO₂ and N fertilization did not significantly affect C and N contents in the bulk soil. Calculated mean residence time (MRT) of C and N revealed the qualitative differences of SOM density fractions: (i) the shortest MRT_C and MRT_N in fPOM confirmed high availability of this fraction to decomposition. Larger C/N ratio of fPOM under elevated vs. ambient CO₂ indicated an increasing recalcitrance of FACE-derived plant residues. (ii) There was no difference in MRT of C and N between lighter and heavier oPOMs probably due to short turnover time of soil aggregates which led to oPOM mixing. The increase of MRT_C and MRT_N in both oPOMs during the experiment confirmed the progressive degradation of organic material within aggregates. (iii) Constant turnover rates of C in the mineral fraction neither confirmed nor rejected the assumed stabilization of SOM to take place in the mineral fraction. Moreover, a trend of decreasing of C and N amounts in the Min fraction throughout the experiment was especially pronounced for C under elevated CO₂. Hence, along with the progressive increase of C_FACE in the Min fraction the overall losses of C under elevated CO₂ may occur at the expense of older “pre-FACE” C.     

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.     

Palatability trials on hardwood leaf litter grown under elevated CO2: a stable carbon isotope study

The palatability to isopods and microbes of a broad range of hardwood leaf litter, derived from three field CO₂-enrichment experiments in the USA, was investigated, using δ¹³C, to trace the C flow from litter to isopods and to CO₂ respired by microbial decomposition. Leaf litter grown under elevated CO₂ had δ¹³C values ranging from −39 to −45‰, which were significantly different from ambient litter δ¹³C values of around −30‰. Litter palatability to isopods of the Porcellio sp. was tested by incubating ambient- and elevated-CO₂ litter, and a mixture of the two, in the presence of isopods for 14 days, under environmentally controlled conditions; δ¹³C was measured on litter and isopods' body before and after incubation. In an additional experiment, litter was incubated in the absence of fauna for 30 days, and on five occasions the δ¹³C of the CO₂ respired from litter was measured. The ¹³C label was clearly carried from the litter source to the isopods' bodies, and their faeces. For microbial-respired CO₂, δ¹³C was significantly higher than that of the litter source, suggesting preferential degradation of substrates enriched in ¹³C as compared to those in the overall litter. With the exception of Quercus myrtifolia leaf litter, elevated CO₂ did not affect the palatability to isopods nor the microbial degradation of any of the litters, possibly as a result of unaltered litter N concentration. However, significant differences in litter palatability and decay rates were observed among the different species. With this study, the use of isotopically labelled litter material was confirmed as a key methodology that can significantly contribute to the advancement of the understanding of litter decomposition and of the quantification of C fluxes in the process.