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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

Natural N abundances in a tallgrass prairie ecosystem exposed to 8-y of elevated atmospheric CO2

After 8-y of elevated CO₂, we previously detected greater amounts of total soil nitrogen, suggesting that rates of ecosystem N flux into or out of tallgrass prairie had been altered. Denitrification and associative N fixation rates are the two primary biological processes that are known to control N loss and accumulation in tallgrass prairie soil. Therefore, our objective was to assess the natural abundance of plant and soil ¹⁵N isotopes as a cumulative index of potential change in efflux or influx of N into and out of the tallgrass prairie after 8-y of exposure to elevated CO₂. Aboveground plant delta ¹⁵N values of Andropogon gerardii were close to zero and more positive as a result of elevated CO₂, but whole-soil values at the 5-30 cm depth were significantly reduced (6.8 vs 7.3; P<0.05) under elevated CO₂-chamber (EC) relative to ambient CO₂- chamber (AC). Total, aboveground plant biomass, root-in-growth, extractable N, microbial biomass N, and soil pools collectively exhibited a range of delta ¹⁵N values from −2.8 to 7.3. Measurements of surface soil ¹⁵N indicate that a change in N inputs and outputs has occurred as a result of elevated atmospheric CO₂. In addition to possible changes in denitrification and N₂ fixation, other sources of N such as the re-translocation of N to the surface from deeper soil layers are needed to explain how soil N accrues in surface soils as a consequence of elevated CO₂. Our results support the notion that C accrual may promote N accrual, possibly driven by high plant and microbial N demand amplified by soil N limitation.     

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.     

Elevated CO2 concentration and nitrogen fertilisation effects on N2O and CH4 fluxes and biomass production of Phleum pratense on farmed peat soil

The effects of elevated CO₂ supply on N₂O and CH₄ fluxes and biomass production of Phleum pratense were studied in a greenhouse experiment. Three sets of 12 farmed peat soil mesocosms (10 cm dia, 47 cm long) sown with P. pratense and equally distributed in four thermo-controlled greenhouses were fertilised with a commercial fertiliser in order to add 2, 6 or 10 g N m⁻². In two of the greenhouses, CO₂ concentration was kept at atmospheric concentration (360 μmol mol⁻¹) and in the other two at doubled concentration (720 μmol mol⁻¹). Soil temperature was kept at 15 °C and air temperature at 20 °C. Natural lighting was supported by artificial light and deionized water was used to regulate soil moisture. Forage was harvested and the plants fertilised three times during the basic experiment, followed by an extra fertilisations and harvests. At the end of the experiment CH₄ production and CH₄ oxidation potentials were determined; roots were collected and the biomass was determined. From the three first harvests the amount of total N in the aboveground biomass was determined. N₂O and CH₄ exchange was monitored using a closed chamber technique and a gas chromatograph. The highest N₂O fluxes (on average, 255 μg N₂O m⁻² h⁻¹ during period IV) occurred just after fertilisation at high water contents, and especially at the beginning of the growing season (on average, 490 μg N₂O m⁻² h⁻¹ during period I) when the competition of vegetation for N was low. CH₄ fluxes were negligible throughout the experiment, and for all treatments the production and oxidation potentials of CH₄ were inconsequential. Especially at the highest rates of fertilisation, the elevated supply of CO₂ increased above- and below-ground biomass production, but both at the highest and lowest rates of fertilisation, decreased the total amount of N in the aboveground dry biomass. N₂O fluxes tended to be higher under doubled CO₂ concentrations, indicating that increasing atmospheric CO₂ concentration may affect N and C dynamics in farmed peat soil.     

Emission of N2O, N2 and CO2 from soil fertilized with nitrate: effect of compaction, soil moisture and rewetting

Soil compaction and soil moisture are important factors influencing denitrification and N₂O emission from fertilized soils. We analyzed the combined effects of these factors on the emission of N₂O, N₂ and CO₂ from undisturbed soil cores fertilized with (150 kg N ha⁻¹) in a laboratory experiment. The soil cores were collected from differently compacted areas in a potato field, i.e. the ridges (ρ_D=1.03 g cm⁻³), the interrow area (ρ_D=1.24 g cm⁻³), and the tractor compacted interrow area (ρ_D=1.64 g cm⁻³), and adjusted to constant soil moisture levels between 40 and 98% water-filled pore space (WFPS).High N₂O emissions were a result of denitrification and occurred at a WFPS≥70% in all compaction treatments. N₂ production occurred only at the highest soil moisture level (≥90% WFPS) but it was considerably smaller than the N₂O-N emission in most cases. There was no soil moisture effect on CO₂ emission from the differently compacted soils with the exception of the highest soil moisture level (98% WFPS) of the tractor-compacted soil in which soil respiration was significantly reduced. The maximum N₂O emission rates from all treatments occurred after rewetting of dry soil. This rewetting effect increased with the amount of water added. The results show the importance of increased carbon availability and associated respiratory O₂ consumption induced by soil drying and rewetting for the emissions of N₂O.     

Rhizodeposition-induced decomposition increases N availability to wild and cultivated wheat genotypes under elevated CO2

Elevated CO₂ may increase nutrient availability in the rhizosphere by stimulating N release from recalcitrant soil organic matter (SOM) pools through enhanced rhizodeposition. We aimed to elucidate how CO₂-induced increases in rhizodeposition affect N release from recalcitrant SOM, and how wild versus cultivated genotypes of wheat mediated differential responses in soil N cycling under elevated CO₂. To quantify root-derived soil carbon (C) input and release of N from stable SOM pools, plants were grown for 1 month in microcosms, exposed to ¹³C labeling at ambient (392 μmol mol⁻¹) and elevated (792 μmol mol⁻¹) CO₂ concentrations, in soil containing ¹⁵N predominantly incorporated into recalcitrant SOM pools. Decomposition of stable soil C increased by 43%, root-derived soil C increased by 59%, and microbial-¹³C was enhanced by 50% under elevated compared to ambient CO₂. Concurrently, plant ¹⁵N uptake increased (+7%) under elevated CO₂ while ¹⁵N contents in the microbial biomass and mineral N pool decreased. Wild genotypes allocated more C to their roots, while cultivated genotypes allocated more C to their shoots under ambient and elevated CO₂. This led to increased stable C decomposition, but not to increased N acquisition for the wild genotypes. Data suggest that increased rhizodeposition under elevated CO₂ can stimulate mineralization of N from recalcitrant SOM pools and that contrasting C allocation patterns cannot fully explain plant mediated differential responses in soil N cycling to elevated CO₂.     

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.     

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

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

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.     

Interactive effects of elevated CO2 and nitrogen fertilization levels on photosynthesized carbon allocation in a temperate spring wheat and soil system

Increasing atmospheric CO2 concentration impacts the terrestrial carbon(C) cycle by affecting plant photosynthesis, the flow of photosynthetically fixed C belowground, and soil C pool turnover. For managed agroecosystems, how and to what extent the interactions between elevated CO2 and N fertilization levels influence the accumulation of photosynthesized C in crops and the incorporation of photosynthesized C into arable soil are in urgent need of exploration.We conducted an experiment simulating elevated CO2 with spring wheat(Triticum aestivum L.) planted in growth chambers.13C-enriched CO2 with an identical 13C abundance was continuously supplied at ambient and elevated CO2 concentrations(350 and 600 μmol mol-1, respectively) until wheat harvest.Three levels of N fertilizer application(equivalent to 80, 120, and 180 kg N ha-1 soil) were supplied for wheat growth at both CO2 concentrations. During the continuous 62-d 13CO2 labeling period, elevated CO2 and increased N fertilizer application increased photosynthesized C accumulation in wheat by 14%–24% and 11%–20%, respectively, as indicated by increased biomass production, whereas the C/N ratio in the roots increased under elevated CO2 but declined with increasing N fertilizer application levels. Wheat root deposition induced 1%–2.5% renewal of soil C after 62 d of 13CO2 labeling. Compared to ambient CO2, elevated CO2 increased the amount of photosynthesized C incorporated into soil by 20%–44%. However, higher application rates of N fertilizer reduced the net input of root-derived C in soil by approximately 8% under elevated CO2. For the wheat-soil system, elevated CO2 and increased N fertilizer application levels synergistically increased the amount of photosynthesized C. The pivotal role of plants in photosynthesized C accumulation under elevated CO2 was thereby enhanced in the short term by the increased N application. Therefore, robust N management could mediate C cycling and sequestration by influencing the interactions between plants and soil in agroecosystems under elevated CO2.     

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。