Interannual and interseasonal soil CO2 efflux and VOC exchange rates in a Mediterranean holm oak forest in response to experimental drought

Climate models predict drier conditions in the next decades in the Mediterranean basin. Given the importance of soil CO₂ efflux in the global carbon balance and the important role of soil monoterpene and volatile organic compounds (VOCs) in soil ecology, we aimed to study the effects of the predicted drought on soil CO₂, monoterpenes and other VOC exchange rates and their seasonal and interannual variations. We decreased soil water availability in a Mediterranean holm oak forest soil by means of an experimental drought system performed since 1999 to the present. Measurements of soil gas exchange were carried out with IRGA, GC and PTR-MS techniques during two annual campaigns of contrasting precipitation. Soil respiration was twice higher the wet year than the dry year (2.27±0.26 and 1.05±0.15, respectively), and varied seasonally from 3.76±0.85 μmol m⁻² s⁻¹ in spring, to 0.13±0.01 μmol m⁻² s⁻¹ in summer. These results highlight the strong interannual and interseasonal variation in CO₂ efflux in Mediterranean ecosystems. The drought treatment produced a significant soil respiration reduction in drought plots in the wet sampling period. This reduction was even higher in wet springs (43% average reduction). These results show (1) that soil moisture is the main factor driving seasonal and interannual variations in soil respiration and (2) that the response of soil respiration to increased temperature is constrained by soil moisture. The results also show an additional control of soil CO₂ efflux by physiology and phenology of trees and animals. Soil monoterpene exchange rates ranged from −0.01 to 0.004 nmol m⁻² s⁻¹, thus the contribution of this Mediterranean holm oak forest soil to the total monoterpenes atmospheric budget seems to be very low. Responses of individual monoterpenes and VOCs to the drought treatment were different depending on the compound. This suggests that the effect of soil moisture reduction in the monoterpenes and VOC exchange rates seems to be dependent on monoterpene and VOC type. In general, soil monoterpene and other VOC exchange rates were not correlated with soil CO₂ efflux. In all cases, only a low proportion of variance was explained by the soil moisture changes, since almost all VOCs increased their emission rates in summer 2005, probably due to the effect of high soil temperature. Results indicate thus that physical and biological processes in soil are controlling soil VOC exchange but further research is needed on how these factors interact to produce the observed VOCs exchange responses.     

Grazing intensity alters soil respiration in an alpine meadow on the Tibetan plateau

Grazing intensity may alter the soil respiration rate in grassland ecosystems. The objectives of our study were to (1) determine the influence of grazing intensity on temporal variations in soil respiration of an alpine meadow on the northeastern Tibetan Plateau; and (2) characterise the temperature response of soil respiration under different grazing intensities. Diurnal and seasonal soil respiration rates were measured for two alpine meadow sites with different grazing intensities. The light grazing (LG) meadow site had a grazing intensity of 2.55 sheep ha⁻¹, while the grazing intensity of the heavy grazing (HG) meadow site, 5.35 sheep ha⁻¹, was approximately twice that of the LG site. Soil respiration measurements showed that CO₂ efflux was almost twice as great at the LG site as at the HG site during the growing season, but the diurnal and seasonal patterns of soil respiration rate were similar for the two sites. Both exhibited the highest annual soil respiration rate in mid-August and the lowest in January. Soil respiration rate was highly dependent on soil temperature. The Q₁₀ value for annual soil respiration was lower for the HG site (2.75) than for the LG site (3.22). Estimates of net ecosystem CO₂ exchange from monthly measurements of biomass and soil respiration revealed that during the period from May 1998 to April 1999, the LG site released 2040 g CO₂ m⁻² y⁻¹ to the atmosphere, which was about one third more than the 1530 g CO₂ m⁻² y⁻¹ released at the HG site. The results suggest that (1) grazing intensity alters not only soil respiration rate, but also the temperature dependence of soil CO₂ efflux; and (2) soil temperature is the major environmental factor controlling the temporal variation of soil respiration rate in the alpine meadow ecosystem.     

Subtropical plantations are large carbon sinks: Evidence from two monoculture plantations in South China

Quantifying the net carbon (C) storage of forest plantations is required to assess their potential to offset fossil fuel emissions. In this study, a biometric approach was used to estimate net ecosystem productivity (NEP) for two monoculture plantations in South China: Acacia crassicarpa and Eucalyptus urophylla. This approach was based on stand-level net primary productivity (NPP, based on direct biometric inventory) and heterotrophic respiration (R_h). In comparisons of R_h determination based on trenching vs. tree girdling, both trenching and tree girdling changed soil temperature and soil moisture relative to undisturbed control plots, and we assess the effects of corrections for disturbances of soil moisture and soil moisture on the estimation of soil CO₂ efflux partitioning. Soil microbial biomass and dissolved organic carbon were significantly lower in trenched plots than in tree girdled plots for both plantations. Annual soil CO₂ flux in trenched plots (R_h-t) was significantly lower than in tree-girdled plots (R_h-g) in both plantations. The estimates of R_h-t and R_h-g, expressed as a percentage of total soil respiration, were 58 ± 4% and 74 ± 6%, respectively, for A. crassicarpa, and 64 ± 3% and 78 ± 5%, respectively, for E. urophylla. By the end of experiment, the difference in soil CO₂ efflux between the trenched plots and tree-girdled plots had become small for both plantations. Annual R_h (mean of the annual R_h-t and R_h-g) and net primary production (NPP) were 470 ± 25 and 800 ± 118 g C m⁻² yr⁻¹, respectively, for A. crassicarpa, and 420 ± 35 and 2380 ± 187 g C m⁻² yr⁻², respectively, for E. urophylla. The two plantations in the developmental stage were large carbon sinks: NEP was 330 ± 76 C m⁻² yr⁻¹ for A. crassicarpa and 1960 ± 178 g C m⁻² yr⁻¹ for E. urophylla.     

Explaining temporal variation in soil CO2 efflux in a mature spruce forest in Southern Germany

An open dynamic chamber system was used to measure the soil CO₂ efflux intensively and continuously throughout a growing season in a mature spruce forest (Picea abies) in Southern Germany. The resulting data set contained a large amount of temporally highly resolved information on the variation in soil CO₂ efflux together with environmental variables. Based on this background, the dependencies of the soil CO₂ efflux rate on the controlling environmental factors were analysed in-depth. Of the abiotic factors, soil temperature alone explained 72% of the variation in the efflux rate, and including soil water content (SWC) as an additional variable increased the explained variance to about 83%. Between April and December, average rates ranged from 0.43 to 5.15 μmol CO₂ m⁻² s⁻¹ (in November and July, respectively) with diurnal variations of up to 50% throughout the experiment. The variability in wind speed above the forest floor influenced the CO₂ efflux rates for measuring locations with a litter layer of relatively low bulk density (and hence relatively high proportions of pore spaces). For the temporal integration of flux rates for time scales of hours to days, however, wind velocities were of no effect, reflecting the fact that wind forcing acts on the transport, but not the production of CO₂ in the soil. The variation in both the magnitude of the basal respiration rate and the temperature sensitivity throughout the growing season was only moderate (coefficient of variation of 15 and 25%, respectively). Soil water limitation of the CO₂ production in the soil could be best explained by a reduction in the temperature-insensitive basal respiration rate, with no discernible effect on the temperature sensitivity. Using a soil CO₂ efflux model with soil temperature and SWC as driving variables, it was possible to calculate the annual soil CO₂ efflux for four consecutive years for which meteorological data were available. These simulations indicate an average efflux sum of 560 g C m⁻² yr⁻¹ (SE=22 g C m⁻² yr⁻¹). An alternative model derived from the same data but using temperature alone as a driver over-estimated the annual flux sum by about 7% and showed less inter-annual variability. Given a likely shift in precipitation patterns alongside temperature changes under projected global change scenarios, these results demonstrate the necessity to include soil moisture in models that calculate the evolution of CO₂ from temperate forest soils.     

Winter soil CO2 efflux and its contribution to annual soil respiration in different ecosystems of a forest-steppe ecotone, north China

Most soil respiration measurements are conducted during the growing season. In tundra and boreal forest ecosystems, cumulative winter soil CO₂ fluxes are reported to be a significant component of their annual carbon budgets. However, little information on winter soil CO₂ efflux is known from mid-latitude ecosystems. Therefore, comparing measurements of soil respiration taken annually versus during the growing season will improve the accuracy of ecosystem carbon budgets and the response of soil CO₂ efflux to climate changes. In this study we measured winter soil CO₂ efflux and its contribution to annual soil respiration for seven ecosystems (three forests: Pinus sylvestris var. mongolica plantation, Larix principis-rupprechtii plantation and Betula platyphylla forest; two shrubs: Rosa bella and Malus baccata; and two meadow grasslands) in a forest-steppe ecotone, north China. Overall mean winter and growing season soil CO₂ effluxes were 0.15-0.26 μmol m⁻² s⁻¹ and 2.65-4.61 μmol m⁻² s⁻¹, respectively, with significant differences in the growing season among the different ecosystems. Annual Q₁₀ (increased soil respiration rate per 10 °C increase in temperature) was generally higher than the growing season Q₁₀. Soil water content accounted for 84% of the variations in growing season Q₁₀ and soil temperature range explained 88% of the variation in annual Q₁₀. Soil organic carbon density to 30 cm depth was a good surrogate for SR₁₀ (basal soil respiration at a reference temperature of 10 °C). Annual soil CO₂ efflux ranged from 394.76 g C m⁻² to 973.18 g C m⁻² using observed ecosystem-specific response equations between soil respiration and soil temperature. Estimates ranged from 424.90 g C m⁻² to 784.73 g C m⁻² by interpolating measured soil respiration between sampling dates for every day of the year and then computing the sum to obtain the annual value. The contributions of winter soil CO₂ efflux to annual soil respiration were 3.48-7.30% and 4.92-7.83% using interpolated and modeled methods, respectively. Our results indicate that in mid-latitude ecosystems, soil CO₂ efflux continues throughout the winter and winter soil respiration is an important component of annual CO₂ efflux.     

Long-term effects of seasonal and diurnal temperature fluctuations on carbon dioxide efflux from a forest soil

Temperature fluctuations are a fundamental entity of the soil environment in the temperate zone and show fast (diurnal) and slow (seasonal) dynamics. Responses of soil respiration to temperature fluctuations were investigated in a root-free soil of a mid-European beech-oak forest. First, in laboratory we analysed the efflux of CO₂ from soil microcosms exposed to seasonal (±5 °C of the annual mean) and diurnal fluctuations (±5 °C of the seasonal levels) in a two-factorial design. Second, in field microcosms we investigated effects of smoothing diurnal temperature fluctuations in soil (simulating a possible global trend) on CO₂ efflux. Third, the natural temperature regime was simulated in laboratory microcosms and their CO₂ efflux was compared to the one in the field. The experiments lasted for 1 year to differentiate seasonal and annual responses.Dynamics of CO₂ efflux, microbial basal respiration, biomass and qO₂ varied with seasonal temperature regime. However, in the laboratory the annual cumulative CO₂-C production did not differ between treatments and varied between 10.9% and 11.7% of the total microcosm C, disregarding seasonal and/or diurnal fluctuations. The similarity of cumulative C production suggests that the availability of microbially mobilisable carbon pools rather than the temperature regime limited soil respiration. Diurnal fluctuations generally did not affect CO₂ efflux and microbial activity, though winter Q₁₀ values were increased in their absence. Simulation of the natural temperature regime in the laboratory resulted in CO₂ efflux similar to field microcosms. In the field, rates of CO₂ efflux and microbial activity, seasonal and annual cumulative CO₂-C production were significantly higher at smoothed than at natural temperature conditions (annually 13.1% and 11.0% of total C was respired, respectively). Facing global climate changes the mechanisms regulating responses of soil respiration to temperature fluctuations need further investigation.     

Short-term effects of clearfelling on soil CO2, CH4, and N2O fluxes in a Sitka spruce plantation

We examined the effects of forest clearfelling on the fluxes of soil CO₂, CH₄, and N₂O in a Sitka spruce (Picea sitchensis (Bong.) Carr.) plantation on an organic-rich peaty gley soil, in Northern England. Soil CO₂, CH₄, N₂O as well as environmental factors such as soil temperature, soil water content, and depth to the water table were recorded in two mature stands for one growing season, at the end of which one of the two stands was felled and one was left as control. Monitoring of the same parameters continued thereafter for a second growing season. For the first 10 months after clearfelling, there was a significant decrease in soil CO₂ efflux, with an average efflux rate of 4.0 g m⁻² d⁻¹ in the mature stand (40-year) and 2.7 g m⁻² d⁻¹ in clearfelled site (CF). Clearfelling turned the soil from a sink (−0.37 mg m⁻² d⁻¹) for CH₄ to a net source (2.01 mg m⁻² d⁻¹). For the same period, soil N₂O fluxes averaged 0.57 mg m⁻² d⁻¹ in the CF and 0.23 mg m⁻² d⁻¹ in the 40-year stand. Clearfelling affected environmental factors and lead to higher daily soil temperatures during the summer period, while it caused an increase in the soil water content and a rise in the water table depth. Despite clearfelling, CO₂ remained the dominant greenhouse gas in terms of its greenhouse warming potential.     

Carbon exchange in a freshwater marsh in the Sanjiang Plain, northeastern China

Northern wetlands are critically important to global change because of their role in modulating atmospheric concentrations of greenhouse gases, especially CO₂ and CH₄. At present, continuous observations for CO₂ and CH₄ fluxes from northern wetlands in Asia are still very limited. In this paper, two growing season measurements for CO₂ flux by eddy covariance technique and CH₄ flux by static chamber technique were conducted in 2004 and 2005, at a permanently inundated marsh in the Sanjiang Plain, northeastern China. The seasonal variations of CO₂ exchange and CH₄ flux and the environmental controls on them were investigated. During the growing seasons, large variations in net ecosystem CO₂ exchange (NEE) and gross ecosystem productivity (GEP) were observed with the range of −4.0 to 2.2 (where negative exchange is a gain of carbon from the atmosphere) and 0-7.6 g C m⁻² d⁻¹, respectively. Ecosystem respiration (RE) displayed relatively smooth seasonal pattern with the range of 0.8-4.2 g C m⁻² d⁻¹. More than 70% of the total GEP was consumed by respiration, which resulted in a net CO₂ uptake of 143 ± 9.8 and 100 ± 9.2 g C m⁻² for the marsh over the growing seasons of 2004 and 2005, respectively. A significant portion of the accumulated NEE-C was lost by CH₄ emission during the growing seasons, indicating the great potential of CH₄ emission from the inundated marsh. Air temperature and leaf area index jointly affected the seasonal variation of GEP and the seasonal dynamic of RE was mainly controlled by soil temperature and leaf area index. Soil temperature also exerted the dominant influence over variation of CH₄ flux while no significant relationship was found between CH₄ emission and water table level. The close relationships between carbon fluxes and temperature can provide insights into the response of marsh carbon exchange to a changing climate. Future long term flux measurements over the freshwater marsh ecosystems are undoubtedly necessary.     

Uncoupling of microbial CO2 production and release in frozen soil and its implications for field studies of arctic C cycling

Knowledge of seasonal trends and controls of soil CO₂ emissions to the atmosphere is important for simulating atmospheric CO₂ concentrations and for understanding and predicting the global carbon cycle. This is particularly the case for high arctic soils subject to extreme fluctuating environmental conditions. Based on field measurements of soil CO₂ efflux, temperature, water content, pore gas composition in soil and frozen cores as well as detailed temperature experiments performed in the laboratory, we evaluated seasonal controls of CO₂ effluxes from a well-drained tundra heath site in NE-Greenland. During the growing season, near-surface temperatures correlated well with observed CO₂ effluxes (r²>0.9). However, during intensive thawing of near-surface layers we observed up to 1.5-fold higher effluxes than expected due to temperature alone. These high rates were consistent with high CO₂ concentrations in frozen soil (>10% CO₂) and suggested a spring burst event during soil thawing and a corresponding trapping of produced CO₂ during winter. Laboratory experiments revealed that microbial soil respiration continued down to a least −18 °C and that up to 80% of the produced CO₂ was trapped in soil at temperatures between 0 and −9 °C. The trapping of CO₂ in frozen soil was positively correlated with soil moisture (r²=0.85) and led to an abrupt change of the temperature sensitivity (Q₁₀) observed for soil CO₂ release at 0 °C with Q₁₀ values below 0 °C being up to 100-fold higher than above 0 °C. The results of sub-zero CO₂ production allowed us to predict the microbial soil respiration throughout the year and to evaluate to what extent burst events during thawing can be explained by the release of CO₂ being produced and trapped during winter. Taking only the upper 20 cm of the soil into account, winter soil respiration accounted for about 40% of the annual soil respiration. At least 14% of the winter CO₂ production was trapped during the winter 2000-2001 and observed to be released upon thawing. Thus, the site-specific winter soil respiration is an important part of the annual C cycle and CO₂ trapping should be accounted for in future field and modelling studies of soil respiration dynamics in arctic ecosystems. In conclusion, we have discovered a soil moisture dependent uncoupling of CO₂ production and release in frozen soils with important implications for future field studies of Arctic C cycling.     

Influence of warming and enchytraeid activities on soil CO2 and CH4 fluxes

To determine the sum of ‘direct’ and ‘indirect’ effects of climatic change on enchytraeid activity and C fluxes from an organic soil we assessed the influence of temperature (4, 10 and 15 °C incubations) on enchytraeid populations and soil CO₂ and CH₄ fluxes over 116 days. Moisture was maintained at 60% of soil dry weight during the experimental period and measurements of enchytraeid biomass and numbers, and CO₂ and CH₄ fluxes were made after 3, 16, 33, 44, 65, 86 and 116 days. Enchytraeid population numbers and biomass increased in all temperature treatments with the greatest increase produced at 15 °C (to over threefold initial values by day 86). Results also showed that enchytraeid activity increased CO₂ fluxes by 10.7±4.5, 3.4±4.0 and 26.8±2.6% in 4, 10 and 15 °C treatments, respectively, with the greatest CO₂ production observed at 15 °C for the entire 116 day incubation period (P<0.05). The soil respiratory quotient analyses at lower temperatures (i.e. 4-10 °C) gave a Q₁₀ of 1.7 and 1.9 with and without enchytraeids, respectively. At temperatures above 10 °C (i.e. 10-15 °C) Q₁₀ significantly increased (P<0.01) and was 25% greater in the presence of enchytraeids (Q₁₀=3.4) than without (Q₁₀=2.6). In contrast to CO₂ production, no significant relationships were observed between net CH₄ fluxes and temperature and only time showed a significant effect on CH₄ production (P<0.01).Total soil CO₂ production was positively linked with enchytraeid biomass and mean soil CO₂-C production was 77.01±6.05 CO₂-C μg mg enchytraeid tissue⁻¹ day⁻¹ irrespective of temperature treatment. This positive relationship was used to build a two step regression model to estimate the effects of temperature on enchytraeid biomass and soil CO₂ respiration in the field. Predictions of potential CO₂ production were made using enchytraeid biomass data obtained in the field from two upland grassland sites (Sourhope and Great Dun Fell at the Moor House Nature Reserve, both in the UK). The findings of this work suggest that a 5 °C increase in atmospheric temperature above mean ambient temperature could have the potential to produce a significant increase in enchytraeid biomass resulting in a near twofold increase in soil CO₂ release from both soil types. The interaction between temperature and soil biology will clearly be an important determinant of soil respiration responses to global warming.     

Plant biomass influences rhizosphere priming effects on soil organic matter decomposition in two differently managed soils

We used a continuous labeling method of naturally ¹³C-depleted CO₂ in a growth chamber to test for rhizosphere effects on soil organic matter (SOM) decomposition. Two C3 plant species, soybean (Glycine max) and sunflower (Helianthus annus), were grown in two previously differently managed soils, an organically farmed soil and a soil from an annual grassland. We maintained a constant atmospheric CO₂ concentration at 400±5 ppm and δ¹³C signature at −24.4‰ by regulating the flow of naturally ¹³C-depleted CO₂ and CO₂-free air into the growth chamber, which allowed us to separate new plant-derived CO₂-C from original soil-derived CO₂-C in soil respiration. Rhizosphere priming effects on SOM decomposition, i.e., differences in soil-derived CO₂-C between planted and non-planted treatments, were significantly different between the two soils, but not between the two plant species. Soil-derived CO₂-C efflux in the organically farmed soil increased up to 61% compared to the no-plant control, while the annual grassland soil showed a negligible increase (up to 5% increase), despite an overall larger efflux of soil-derived CO₂-C and total soil C content. Differences in rhizosphere priming effects on SOM decomposition between the two soils could be largely explained by differences in plant biomass, and in particular leaf biomass, explaining 49% and 74% of the variation in primed soil C among soils and plant species, respectively. Nitrogen uptake rates by soybean and sunflower was relatively high compared to soil C respiration and associated N mineralization, while inorganic N pools were significantly depleted in the organic farm soil by the end of the experiment. Despite relatively large increases in SOM decomposition caused by rhizosphere effects in the organic farm soil, the fast-growing soybean and sunflower plants gained little extra N from the increase in SOM decomposition caused by rhizosphere effects. We conclude that rhizosphere priming effects of annual plants on SOM decomposition are largely driven by plant biomass, especially in soils of high fertility that can sustain high plant productivity.     

Temperature dependence of soil CO2 efflux is strongly modulated by seasonal patterns of moisture availability in a Mediterranean ecosystem

Extensive research has focused on the temperature sensitivity of soil respiration. However, in Mediterranean ecosystems, soil respiration may have a pulsed response to precipitation events, especially during prolonged dry periods. Here, we investigate temporal variations in soil respiration (R_s), soil temperature (T) and soil water content (SWC) under three different land uses (a forest area, an abandoned agricultural field and a rainfed olive grove) in a dry Mediterranean area of southeast Spain, and evaluate the relative importance of soil temperature and water content as predictors of R_s. We hypothesize that soil moisture content, rather than soil temperature, becomes the major factor controlling CO₂ efflux rates in this Mediterranean ecosystem during the summer dry season. Soil CO₂ efflux was measured monthly between January 2006 and December 2007 using a portable soil respiration instrument fitted with a soil respiration chamber (LI-6400-09). Mean annual soil respiration rates were 2.06 ± 0.07, 1.71 ± 0.09, and 1.12 ± 0.12 μmol m⁻² s⁻¹ in the forest, abandoned field and olive grove, respectively. R_s was largely controlled by soil temperature above a soil water content threshold value of 10% at 0-15 cm depth for forest and olive grove, and 15% for abandoned field. However, below those thresholds R_s was controlled by soil moisture. Exponential and linear models adequately described R_s responses to environmental variables during the growing and dry seasons. Models combining abiotic (soil temperature and soil rewetting index) and biotic factors (above-ground biomass index and/or distance from the nearest tree) explained between 39 and 73% of the temporal variability of R_s in the forest and olive grove. However, in the abandoned field, a single variable - either soil temperature (growing season) or rewetting index (dry season) - was sufficient to explain between 51 and 63% of the soil CO₂ efflux. The fact that the rewetting index, rather than soil water content, became the major factor controlling soil CO₂ efflux rates during the prolonged summer drought emphasizes the need to quantify the effects of rain pulses in estimates of net annual carbon fluxes from soil in Mediterranean ecosystems.     

Increased N deposition retards mineralization of old soil organic matter

The aim of this study was to investigate the effects of increased N deposition on new and old pools of soil organic matter (SOM). We made use of a 4-yr experiment, where spruce and beech growing on an acidic loam and a calcareous sand were exposed to increased N deposition (7 vs. 70 kg N ha⁻¹ yr⁻¹) and to elevated atmospheric CO₂. The added CO₂ was depleted in ¹³C, which enabled us to distinguish between old and new C in SOM-pools fractionated into particle sizes. Elevated N deposition for 4 yr increased significantly the contents of total SOM in 0-10 cm depth of the acidic loam (+9%), but not in the calcareous sand. Down to 25 cm soil depth, C storage in the acidic loam was between 100 and 300 g C m⁻² larger under high than under low N additions. However, this increase was small as compared with the SOM losses of 600-700 g C g C 0.25 m⁻¹ m⁻² from the calcareous sand resulting from the disturbance of soils during setting up of the experiment. The amounts of new, less than 4 yr old SOM in the sand fractions of both soils were greater under high N deposition, showing that C inputs from trees into soils increased. Root biomass in the acidic loam was larger under N additions (+25%). Contents of old, more than 4 yr old C in the clay and silt fractions of both soils were significantly greater under high than under low N deposition. Since clay- and silt-bound SOM consists of humified compounds, this indicates that N additions retarded mineralization of old and humified SOM. The retardation of C mineralization in the clay and silt fraction accounted for 60-80 g C m⁻² 4 yr⁻¹, which corresponds to about 40% of the old SOM mineralized in these fraction. As a consequence, preservation of old and humified SOM under elevated N deposition might be a process that could lead to an increased soil C storage in the long-term.     

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.     

Annual soil CO2 effluxes in the High Arctic: The role of snow thickness and vegetation type

Little work has been done to quantify annual soil CO₂ effluxes in the High Arctic region because of the difficulty in taking winter measurements. Since the effects of climate change are expected to be higher in Arctic than in temperate ecosystems, it is important that summer measurements are extended to cover the entire year. This study evaluates the quantity and quality of soil organic C (SOC) and seasonal controls of soil CO₂ effluxes in three soils under three dominating types of vegetation (Dryas, Cassiope, and Salix) at Svalbard. Measurements included soil CO₂ effluxes in the field and the laboratory, temperature, water content, and snow thickness. About 90% of the variation in soil respiration throughout 1 year was due to near-surface soil temperatures which ranged from −12 to +12 °C. Total annual soil CO₂ effluxes varied from 103 g C m⁻² at soils under Cassiope, 152 g C m⁻² under Dryas sites, and 176 g C m⁻² under Salix, with 20%, 14%, and 30%, respectively, being released during a 6-month winter period. The sensitivity of soil respiration with respect to soil temperature was the same year round and differences in winter CO₂ effluxes at the three vegetation types were mainly related to subsurface soil temperatures controlled by snow depth. The quantity and quality of soil organic matter varied under the different vegetation types. Soils under Salix had the largest and most labile pool of SOC and were characterized by a long period of snow cover. In contrast, soils under Cassiope were more nutrient-poor, more acidic and held the smallest amount of total and labile SOC, whereas soils under Dryas remained snow-free most of the winter and therefore had the coldest winter conditions. Thus, winter soil respiration rates under Dryas and Cassiope were significantly lower than those under Salix; under Dryas this was mainly due to snow depth, under Cassiope this was a combination of snow depth and poor litter quality. It is concluded that winter respiration is highly variable across Arctic landscapes and depends on the spatial distribution of snow, which acts as a direct control on soil temperatures and indirect on vegetation types and thereby, the amount and quality of soil organic matter, which serve as additional important drivers of soil respiration.     

Microbial activity in soils frozen to below −39 °C

Recent research on life in extreme environments has shown that some microorganisms metabolize at extremely low temperatures in Arctic and Antarctic ice and permafrost. Here, we present kinetic data on CO₂ and ¹⁴CO₂ release from intact and ¹⁴C-glucose amended tundra soils (Barrow, Alaska) incubated for up to a year at 0 to −39°C. The rate of CO₂ production declined exponentially with temperature but it remained positive and measurable, e.g. 2-7 ng CO₂-C cm⁻³ soil d⁻¹, at −39 °C. The variation of CO₂ release rate (v) was adequately explained by the double exponential dependence on temperature (T) and unfrozen water content (W) (r²>0.98): v=A exp(λT+kW) and where A, λ and k are constants. The rate of ¹⁴CO₂ release from added glucose declined more steeply with cooling as compared with the release of total CO₂, indicating that (a) there could be some abiotic component in the measured flux of CO₂ or (b) endogenous respiration is more cold-resistant than substrate-induced respiration. The respiration activity was completely eliminated by soil sterilization (1 h, 121 °C), stimulated by the addition of oxidizable substrate (glucose, yeast extract), and reduced by the addition of acetate, which inhibits microbial processes in acidic soils (pH 3-5). The tundra soil from Barrow displayed higher below-zero activity than boreal soils from West Siberia and Sweden. The permafrost soils (20-30 cm) were more active than the samples from seasonally frozen topsoil (0-10 cm, Barrow). Finding measurable respiration to −39 °C is significant for determining, understanding, and predicting current and future CO₂ emission to the atmosphere and for understanding the low temperature limits of microbial activity on the Earth and on other planets.     

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.     

Soil CO2 flux in six monospecific forest plantations in Southern Rwanda

Forest soils contain the largest carbon stock of all terrestrial biomes and are probably the most important source of carbon dioxide (CO₂) to atmosphere. Soil CO₂ fluxes from 54 to 72-year-old monospecific stands in Rwanda were quantified from March 2006 to December 2007. The influences of soil temperature, soil water content, soil carbon (C) and nitrogen (N) stocks, soil pH, and stand characteristics on soil CO₂ flux were investigated. The mean annual soil CO₂ flux was highest under Eucalyptus saligna (3.92 μmol m⁻² s⁻¹) and lowest under Entandrophragma excelsum (3.13 μmol m⁻² s⁻¹). The seasonal variation in soil CO₂ flux from all stands followed the same trend and was highest in rainy seasons and lowest in dry seasons. Soil CO₂ flux was mainly correlated to soil water content (R² = 0.36-0.77), stand age (R² = 0.45), soil C stock (R² = 0.33), basal area (R² = 0.21), and soil temperature (R² = 0.06-0.17). The results contribute to the understanding of factors that influence soil CO₂ flux in monocultural plantations grown under the same microclimatic and soil conditions. The results can be used to construct models that predict soil CO₂ emissions in the tropics.     

Strong temperature dependence and no moss photosynthesis in winter CO2 flux for a Kobresia meadow on the Qinghai-Tibetan plateau

We examined the CO₂ exchange of a Kobresia meadow ecosystem on the Qinghai-Tibetan plateau using a chamber system. CO₂ efflux from the ecosystem was strongly dependence on soil surface temperature. The CO₂ efflux-temperature relationship was identical under both light and dark conditions, indicating that no photosynthesis could be detected under light conditions during the measurement period. The temperature sensitivity (Q₁₀) of the CO₂ efflux showed a marked transition around −1.0 °C; Q₁₀ was 2.14 at soil surface temperatures above and equal to −1.0 °C but was 15.3 at temperatures below −1.0 °C. Our findings suggest that soil surface temperature was the major factor controlling winter CO₂ flux for the alpine meadow ecosystem and that freeze-thaw cycles at the soil surface layer play an important role in the temperature dependence of winter CO₂ flux.     

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.