Soil 13C–15N dynamics in an N2-fixing clover system under long-term exposure to elevated atmospheric CO2

Reduced soil N availability under elevated CO₂ may limit the plant's capacity to increase photosynthesis and thus the potential for increased soil C input. Plant productivity and soil C input should be less constrained by available soil N in an N₂‐fixing system. We studied the effects of Trifolium repens (an N₂‐fixing legume) and Lolium perenne on soil N and C sequestration in response to 9 years of elevated CO₂ under FACE conditions. ¹⁵N‐labeled fertilizer was applied at a rate of 140 and 560 kg N ha^?1 yr^?1 and the CO₂ concentration was increased to 60 Pa pCO₂ using ¹³C‐depleted CO₂. The total soil C content was unaffected by elevated CO₂, species and rate of ¹⁵N fertilization. However, under elevated CO₂, the total amount of newly sequestered soil C was significantly higher under T. repens than under L. perenne. The fraction of fertilizer‐N (f_N) of the total soil N pool was significantly lower under T. repens than under L. perenne. The rate of N fertilization, but not elevated CO₂, had a significant effect on f_N values of the total soil N pool. The fractions of newly sequestered C (f_C) differed strongly among intra‐aggregate soil organic matter fractions, but were unaffected by plant species and the rate of N fertilization. Under elevated CO₂, the ratio of fertilizer‐N per unit of new C decreased under T. repens compared with L. perenne. The L. perenne system sequestered more ¹⁵N fertilizer than T. repens: 179 vs. 101 kg N ha^?1 for the low rate of N fertilization and 393 vs. 319 kg N ha^?1 for the high N‐fertilization rate. As the loss of fertilizer‐¹⁵N contributed to the ¹⁵N‐isotope dilution under T. repens, the input of fixed N into the soil could not be estimated. Although N₂ fixation was an important source of N in the T. repens system, there was no significant increase in total soil C compared with a non‐N₂‐fixing L. perenne system. This suggests that N₂ fixation and the availability of N are not the main factors controlling soil C sequestration in a T. repens system.     

Elevated CO2 increases microbial carbon substrate use and nitrogen cycling in Mojave Desert soils

Identifying soil microbial responses to anthropogenically driven environmental changes is critically important as concerns intensify over the potential degradation of ecosystem function. We assessed the effects of elevated atmospheric CO₂ on microbial carbon (C) and nitrogen (N) cycling in Mojave Desert soils using extracellular enzyme activities (EEAs), community‐level physiological profiles (CLPPs), and gross N transformation rates. Soils were collected from unvegetated interspaces between plants and under the dominant shrub (Larrea tridentata) during the 2004–2005 growing season, an above‐average rainfall year. Because most measured variables responded strongly to soil water availability, all significant effects of soil water content were used as covariates to remove potential confounding effects of water availability on microbial responses to experimental treatment effects of cover type, CO₂, and sampling date. Microbial C and N activities were lower in interspace soils compared with soils under Larrea, and responses to date and CO₂ treatments were cover specific. Over the growing season, EEAs involved in cellulose (cellobiohydrolase) and orthophosphate (alkaline phosphatase) degradation decreased under ambient CO₂, but increased under elevated CO₂. Microbial C use and substrate use diversity in CLPPs decreased over time, and elevated CO₂ positively affected both. Elevated CO₂ also altered microbial C use patterns, suggesting changes in the quantity and/or quality of soil C inputs. In contrast, microbial biomass N was higher in interspace soils than soils under Larrea, and was lower in soils exposed to elevated CO₂. Gross rates of NH₄⁺ transformations increased over the growing season, and late‐season NH₄⁺ fluxes were negatively affected by elevated CO₂. Gross NO₃⁻ fluxes decreased over time, with early season interspace soils positively affected by elevated CO₂. General increases in microbial activities under elevated CO₂ are likely attributable to greater microbial biomass in interspace soils, and to increased microbial turnover rates and/or metabolic levels rather than pool size in soils under Larrea. Because soil water content and plant cover type dominates microbial C and N responses to CO₂, the ability of desert landscapes to mitigate or intensify the impacts of global change will ultimately depend on how changes in precipitation and increasing atmospheric CO₂ shift the spatial distribution of Mojave Desert plant communities.     

The response of soil CO2 flux to changes in atmospheric CO2, nitrogen supply and plant diversity

We measured soil CO₂ flux over 19 sampling periods that spanned two growing seasons in a grassland Free Air Carbon dioxide Enrichment (FACE) experiment that factorially manipulated three major anthropogenic global changes: atmospheric carbon dioxide (CO₂) concentration, nitrogen (N) supply, and plant species richness. On average, over two growing seasons, elevated atmospheric CO₂ and N fertilization increased soil CO₂ flux by 0.57 µmol m^?2 s^?1 (13% increase) and 0.37 µmol m^?2 s^?1 (8% increase) above average control soil CO₂ flux, respectively. Decreases in planted diversity from 16 to 9, 4 and 1 species decreased soil CO₂ flux by 0.23, 0.41 and 1.09 µmol m^?2 s^?1 (5%, 8% and 21% decreases), respectively. There were no statistically significant pairwise interactions among the three treatments. During 19 sampling periods that spanned two growing seasons, elevated atmospheric CO₂ increased soil CO₂ flux most when soil moisture was low and soils were warm. Effects on soil CO₂ flux due to fertilization with N and decreases in diversity were greatest at the times of the year when soils were warm, although there were no significant correlations between these effects and soil moisture. Of the treatments, only the N and diversity treatments were correlated over time; neither were correlated with the CO₂ effect. Models of soil CO₂ flux will need to incorporate ecosystem CO₂ and N availability, as well as ecosystem plant diversity, and incorporate different environmental factors when determining the magnitude of the CO₂, N and diversity effects on soil CO₂ flux.     

Simulated chronic NO3− deposition reduces soil respiration in northern hardwood forests

Chronic N additions to forest ecosystems can enhance soil N availability, potentially leading to reduced C allocation to root systems. This in turn could decrease soil CO₂ efflux. We measured soil respiration during the first, fifth, sixth and eighth years of simulated atmospheric NO₃^? deposition (3 g N m^?2 yr^?1) to four sugar maple‐dominated northern hardwood forests in Michigan to assess these possibilities. During the first year, soil respiration rates were slightly, but not significantly, higher in the NO₃^?‐amended plots. In all subsequent measurement years, soil respiration rates from NO₃^?‐amended soils were significantly depressed. Soil temperature and soil matric potential were measured concurrently with soil respiration and used to develop regression relationships for predicting soil respiration rates. Estimates of growing season and annual soil CO₂ efflux made using these relationships indicate that these C fluxes were depressed by 15% in the eighth year of chronic NO₃^? additions. The decrease in soil respiration was not due to reduced C allocation to roots, as root respiration rates, root biomass, and root turnover were not significantly affected by N additions. Aboveground litter also was unchanged by the 8 years of treatment. Of the remaining potential causes for the decline in soil CO₂ efflux, reduced microbial respiration appears to be the most likely possibility. Documented reductions in microbial biomass and the activities of extracellular enzymes used for litter degradation on the NO₃^?‐amended plots are consistent with this explanation.     

Estimating soil carbon sequestration under elevated CO2 by combining carbon isotope labelling with soil carbon cycle modelling

Elevated CO₂ concentrations generally stimulate grassland productivity, but herbaceous plants have only a limited capacity to sequester extra carbon (C) in biomass. However, increased primary productivity under elevated CO₂ could result in increased transfer of C into soils where it could be stored for prolonged periods and exercise a negative feedback on the rise in atmospheric CO₂. Measuring soil C sequestration directly is notoriously difficult for a number of methodological reasons. Here, we present a method that combines C isotope labelling with soil C cycle modelling to partition net soil sequestration into changes in new C fixed over the experimental duration (C_new) and pre‐experimental C (C_old). This partitioning is advantageous because the C_new accumulates whereas C_old is lost in the course of time (ΔC_new>0 whereas ΔC_old<0). We applied this method to calcareous grassland exposed to 600 μL CO₂ L^?1 for 6 years. The CO₂ used for atmospheric enrichment was depleted in ¹³C relative to the background atmosphere, and this distinct isotopic signature was used to quantify net soil C_new fluxes under elevated CO₂. Using ¹³C/¹²C mass balance and inverse modelling, the Rothamsted model ‘RothC’ predicted gross soil C_new inputs under elevated CO₂ and the decomposition of C_old. The modelled soil C pools and fluxes were in good agreement with experimental data. C isotope data indicated a net sequestration of ≈90 g C_new m^?2 yr^?1 in elevated CO₂. Accounting for C_old‐losses, this figure was reduced to ≈30 g C m^?2 yr^?1 at elevated CO₂; the elevated CO₂‐effect on net C sequestration was in the range of≈10 g C m^?2 yr^?1. A sensitivity and error analysis suggests that the modelled data are relatively robust. However, elevated CO₂‐specific mechanisms may necessitate a separate parameterization at ambient and elevated CO₂; these include increased soil moisture due to reduced leaf conductance, soil disaggregation as a consequence of increased soil moisture, and priming effects. These effects could accelerate decomposition of C_old in elevated CO₂ so that the CO₂ enrichment effect may be zero or even negative. Overall, our findings suggest that the C sequestration potential of this grassland under elevated CO₂ is rather limited.     

CO2 flux from soil in pastures and forests in southwestern Amazonia

Stocks of carbon in Amazonian forest biomass and soils have received considerable research attention because of their potential as sources and sinks of atmospheric CO₂. Fluxes of CO₂ from soil to the atmosphere, on the other hand, have not been addressed comprehensively in regard to temporal and spatial variations and to land cover change, and have been measured directly only in a few locations in Amazonia. Considerable variation exists across the Amazon Basin in soil properties, climate, and management practices in forests and cattle pastures that might affect soil CO₂ fluxes. Here we report soil CO₂ fluxes from an area of rapid deforestation in the southwestern Amazonian state of Acre. Specifically we addressed (1) the seasonal variation of soil CO₂ fluxes, soil moisture, and soil temperature; (2) the effects of land cover (pastures, mature, and secondary forests) on these fluxes; (3) annual estimates of soil respiration; and (4) the relative contributions of grass‐derived and forest‐derived C as indicated by δ¹³CO₂. Fluxes were greatest during the wet season and declined during the dry season in all land covers. Soil respiration was significantly correlated with soil water‐filled pore space but not correlated with temperature. Annual fluxes were higher in pastures compared with mature and secondary forests, and some of the pastures also had higher soil C stocks. The δ¹³C of CO₂ respired in pasture soils showed that high respiration rates in pastures were derived almost entirely from grass root respiration and decomposition of grass residues. These results indicate that the pastures are very productive and that the larger flux of C cycling through pasture soils compared with forest soils is probably due to greater allocation of C belowground. Secondary forests had soil respiration rates similar to mature forests, and there was no correlation between soil respiration and either forest age or forest biomass. Hence, belowground allocation of C does not appear to be directly related to the stature of vegetation in this region. Variation in seasonal and annual rates of soil respiration of these forests and pastures is more indicative of flux of C through the soil rather than major net changes in ecosystem C stocks.     

Immobilization, stabilization and remobilization of nitrogen in forest soils at elevated CO2: a 15N and 13C tracer study

The fate of immobilized N in soils is one of the great uncertainties in predicting C sequestration at increased CO₂ and N deposition. In a dual isotope tracer experiment (¹³C, ¹⁵N) within a 4‐year CO₂ enrichment (+200 ppm_v) study with forest model ecosystems, we (i) quantified the effects of elevated CO₂ on the partitioning of N; (ii) traced immobilized N into physically separated pools of soil organic matter (SOM) with turnover rates known from their ¹³C signals; and (iii) estimated the remobilization and thus, the bio‐availability of newly sequestered C and N. (1) CO₂ enrichment significantly decreased NO₃^? concentrations in soil waters and export from 1.5 m deep lysimeters by 30–80%. Consequently, elevated CO₂ increased the overall retention of N in the model ecosystems. (2) About 60–80% of added ¹⁵NH₄¹⁵NO₃ were retained in soils. The clay fraction was the greatest sink for the immobilized ¹⁵N sequestering 50–60% of the total new soil N. SOM associated with clay contained only 25% of the total new soil C pool and had small C/N ratios (<13), indicating that it consists of humified organic matter with a relatively slow turn over rate. This implies that added ¹⁵N was mainly immobilized in stable mineral‐bound SOM pools. (3) Incubation of soils for 1 year showed that the remobilization of newly sequestered N was three to nine times smaller than that of newly sequestered C. Thus, inorganic inputs of N were stabilized more effectively in soils than C. Significantly less newly sequestered N was remobilized from soils previously exposed to elevated CO₂. In summary, our results show firstly that a large fraction of inorganic N inputs becomes effectively immobilized in relative stable SOM pools and secondly that elevated CO₂ can increase N retention in soils and hence it may tighten N cycling and diminish the risk of nitrate leaching to groundwater.     

Total soil C and N sequestration in a grassland following 10 years of free air CO2 enrichment

Soil C sequestration may mitigate rising levels of atmospheric CO_2. However, it has yet to be determined whether net soil C sequestration occurs in N‐rich grasslands exposed to long‐term elevated CO₂. This study examined whether N‐fertilized grasslands exposed to elevated CO₂ sequestered additional C. For 10 years, Lolium perenne, Trifolium repens, and the mixture of L. perenne/T. repens grasslands were exposed to ambient and elevated CO₂ concentrations (35 and 60 Pa pCO₂). The applied CO₂ was depleted in δ¹³C and the grasslands received low (140 kg ha^?1) and high (560 kg ha^?1) rates of ¹⁵N‐labeled fertilizer. Annually collected soil samples from the top 10 cm of the grassland soils allowed us to follow the sequestration of new C in the surface soil layer. For the first time, we were able to collect dual‐labeled soil samples to a depth of 75 cm after 10 years of elevated CO₂ and determine the total amount of new soil C and N sequestered in the whole soil profile. Elevated CO₂, N‐fertilization rate, and species had no significant effect on total soil C. On average 9.4 Mg new C ha^?1 was sequestered, which corresponds to 26.5% of the total C. The mean residence time of the C present in the 0–10 cm soil depth was calculated at 4.6±1.5 and 3.1±1.1 years for L. perenne and T. repens soil, respectively. After 10 years, total soil N and C in the 0–75 cm soil depth was unaffected by CO₂ concentration, N‐fertilization rate and plant species. The total amount of ¹⁵N‐fertilizer sequestered in the 0–75 cm soil depth was also unaffected by CO₂ concentration, but significantly more ¹⁵N was sequestered in the L. perenne compared with the T. repens swards: 620 vs. 452 kg ha^?1 at the high rate and 234 vs. 133 kg ha^?1 at the low rate of N fertilization. Intermediate values of ¹⁵N recovery were found in the mixture. The fertilizer derived N amounted to 2.8% of total N for the low rate and increased to 8.6% for the high rate of N application. On average, 13.9% of the applied ¹⁵N‐fertilizer was recovered in the 0–75 cm soil depth in soil organic matter in the L. perenne sward, whereas 8.8% was recovered under the T. repens swards, indicating that the N₂‐fixing T. repens system was less effective in sequestering applied N than the non‐N₂‐fixing L. perenne system. Prolonged elevated CO₂ did not lead to an increase in whole soil profile C and N in these fertilized pastures. The potential use of fertilized and regular cut pastures as a net soil C sink under long‐term elevated CO₂ appears to be limited and will likely not significantly contribute to the mitigation of anthropogenic C emissions.     

Long-term CO2 production from deeply weathered soils of a tropical rain forest: evidence for a potential positive feedback to climate warming

Currently, it is unknown what role tropical forest soils will play in the future global carbon cycle under higher temperatures. Many tropical forests grow on deeply weathered soils and although it is generally accepted that soil carbon decomposition increases with higher temperatures, it is not known whether subsurface carbon pools are particularly responsive to increasing soil temperatures. Carbon dioxide (CO₂) diffusing out of soils is an important flux in the global carbon. Although soil CO₂ efflux has been the subject of many studies in recent years, it remains difficult to deduct controls of this flux because of the different sources that produce CO₂ and because potential environmental controls like soil temperature and soil moisture often covary. Here, we report results of a 5‐year study in which we measured soil CO₂ production on two deeply weathered soil types at different depths in an old‐growth tropical wet forest in Costa Rica. Three sites were developed on old river terraces (old alluvium) and the other three were developed on old lava flows (residual). Annual soil CO₂ efflux varied between 2.8–3.6 μmol CO₂‐C m^?2 s^?1 (old alluvium) and 3.4–3.9 μmol CO₂‐C m^?2 s^?1 (residual). More than 75% of the CO₂ was produced in the upper 0.5 m (including litter layer) and less than 7% originated from the soil below 1 m depth. This low contribution was explained by the lack of water stress in this tropical wet forest which has resulted in very low root biomass below 2 m depth. In the top 0.5 m CO₂ production was positively correlated with both temperature and soil moisture; between 0.6 and 2 m depth CO₂ production correlated negatively with soil moisture in one soil and positively with photosynthetically active radiation in the other soil type. Below 2 m soil CO₂ production strongly increased with increasing temperature. In combination with reduced tree growth that has been shown for this ecosystem, this would be a strong positive feedback to ecosystem warming.     

Six years of in situ CO2 enrichment evoke changes in soil structure and soil biota of nutrient-poor grassland

Nutrient‐poor grassland on a silty clay loam overlying calcareous debris was exposed to elevated CO₂ for six growing seasons. The CO₂ exchange and productivity were persistently increased throughout the experiment, suggesting increases in soil C inputs. At the same time, elevated CO₂ lead to increased soil moisture due to reduced evapotransporation. Measurements related to soil microflora did not indicate increased soil C fluxes under elevated CO₂. Microbial biomass, soil basal respiration, and the metabolic quotient for CO₂ (qCO₂) were not altered significantly. PLFA analysis indicated no significant shift in the ratio of fungi to bacteria. 0.5 m KCl extractable organic C and N, indicators of changed DOC and DON concentrations, also remained unaltered. Microbial grazer populations (protozoa, bacterivorous and fungivorous nematodes, acari and collembola) and root feeding nematodes were not affected by elevated CO₂. However, total nematode numbers averaged slightly lower under elevated CO₂ (?16%, ns) and nematode mass was significantly reduced (?43%, P = 0.06). This reduction reflected a reduction in large‐diameter nematodes classified as omnivorous and predacious. Elevated CO₂ resulted in a shift towards smaller aggregate sizes at both micro‐ and macro‐aggregate scales; this was caused by higher soil moisture under elevated CO₂. Reduced aggregate sizes result in reduced pore neck diameters. Locomotion of large‐diameter nematodes depends on the presence of large enough pores; the reduction in aggregate sizes under elevated CO₂ may therefore account for the decrease in large nematodes. These animals are relatively high up the soil food web; this decline could therefore trigger top‐down effects on the soil food web. The CO₂ enrichment also affected the nitrogen cycle. The N stocks in living plants and surface litter increased at elevated CO₂, but N in soil organic matter and microbes remained unaltered. Nitrogen mineralization increased markedly, but microbial N did not differ between CO₂ treatments, indicating that net N immobilization rates were unaltered. In summary, this study did not provide evidence that soils and soil microbial communities are affected by increased soil C inputs under elevated CO₂. On the contrary, available data (¹³C tracer data, minirhizotron observations, root ingrowth cores) suggests that soil C inputs did not increase substantially. However, we provide first evidence that elevated CO₂ can reduce soil aggregation at the scale from µ m to mm scale, and that this can affect soil microfaunal populations.     

Net grassland carbon flux over a subambient to superambient CO2 gradient

Increasing atmospheric CO₂ concentrations may have a profound effect on the structure and function of plant communities. A previously grazed, central Texas grassland was exposed to a 200‐µmol mol^?1 to 550 µmol mol^?1 CO₂ gradient from March to mid‐December in 1998 and 1999 using two, 60‐m long, polyethylene‐ covered chambers built directly onto the site. One chamber was operated at subambient CO₂ concentrations (200–360 µmol mol^?1 daytime) and the other was regulated at superambient concentrations (360–550 µmol mol^?1). Continuous CO₂ gradients were maintained in each chamber by photosynthesis during the day and respiration at night. Net ecosystem CO₂ flux and end‐of‐year biomass were measured in each of 10, 5‐m long sections in each chamber. Net CO₂ fluxes were maximal in late May (c. day 150) in 1998 and in late August in 1999 (c. day 240). In both years, fluxes were near zero and similar in both chambers at the beginning and end of the growing season. Average daily CO₂ flux in 1998 was 13 g CO₂ m^?2 day^?1 in the subambient chamber and 20 g CO₂ m^?2 day^?1 in the superambient chamber; comparable averages were 15 and 26 g CO₂ m^?2 day^?1 in 1999. Flux was positively and linearly correlated with end‐of‐year above‐ground biomass but flux was not linearly correlated with CO₂ concentration; a finding likely to be explained by inherent differences in vegetation. Because C₃ plants were the dominant functional group, we adjusted average daily flux in each section by dividing the flux by the average percentage C₃ cover. Adjusted fluxes were better correlated with CO₂ concentration, although scatter remained. Our results indicate that after accounting for vegetation differences, CO₂ flux increased linearly with CO₂ concentration. This trend was more evident at subambient than superambient CO₂ concentrations.     

Interactions between plant growth and soil nutrient cycling under elevated CO2: a meta-analysis

free air carbon dioxide enrichment (FACE) and open top chamber (OTC) studies are valuable tools for evaluating the impact of elevated atmospheric CO₂ on nutrient cycling in terrestrial ecosystems. Using meta‐analytic techniques, we summarized the results of 117 studies on plant biomass production, soil organic matter dynamics and biological N₂ fixation in FACE and OTC experiments. The objective of the analysis was to determine whether elevated CO₂ alters nutrient cycling between plants and soil and if so, what the implications are for soil carbon (C) sequestration. Elevated CO₂ stimulated gross N immobilization by 22%, whereas gross and net N mineralization rates remained unaffected. In addition, the soil C : N ratio and microbial N contents increased under elevated CO₂ by 3.8% and 5.8%, respectively. Microbial C contents and soil respiration increased by 7.1% and 17.7%, respectively. Despite the stimulation of microbial activity, soil C input still caused soil C contents to increase by 1.2% yr^?1. Namely, elevated CO₂ stimulated overall above‐ and belowground plant biomass by 21.5% and 28.3%, respectively, thereby outweighing the increase in CO₂ respiration. In addition, when comparing experiments under both low and high N availability, soil C contents (+2.2% yr^?1) and above‐ and belowground plant growth (+20.1% and+33.7%) only increased under elevated CO₂ in experiments receiving the high N treatments. Under low N availability, above‐ and belowground plant growth increased by only 8.8% and 14.6%, and soil C contents did not increase. Nitrogen fixation was stimulated by elevated CO₂ only when additional nutrients were supplied. These results suggest that the main driver of soil C sequestration is soil C input through plant growth, which is strongly controlled by nutrient availability. In unfertilized ecosystems, microbial N immobilization enhances acclimation of plant growth to elevated CO₂ in the long‐term. Therefore, increased soil C input and soil C sequestration under elevated CO₂ can only be sustained in the long‐term when additional nutrients are supplied.     

Linking microbial activity and soil organic matter transformations in forest soils under elevated CO2

Soil organic matter (SOM) dynamics ultimately govern the ability of soil to provide long‐term C sequestration and the nutrients required for ecosystem productivity. Predicting belowground responses to elevated CO₂ requires an integrated understanding of SOM transformations and the microbial activity that governs them. It remains unclear how the microorganisms upon which these transformations depend will function in an elevated CO₂ world. This study examines SOM transformations and microbial metabolism in soils from the Duke Free Air Carbon Enrichment site in North Carolina, USA. We assessed microbial respiration and net nitrogen (N) mineralization in soils with and without elevated CO₂ exposure during a 100‐day incubation. We also traced the depleted C isotopic signature of the supplemental CO₂ into SOM and the soils' phospholipid fatty acids (PLFA), which serve as biomarkers for living cells. Cumulative net N mineralization in elevated CO₂ soils was 50% that in control soils after a 100‐day incubation. Respiration was not altered with elevated CO₂. C : N ratios of bulk SOM did not change with elevated CO₂, but incubation data suggest that the C : N ratios of mineralized organic matter increased with elevated CO₂. Values of SOM δ¹³C were depleted with elevated CO₂ (?26.7±0.2 vs. ?30.2±0.3‰), reflecting the depleted signature of the supplemental CO₂. We compared δ¹³C of individual PLFA with the δ¹³C of SOM to discern incorporation of the depleted C isotopic signature into soil microbial groups in elevated CO₂ plots. PLFA i15:0, a15:0, and 10Met18:0 reflected significant incorporation of recently produced photosynthate, suggesting that the bacterial groups defined by these biomarkers are active metabolizers in elevated CO₂ soils. At least one of these groups (actinomycetes, 10Met18:0) specializes in metabolizing less labile substrates. Because control plots did not receive an equivalent ¹³C tracer, we cannot determine from these data whether this group of organisms was stimulated by elevated CO₂ compared with these organisms in control soils. Stimulation of this group, if it occurred in the elevated CO₂ plot, would be consistent with a decline in the availability of mineralizable organic matter with elevated CO₂, which incubation data suggest may be the case in these soils.     

Interactions of tropospheric CO2 and O3 enrichments and moisture variations on microbial biomass and respiration in soil

Soil microbial biomass C (C_mic) is a sensitive indicator of trends in organic matter dynamics in terrestrial ecosystems. This study was conducted to determine the effects of tropospheric CO₂ or O₃ enrichments and moisture variations on total soil organic C (C_org), mineralizable C fraction (C_Min), C_mic, maintenance respiratory (qCO₂) or C_mic death (qD) quotients, and their relationship with basal respiration (BR) rates and field respiration (FR) fluxes in wheat‐soybean agroecosystems. Wheat (Triticum aestivum L.) and soybean (Glycine max. L. Merr) plants were grown to maturity in 3‐m dia open‐top field chambers and exposed to charcoal‐filtered (CF) air at 350 μL CO₂ L^?1; CF air + 150 μL CO₂ L^?1; nonfiltered (NF) air + 35 nL O₃ L^?1; and NF air + 35 nL O₃ L^?1 + 150 μL CO₂ L^?1 at optimum (? 0.05 MPa) and restricted soil moisture (? 1.0 ± 0.05 MPa) regimes. The + 150 μL CO₂ L^?1 additions were 18 h d^?1 and the + 35 nL O₃ L^?1 treatments were 7 h d^?1 from April until late October. While C_org did not vary consistently, C_Min, C_mic and C_mic fractions increased in soils under tropospheric CO₂ enrichment (500 μL CO₂ L^?1) and decreased under high O₃ exposures (55 ± 6 nL O₃ L^?1 for wheat; 60 ± 5 nL O₃ L^?1 for soybean) compared to the CF treatments (25 ± 5 nL O₃ L^?1). The qCO₂ or qD quotients of C_mic were also significantly decreased in soils under high CO₂ but increased under high O₃ exposures compared to the CF control. The BR rates did not vary consistently but they were higher in well‐watered soils. The FR fluxes were lower under high O₃ exposures compared to soils under the CF control. An increase in C_mic or C_mic fractions and decrease in qCO₂ or qD observed under high CO₂ treatment suggest that these soils were acting as C sinks whereas, reductions in C_mic or C_mic fractions and increase in qCO₂ or qD in soils under elevated tropospheric O₃ exposures suggest the soils were serving as a source of CO₂.     

Changes in soil carbon and nitrogen properties and microbial communities in relation to growth of Pinus radiata and Nothofagus fusca trees after 6 years at ambient and elevated atmospheric CO2

Increasing atmospheric CO₂ concentration can influence the growth and chemical composition of many plant species, and thereby affect soil organic matter pools and nutrient fluxes. Here, we examine the effects of ambient (initially 362 μL L^?1) and elevated (654 μL L^?1) CO₂ in open‐top chambers on the growth after 6 years of two temperate evergreen forest species: an exotic, Pinus radiata D. Don, and a native, Nothofagus fusca (Hook. F.) Oerst. (red beech). We also examine associated effects on selected carbon (C) and nitrogen (N) properties in litter and mineral soil, and on microbial properties in rhizosphere and hyphosphere soil. The soil was a weakly developed sand that had a low initial C concentration of about 1.0 g kg^?1 at both 0–100 and 100–300 mm depths; in the N. fusca system, it was initially overlaid with about 50 mm of forest floor litter (predominantly FH material) taken from a Nothofagus forest. A slow‐release fertilizer was added during the early stages of plant growth; subsequent foliage analyses indicated that N was not limiting. After 6 years, stem diameters, foliage N concentrations and C/N ratios of both species were indistinguishable (P>0.10) in the two CO₂ treatments. Although total C contents in mineral soil at 0–100 mm depth had increased significantly (P<0.001) after 6 years growth of P. radiata, averaging 80±0.20 g m^?2 yr^?1, they were not significantly influenced by elevated CO₂. However, CO₂‐C production in litter, and CO₂‐C production, microbial C, and microbial C/N ratios in mineral soil (0–100 mm depth) under P. radiata were significantly higher under elevated than ambient CO₂. CO₂‐C production, microbial C, and numbers of bacteria (but not fungi) were also significantly higher under elevated CO₂ in hyphosphere soil, but not in rhizosphere soil. Under N. fusca, some incorporation of the overlaid litter into the mineral soil had probably occurred; except for CO₂‐C production and microbial C in hyphosphere soil, none of the biochemical properties or microbial counts increased significantly under elevated CO₂. Net mineral‐N production, and generally the potential utilization of different substrates by microbial communities, were not significantly influenced by elevated CO₂ under either tree species. Physiological profiles of the microbial communities did, however, differ significantly between rhizosphere and hyphosphere samples and between samples under P. radiata and N. fusca. Overall, results support the concept that a major effect on soil properties after prolonged exposure of trees to elevated CO₂ is an increase in the amounts, and mineralization rate, of labile organic components.     

Indirect effects of soil moisture reverse soil C sequestration responses of a spring wheat agroecosystem to elevated CO2

Increased plant productivity under elevated atmospheric CO₂ concentrations might increase soil carbon (C) inputs and storage, which would constitute an important negative feedback on the ongoing atmospheric CO₂ rise. However, elevated CO₂ often also leads to increased soil moisture, which could accelerate the decomposition of soil organic matter, thus counteracting the positive effects via C cycling. We investigated soil C sequestration responses to 5 years of elevated CO₂ treatment in a temperate spring wheat agroecosystem. The application of ¹³C‐depleted CO₂ to the elevated CO₂ plots enabled us to partition soil C into recently fixed C (C_new) and pre‐experimental C (C_old) by ¹³C/¹²C mass balance. Gross C inputs to soils associated with C_new accumulation and the decomposition of C_old were then simulated using the Rothamsted C model ‘RothC.’ We also ran simulations with a modified RothC version that was driven directly by measured soil moisture and temperature data instead of the original water balance equation that required potential evaporation and precipitation as input. The model accurately reproduced the measured C_new in bulk soil and microbial biomass C. Assuming equal soil moisture in both ambient and elevated CO₂, simulation results indicated that elevated CO₂ soils accumulated an extra ～40–50 g C m^?2 relative to ambient CO₂ soils over the 5 year treatment period. However, when accounting for the increased soil moisture under elevated CO₂ that we observed, a faster decomposition of C_old resulted; this extra C loss under elevated CO₂ resulted in a negative net effect on total soil C of ～30 g C m^?2 relative to ambient conditions. The present study therefore demonstrates that positive effects of elevated CO₂ on soil C due to extra soil C inputs can be more than compensated by negative effects of elevated CO₂ via the hydrological cycle.     

Precipitation pulses and soil CO2 flux in a Sonoran Desert ecosystem

Precipitation is a major driver of biological processes in arid and semiarid ecosystems. Soil biogeochemical processes in these water‐limited systems are closely linked to episodic rainfall events, and the relationship between microbial activity and the amount and timing of rainfall has implications for the whole‐system carbon (C) balance. Here, the influences of storm size and time between events on pulses of soil respiration were explored in an upper Sonoran Desert ecosystem. Immediately following experimental rewetting in the field, CO₂ efflux increased up to 30‐fold, but generally returned to background levels within 48 h. CO₂ production integrated over 48 h ranged from 2.5 to 19.3 g C m⁻² and was greater beneath shrubs than in interplant spaces. When water was applied on sequential days, postwetting losses of CO₂ were only half a large as initial fluxes, and the size of the second pulse increased with time between consecutive events. Soil respiration was more closely linked to the organic matter content of surface soils than to rainfall amount. Beneath shrubs, rates increased nonlinearly with storm size, reaching an asymptote at approximately 0.5 cm simulated storms. This nonlinear relationship stems from (1) resource limitation of microbial activity that is manifest at small time scales, and (2) greatly reduced process rates in deeper soil strata. Thus, beyond some threshold in storm size, increasing the duration or depth of soil moisture has little consequence for short‐term losses of CO₂. In addition, laboratory rewetting across a broad range in soil water content suggest that microbial activity and CO₂ efflux following rainfall may be further modified by the routing and redistribution of water along hillslopes. Finally, analysis of long‐term precipitation data suggests that half the monsoon storms in this system are sufficient to induce soil heterotrophic activity and C losses, but are not large enough to elicit autotrophic activity and C accrual by desert shrubs.     

Carbon and nitrogen pools and mineralization in a grassland gley soil under elevated carbon dioxide at a natural CO2 spring

The growth and chemical composition of most plants are influenced by elevated CO₂, but accompanying effects on soil organic matter pools and mineralization are less clearly defined, partly because of the short‐term nature of most studies. Herein we describe soil properties from a naturally occurring cold CO₂ spring (Hakanoa) in Northland, New Zealand, at which the surrounding vegetation has been exposed to elevated CO₂ for at least several decades. The mean annual temperature at this site is ≈ 15.5 °C and rainfall ≈ 1550 mm. The site was unfertilized and ungrazed, with a vegetation of mainly C3 and C4 grasses, and had moderate levels of ‘available’ P. Two soils were present ? a gley soil and an organic soil – but only the gley soil is examined here. Average atmospheric CO₂ concentrations at 17 sampling locations in the gley soil area ranged from 372 to 670 ppmv. In samples at 0–5 cm depth, pH averaged 5.4; average values for organic C were 150 g, total N 11 g, microbial C 3.50 g, and microbial N 0.65 g kg^?1, respectively. Under standardized moisture conditions at 25 °C, average rates of CO₂‐C production (7–14 days) were 5.4 mg kg^?1 h^?1 and of net mineral‐N production (14 ?42 days) 0.40 mg kg^?1 h^?1. These properties were all correlated positively and significantly (P < 0.10) with atmospheric CO₂ concentrations, but not with soil moisture (except for CO₂‐C production) or with clay content; they were, however, correlated negatively and mainly significantly with soil pH. In spite of uncertainties associated with the uncontrolled environment of naturally occurring springs, we conclude that storage of C and N can increase under prolonged exposure to elevated CO₂, and may include an appreciable labile fraction in mineral soil with an adequate nutrient supply.     

CO2 exchange in three Canadian High Arctic ecosystems: response to long-term experimental warming

Carbon dioxide exchange, soil C and N, leaf mineral nutrition and leaf carbon isotope discrimination (LCID‐Δ) were measured in three High Arctic tundra ecosystems over 2 years under ambient and long‐term (9 years) warmed (～2°C) conditions. These ecosystems are located at Alexandra Fiord (79°N) on Ellesmere Island, Nunavut, and span a soil water gradient; dry, mesic, and wet tundra. Growing season CO₂ fluxes (i.e., net ecosystem exchange (NEE), gross ecosystem photosynthesis (GEP), and ecosystem respiration (R_e)) were measured using an infrared gas analyzer and winter C losses were estimated by chemical absorption. All three tundra ecosystems lost CO₂ to the atmosphere during the winter, ranging from 7 to 12 g CO₂‐C m^?2 season^?1 being highest in the wet tundra. The period during the growing season when mesic tundra switch from being a CO₂ source to a CO₂ sink was increased by 2 weeks because of warming and increases in GEP. Warming during the summer stimulated dry tundra GEP more than R_e and thus, NEE was consistently greater under warmed as opposed to ambient temperatures. In mesic tundra, warming stimulated GEP with no effect on R_e increasing NEE by ～10%, especially in the first half of the summer. During the ～70 days growing season (mid‐June–mid‐August), the dry and wet tundra ecosystems were net CO₂‐C sinks (30 and 67 g C m^?2 season^?1, respectively) and the mesic ecosystem was a net C source (58 g C m^?2 season^?1) to the atmosphere under ambient temperature conditions, due in part to unusual glacier melt water flooding that occurred in the mesic tundra. Experimental warming during the growing season increased net C uptake by ～12% in dry tundra, but reduced net C uptake by ～20% in wet tundra primarily because of greater rates of R_e as opposed to lower rates of GEP. Mesic tundra responded to long‐term warming with ～30% increase in GEP with almost no change in R_e reducing this tundra type to a slight C source (17 g C m^?2 season^?1). Warming caused LCID of Dryas integrafolia plants to be higher in dry tundra and lower in Salix arctic plants in mesic and wet tundra. Our findings indicate that: (1) High Arctic ecosystems, which occur in similar mesoclimates, have different net CO₂ exchange rates with the atmosphere; (2) long‐term warming can increase the net CO₂ exchange of High Arctic tundra by stimulating GEP, but it can also reduce net CO₂ exchange in some tundra types during the summer by stimulating R_e to a greater degree than stimulating GEP; (3) after 9 years of experimental warming, increases in soil carbon and nitrogen are detectable, in part, because of increases in deciduous shrub cover, biomass, and leaf litter inputs; (4) dry tundra increases in GEP, in response to long‐term warming, is reflected in D. integrifolia LCID; and (5) the differential carbon exchange responses of dry, mesic, and wet tundra to similar warming magnitudes appear to depend, in part, on the hydrologic (soil water) conditions. Annual net ecosystem CO₂‐C exchange rates ranged from losses of 64 g C m^?2 yr^?1 to gains of 55 g C m^?2 yr^?1. These magnitudes of positive NEE are close to the estimates of NPP for these tundra types in Alexandra Fiord and in other High Arctic locations based on destructive harvests.     

Soil CO2 efflux in a tropical forest in the central Amazon

This study investigated the spatial and temporal variation in soil carbon dioxide (CO₂) efflux and its relationship with soil temperature, soil moisture and rainfall in a forest near Manaus, Amazonas, Brazil. The mean rate of efflux was 6.45±0.25 SE μmol CO₂ m^?2s^?1 at 25.6±0.22 SE°C (5 cm depth) ranging from 4.35 to 9.76 μmol CO₂ m^?2s^?1; diel changes in efflux were correlated with soil temperature (r²=0.60). However, the efflux response to the diel cycle in temperature was not always a clear exponential function. During period of low soil water content, temperature in deeper layers had a better relationship with CO₂ efflux than with the temperature nearer the soil surface. Soil water content may limit CO₂ production during the drying‐down period that appeared to be an important factor controlling the efflux rate (r²=0.39). On the other hand, during the rewetting period microbial activity may be the main controlling factor, which may quickly induce very high rates of efflux. The CO₂ flux chamber was adapted to mimic the effects of rainfall on soil CO₂ efflux and the results showed that efflux rates reduced 30% immediately after a rainfall event. Measurements of the CO₂ concentration gradient in the soil profile showed a buildup in the concentration of CO₂ after rain on the top soil. This higher CO₂ concentration developed shortly after rainfall when the soil pores in the upper layers were filled with water, which created a barrier for gas exchange between the soil and the atmosphere.