Microbial physiology and soil CO2 efflux after 9 years of soil warming in a temperate forest – no indications for thermal adaptations

Thermal adaptations of soil microorganisms could mitigate or facilitate global warming effects on soil organic matter (SOM) decomposition and soil CO₂ efflux. We incubated soil from warmed and control subplots of a forest soil warming experiment to assess whether 9 years of soil warming affected the rates and the temperature sensitivity of the soil CO₂ efflux, extracellular enzyme activities, microbial efficiency, and gross N mineralization. Mineral soil (0–10 cm depth) was incubated at temperatures ranging from 3 to 23 °C. No adaptations to long‐term warming were observed regarding the heterotrophic soil CO₂ efflux (R₁₀ warmed: 2.31 ± 0.15 μmol m^?2 s^?1, control: 2.34 ± 0.29 μmol m^?2 s^?1; Q₁₀ warmed: 2.45 ± 0.06, control: 2.45 ± 0.04). Potential enzyme activities increased with incubation temperature, but the temperature sensitivity of the enzymes did not differ between the warmed and the control soils. The ratio of C : N acquiring enzyme activities was significantly higher in the warmed soil. Microbial biomass‐specific respiration rates increased with incubation temperature, but the rates and the temperature sensitivity (Q₁₀ warmed: 2.54 ± 0.23, control 2.75 ± 0.17) did not differ between warmed and control soils. Microbial substrate use efficiency (SUE) declined with increasing incubation temperature in both, warmed and control, soils. SUE and its temperature sensitivity (Q₁₀ warmed: 0.84 ± 0.03, control: 0.88 ± 0.01) did not differ between warmed and control soils either. Gross N mineralization was invariant to incubation temperature and was not affected by long‐term soil warming. Our results indicate that thermal adaptations of the microbial decomposer community are unlikely to occur in C‐rich calcareous temperate forest soils.     

Winter CO2 flux from soil and snow surfaces in a cool-temperate deciduous forest, Japan

We measured diurnal and wintertime changes in CO₂ fluxes from soil and snow surfaces in a Japanese cool-temperate Quercus/Betula forest between December 1994 and May 1995. To evaluate the relationship between these winter fluxes and temperature, flux measurements were made with the open-flow infrared gas analyzer (IRGA) method rather than with the more commonly used closed chamber method or the snow CO₂ profile method. The open-flow IRGA method proved to be more successful in measurements of winter CO₂ fluxes than the two standard methods. Despite colder air temperatures, soil temperature profiles were greater than 0°C because of the thermal insulation effect of deep snowpack. This reveals that soil temperature is satisfactory for microbial respiration throughout the winter. Unfrozen soils under the snowpack showed neither diurnal nor wintertime trends in CO₂ fluxes or in soil surface temperature, although there was a daily snow surface CO₂ flux of 0.18–0.32 g m^–2. By combining this with other reference data, Japanese cool-temperate forest soils in snowy regions can be estimated to emit < 100 g m^–2 carbon over an entire winter, and this value accounts for < 15% of the annual emission. In the present study, when data for all winter fluxes were taken together, fluxes were most highly correlated with deep soil temperatures rather than the soil surface temperature. Such a high correlation can be attributed to the relatively increased respiration of the deep soil where the temperature was higher than the soil surface temperature. Thus, deeper soil temperature is a better predictor of winter CO₂ fluxes in cold and snowy ecosystems.     

中国森林生态系统土壤CO2释放分布规律及其影响因素

联合国气候框架公约的签署提升了人们对全球变暖、碳循环变化的关注。陆地生态系统在全球变暖格局下的地位与作用,尤其是土壤碳库对全球变暖格局的响应是全球变化研究的焦点。土壤CO2释放作为土壤-大气CO2交换的主要途径之一,也就成为各国生态学家研究的重点内容。在对我国森林生态系统CO2释放通量以及相关气候、生物等因子的资料进行收集、整理和分析的基础上,探讨了我国森林生态系统土壤CO2释放的分布规律,以及这种规律性分布的气候、生物影响因素。对于我国这样一个南北跨度大的国家,不同区域的森林生态系统土壤CO2释放通量间存在较大的差异,在全国尺度上,森林生态系统土壤CO2释放通量平均值为（1．79±0．86）gCm^-2d^-1,而且土壤CO2释放通量随着纬度增加逐渐降低。作为一个复杂的生态过程,土壤CO2释放受到生物、非生物因子或独立、或综合的影响。通过分析指出,在全国尺度上,年均温、降雨量、群落净生产力及凋落物量显著地影响森林土壤CO2释放通量。同时,也正是这些影响因子的纬度分布,导致了我国森林生态系统土壤CO2释放通量的纬度分布规律。作为衡量土壤CO2释放对温度敏感性的重要指标,计算了我国森林生态系统土壤CO2释放温度敏感性系数-Q10值,约为1．5,该值显著低于全球平均水平,2．0。     

CO2 efflux from a Mediterranean semi-arid forest soil. II. Effects of soil fauna and surface stoniness

Many forest soils in the Mediterranean basin areshallow and contain high amounts of gravel in theorganic layers. Recent studies on soil organic matteraccumulation have shown high amounts of organic matteroccurring mainly in soils with high levels ofstoniness at the soil surface. The gravel layer mayaffect the microclimatic conditions of the soilsurface and probably the distribution and activity ofsoil fauna.In order to quantify the combined effects soil fauna(epigeic macrofauna and earthworms) and stoniness onthe release of soil CO₂, we performed a threefactor field experiment by using a series ofreconstructed soil profiles. Factors 1 and 2 consistedof the exclusion/presence of soil epigeic macrofaunaand earthworms, and factor 3 of the presence/absenceof a gravel layer intermingled with the H horizon. Weincubated ¹⁴C straw in the H horizon and carriedout three 40 mm rainfall simulations.Soil respiration primarily depended on the season. Theeffects of soil fauna were generally small and did notcoincide with periods of high faunal activity. Thelargest effects of both earthworms and soil epigeicfauna were found after wetting the soil in summer. Theeffects of the earthworms were concentrated in themineral soil while the effects of the epigeic faunawere concentrated in the H horizon and mainly arosetowards the end of the experiment. This suggests thatthe effects of epigeic fauna may have beenunderestimated due to the length of the experiment.The gravel layer increased the effect of faunaprobably by creating more favorable microclimaticconditions. The accumulation of organic matter insoils with high levels of stoniness cannot beexplained by the effect of gravel on soil microclimatenor by its effect on the activity of soil fauna.     

Temperature-independent diel variation in soil respiration observed from a temperate deciduous forest

The response of soil respiration (R_s) to temperature depends largely on the temporal and spatial scales of interest and how other environmental factors interact with this response. They are often represented by empirical exponential equations in many ecosystem analyses because of the difficulties in separating covarying environmental responses and in observing below ground processes. The objective of this study was to quantify a soil temperature‐independent component in R_s by examining the diel variation of an R_s time series measured in a temperate deciduous forest located at Oak Ridge, TN, USA between March and December 2003. By fitting 2 hourly, continuous automatic chamber measurements of CO₂ efflux at the soil surface to a Q₁₀ function to obtain the temperature‐dependent respiration (R_t) and plotting the diel cycles of R_t, R_s, and their difference (R_i), we found that an obvious temperature‐independent component exists in R_s during the growing season. The diel cycle of this component has a distinct day/night pattern and agrees well with diel variations in photosynthetically active radiation (PAR) and air temperature. Elevated canopy CO₂ concentration resulted in similar patterns in the diel cycle of the temperature‐independent component but with different daily average rates in different stages of growing season. We speculate that photosynthesis of the stand is one of the main contributors to this temperature‐independent respiration component although more experiments are needed to draw a firm conclusion. We also found that despite its relatively small magnitude compared with the temperature‐dependent component, the diel variation in the temperature‐independent component can lead to significantly different estimates of the temperature sensitivity of soil respiration in the study forest. As a result, the common practice of using fitted temperature‐dependent function from night‐time measurements to extrapolate soil respiration during the daytime may underestimate daytime soil respiration.     

Changes in seasonal precipitation distribution but not annual amount affect litter decomposition in a secondary tropical forest

In the tropics of South China, climate change induced more rainfall events in the wet season in the last decades. Moreover, there will be more frequently spring drought in the future. However, knowledge on how litter decomposition rate would respond to these seasonal precipitation changes is still limited. In the present study, we conducted a precipitation manipulation experiment in a tropical forest. First, we applied a 60% rainfall exclusion in April and May to defer the onset of wet season and added the same amount of water in October and November to mimic a deferred wet season (DW); second, we increased as much as 25% mean annual precipitation into plots in July and August to simulate a wetter wet season (WW). Five single‐species litters, with their carbon to nitrogen ratio ranged from 27 to 49, and a mixed litter were used to explore how the precipitation change treatments would affect litter decomposition rate. The interaction between precipitation changes and litter species was not significant. The DW treatment marginally accelerated litter decomposition across six litter types. Detailed analysis showed that DW increased litter decomposition rate in the periods of January to March and October to December, when soil moisture was increased by the water addition in the dry season. In contrast, WW did not significantly affect litter decomposition rate, which was consistent with the unchanged soil moisture pattern. In conclusion, the study indicated that regardless of litter types or litter quality, the projected deferred wet season would increase litter decomposition rate, whereas the wetter wet season would not affect litter decomposition rate in the tropical forests. This study improves our knowledge of how tropical forest carbon cycling in response to precipitation change.     

森林生态系统土壤CO2释放随海拔梯度的变化及其影响因子

联合国气候框架公约的签署提升了人们对全球变暖、碳循环的关注。土壤CO2释放作为土壤-大气CO2交换的主要途径之一,成为了各国生态学家研究的重点内容。通过对1800～2155m海拔梯度上森林生态系统土壤CO2释放进行研究,揭示了较小空间尺度上土壤CO2释放的变化规律及其控制机制。在研究区域内,随着海拔梯度的增加,森林土壤CO2释放由(1.94±0.06)μmolm-2s-1逐渐增加至(2.22±0.07)μmolm-2s-1。土壤温度、土壤水分、土壤有机碳(SOC)、全N、全P与土壤CO2释放呈显著正相关(n=14,P<0.05);土壤容重与土壤CO2释放速率呈显著负相关(n=14,P<0.05);土壤pH对土壤CO2释放影响不显著。作为一个复杂的生态学过程,环境因子及其交互作用对土壤CO2释放产生影响,为了减少因子共线性影响,逐步降低因子维数,采用主成分分析(PCA)揭示了土壤温度、土壤水分、SOC、全N、全P、容重6个因子的联合作用,其累积贡献率达到了57%以上;进一步运用逐步回归分析方法,探讨了影响土壤CO2释放沿海拔梯度分布的主导因子,结果表明土壤水分是研究区域森林生态系统土壤CO2释放沿海拔梯度变化的主导因子。     

Separating the effects of moisture and temperature on soil CO2 efflux in a coniferous forest in the Sierra Nevada mountains

Separating the effects of soil temperature and moisture on soil CO₂ efflux is critical to modeling and understanding the belowground carbon dynamics of forest ecosystems. We developed two analytical procedures to separate the effects of soil temperature and moisture, based on continuous measurements of the CO₂ efflux, temperature and moisture of the soil at a ponderosa pine plantation in the Sierra Nevada Mountains in California, from May 1998 to August 1999. We found that the combined effects of temperature and moisture on the seasonal variation of soil CO₂ efflux could be effectively separated and represented with the product of a temperature term and a moisture term. The relationship between soil CO₂ efflux and temperature could be well described using a power function. This relationship was modified by soil moisture which affects only the coefficient, but not the exponent, of the power function. We also found that when soil moisture was held constant, the temperature effect explained 82% of the temporal variation in CO₂ efflux of the soil. Similarly, when temperature was held constant, the moisture effects explained 84% of the variation. Temperature and moisture together explained 89% of the total temporal variations in soil CO₂ efflux. A multiplicative formulation with power functions representing both temperature and moisture dependences was recommended for modeling soil CO₂ efflux. This formulation can be used to model the seasonal trend of soil CO₂ efflux of the forest based on temperature and moisture, two key variables influenced by climate change and management practices.     

Soil CO2 efflux in a boreal pine forest under atmospheric CO2 enrichment and air warming

The response of forest soil CO₂ efflux to the elevation of two climatic factors, the atmospheric concentration of CO₂ (↑CO₂ of 700 μmol mol⁻¹) and air temperature (↑ T with average annual increase of 5°C), and their combination (↑CO₂+↑ T ) was investigated in a 4-year, full-factorial field experiment consisting of closed chambers built around 20-year-old Scots pines ( Pinus sylvestris L.) in the boreal zone of Finland. Mean soil CO₂ efflux in May–October increased with elevated CO₂ by 23–37%, with elevated temperature by 27–43%, and with the combined treatment by 35–59%. Temperature elevation was a significant factor in the combined 4-year efflux data, whereas the effect of elevated CO₂ was not as evident. Elevated temperature had the most pronounced impact early and late in the season, while the influence of elevated CO₂ alone was especially notable late in the season. Needle area was found to be a significant predictor of soil CO₂ efflux, particularly in August, a month of high root growth, thus supporting the assumption of a close link between whole-tree physiology and soil CO₂ emissions. The decrease in the temperature sensitivity of soil CO₂ efflux observed in the elevated temperature treatments in the second year nevertheless suggests the existence of soil response mechanisms that may be independent of the assimilating component of the forest ecosystem. In conclusion, elevated atmospheric CO₂ and air temperature consistently increased forest soil CO₂ efflux over the 4-year period, their combined effect being additive, with no apparent interaction.     

Effect of soil water stress on soil respiration and its temperature sensitivity in an 18-year-old temperate Douglas-fir stand

We analyzed 17 months (August 2005 to December 2006) of continuous measurements of soil CO₂ efflux or soil respiration (R_S) in an 18‐year‐old west‐coast temperate Douglas‐fir stand that experienced somewhat greater than normal summertime water deficit. For soil water content at the 4 cm depth (θ) > 0.11 m³ m^?3 (corresponding to a soil water matric potential of ?2 MPa), R_S was positively correlated to soil temperature at the 2 cm depth (T_S). Below this value of θ, however, R_S was largely decoupled from T_S, and evapotranspiration, ecosystem respiration and gross primary productivity (GPP) began to decrease, dropping to about half of their maximum values when θ reached 0.07 m³ m^?3. Soil water deficit substantially reduced R_S sensitivity to temperature resulting in a Q₁₀ significantly < 2. The absolute temperature sensitivity of R_S (i.e. dR_S/dT_S) increased with θ up to 0.15 m³ m^?3, above which it slowly declined. The value of dR_S/dT_S was nearly 0 for θ < 0.08 m³ m^?3, thereby confirming that R_S was largely unaffected by temperature under soil water stress conditions. Despite the possible effects of seasonality of photosynthesis, root activity and litterfall on R_S, the observed decrease in its temperature sensitivity at low θ was consistent with the reduction in substrate availability due to a decrease in (a) microbial mobility, and diffusion of substrates and extracellular enzymes, and (b) the fraction of substrate that can react at high T_S, which is associated with low θ. We found that an exponential (van't Hoff type) model with Q₁₀ and R₁₀ dependent on only θ explained 92% of the variance in half‐hourly values of R_S, including the period with soil water stress conditions. We hypothesize that relating Q₁₀ and R₁₀ to θ not only accounted for the effects of T_S on R_S and its temperature sensitivity but also accounted for the seasonality of biotic (photosynthesis, root activity, and litterfall) and abiotic (soil moisture and temperature) controls and their interactions.     

Impact of grazing intensity on seasonal variations in soil organic carbon and soil CO2 efflux in two semiarid grasslands in southern Botswana

Biological soil crusts (BSCs) are an important source of organic carbon, and affect a range of ecosystem functions in arid and semiarid environments. Yet the impact of grazing disturbance on crust properties and soil CO₂ efflux remain poorly studied, particularly in African ecosystems. The effects of burial under wind-blown sand, disaggregation and removal of BSCs on seasonal variations in soil CO₂ efflux, soil organic carbon, chlorophyll a and scytonemin were investigated at two sites in the Kalahari of southern Botswana. Field experiments were employed to isolate CO₂ efflux originating from BSCs in order to estimate the C exchange within the crust. Organic carbon was not evenly distributed through the soil profile but concentrated in the BSC. Soil CO₂ efflux was higher in Kalahari Sand than in calcrete soils, but rates varied significantly with seasonal changes in moisture and temperature. BSCs at both sites were a small net sink of C to the soil. Soil CO₂ efflux was significantly higher in sand soils where the BSC was removed, and on calcrete where the BSC was buried under sand. The BSC removal and burial under sand also significantly reduced chlorophyll a, organic carbon and scytonemin. Disaggregation of the soil crust, however, led to increases in chlorophyll a and organic carbon. The data confirm the importance of BSCs for C cycling in drylands and indicate intensive grazing, which destroys BSCs through trampling and burial, will adversely affect C sequestration and storage. Managed grazing, where soil surfaces are only lightly disturbed, would help maintain a positive carbon balance in African drylands.     

土壤温度和水分对油松林土壤呼吸的影响

用LI-COR 6400-09土壤呼吸测定系统,在太原天龙山自然保护区对油松林的土壤呼吸进行了4a测定.结果表明,土壤呼吸具有明显的季节变化特点,最大值出现在8月份,在6～10 μmol m~(-2) s~(-1) 之间,最小值出现在12月份和3月份,在0.5 μmol m~(-2) s~(-1)左右.2005、2006、2007和2008年土壤呼吸CO_2的年平均值分别为(4.71±3.74)、(3.08±2.91)、(2.96±2.58) μmol m~(-2) s~(-1)和(2.12±1.54) μmol m~(-2) s~(-1);4a的CO_2总平均值为(3.27±2.95) μmol m~(-2) s~(-1).4个测定年土壤呼吸的平均值总体差异显著.4个测定年土壤CO_2释放C量分别为1103.5、882.8、918.4 g m~(-2)和666.3 g m~(-2),总C平均值为892.8 g m~(-2),具有明显的年际差异.指数方程可以很好的表达土壤呼吸与10 cm深度土壤温度的关系,R~2值4a分别为0.39,0.60,0.68和0.71,Q_(10)值分别为3.10,4.41、4.05和5.18,用4a全部数据计算的Q_(10)值为4.31.土壤水分对土壤呼吸的作用较弱,R~2值4a分别仅为0.31、0.25、0.13和0.02,但是夏季土壤干旱对土壤呼吸的抑制作用非常明显,可使土壤呼吸下降50%以上.夏季土壤干旱是导致土壤呼吸年际变化的主要原因.4个包括土壤温度和水分的双变量模型均可以很好地模拟土壤呼吸的季节变化, 拟合方程的R~2值从0.58到0.79.     

Effects of rainfall events on soil CO2 flux in a cool temperate deciduous broad-leaved forest

The effects of rainfall events on soil CO₂ fluxes were examined in a cool temperate Quercus/Betula forest in Japan. The soil CO₂ fluxes were measured using an open-flow gas exchange system with an infrared gas analyzer in the snow-free season from August 1999 to November 2000. Soil CO₂ flux showed no significant diurnal trend on days without rain. In contrast, rainfall events caused a significant increase in soil CO₂ flux. To determine the effect of rainfall events and to evaluate more precisely the daily and annual soil carbon flux, we constructed a multiple polynomial regression model that included two variables, soil temperature and soil water content, using the soil CO₂ flux data recorded on sunny days. Daily soil carbon fluxes on sunny days calculated by the model were almost the same as those determined by the field measurements. On the contrary, the fluxes measured on rainy days were significantly higher than those calculated daily from the soil carbon fluxes by the model. Annual soil carbon fluxes in 1999 and 2000 were estimated using models that both do and do not take rainfall effects into consideration. The result indicates that post-rainfall increases in soil CO₂ flux represent approximately 16–21% of the annual soil carbon flux in this cool temperate deciduous forest.     

Response of soil CO2 efflux to water manipulation in a tallgrass prairie ecosystem

Although CO₂ efflux plays a critical role in carbon exchange between the biosphere and atmosphere, our understanding of its regulation by soil moisture is rather limited. This study was designed to examine the relationship between soil CO₂ efflux and soil moisture in a natural ecosystem by taking advantage of the historically long drought period from 29 July to 21 September 2000 in the southern Central Great Plain, USA. At the end of August when soil moisture content at the top 50 mm was reduced to less than 50 g kg^–1 gravimetrically, we applied 8 levels of water treatments (simulated to rainfall of 0, 10, 25, 50, 100, 150, 200, and 300 mm) with three replicates to 24 plots in a Tallgrass Prairie ecosystem in Central Oklahoma, USA. In order to quantify root-free soil CO₂ efflux, we applied the same 8 levels of water treatments to 24 500-mm soil columns using soil from field adjacent to the experimental plots. We characterized dynamic patterns of soil moisture and soil CO₂ efflux over the experimental period of 21 days. Both soil moisture content and CO₂ efflux showed dramatic increases immediately after the water addition, followed by a gradual decline. The time courses in response to water treatments are well described by Y=Y₀+ate^–bt, where Y is either soil moisture or CO₂ efflux, t is time, Y ₀, a, and b are coefficients. Among the 8 water treatments, the maximal soil CO₂ efflux rate occurred at the 50 mm water level in the field and 100 mm in the root-free soil 1 day after the treatment. The maximal soil CO₂ efflux gradually shifted to higher water levels as the experiment continued. We found the relationship between soil CO₂ efflux and soil moisture using the data from the 21-day experiment was highly scattered, suggesting complex mechanisms determining soil CO₂ efflux by soil moisture.     

Spatial variability in forest soil CO2 efflux assessed with a calibrated soda lime technique

Accurate estimates of soil CO₂ efflux are important in the current discussion on the carbon balance of forests, and can be used to validate models and remotely sensed data. Due to the typically large spatial variability, large sample numbers are required to estimate mean forest soil CO₂ efflux with reasonable confidence intervals. Most infrared gas analysis (IRGA) systems are not well suited to simultaneously produce daily means and handle this spatial variability problem. The soda lime technique gives daily means and allows the required large sample numbers, but is less accurate than the IRGA systems. Using an elaborate cross-calibration, we tried to combine the accuracy of an IRGA method with the spatial integration potential of the soda lime technique. This paper reports on the calibration technique used to improve the accuracy of the soda lime technique and confirms the spatial variability in soil CO₂ efflux in a heterogeneous forest.     

CO2‐caused change in plant species composition rivals the shift in vegetation between mid‐grass and tallgrass prairies

Atmospheric CO₂ enrichment usually changes the relative contributions of plant species to biomass production of grasslands, but the types of species favored and mechanisms by which change is mediated differ among ecosystems. We measured changes in the contributions of C₃ perennial forbs and C₄ grasses to aboveground biomass production of tallgrass prairie assemblages grown along a field CO₂ gradient (250–500 μmol mol^?1) in central Texas USA. Vegetation was grown on three soil types and irrigated each season with water equivalent to the growing season mean of precipitation for the area. We predicted that CO₂ enrichment would increase the forb contribution to community production, and favor tall‐grasses over mid‐grasses by increasing soil water content and reducing the frequency with which soil water fell below a limitation threshold. CO₂ enrichment favored forbs over grasses on only one of three soil types, a Mollisol. The grass fraction of production increased dramatically across the CO₂ gradient on all soils. Contribution of the tall‐grass Sorghastrum nutans to production increased at elevated CO₂ on the two most coarse‐textured of the soils studied, a clay Mollisol and sandy Alfisol. The CO₂‐caused increase in Sorghastrum was accompanied by an offsetting decline in production of the mid‐grass Bouteloua curtipendula. Increased CO₂ favored the tall‐grass over mid‐grass by increasing soil water content and apparently intensifying competition for light or other resources (Mollisol) or reducing the frequency with which soil water dipped below threshold levels (Alfisol). An increase in CO₂ of 250 μmol mol^?1 above the pre‐industrial level thus led to a shift in the relative production of established species that is similar in magnitude to differences observed between mid‐grass and tallgrass prairies along a precipitation gradient in the central USA. By reducing water limitation to plants, atmospheric CO₂ enrichment may alter the composition and even structure of grassland vegetation.     

Soil CO2 efflux and its spatial variation in a Florida slash pine plantation

The efflux of CO2 from the soil surface can vary markedly in magnitude both in time and space and its correct determination is crucial in many ecological studies. In this paper, we report results of field measurements, using an open-top dynamic chamber, of soil CO2 efflux in a mature Florida slash pine (Pinus elliottii Engelm. var.elliottii) plantation. The daily average efflux was 0.217 mg CO2 m-2s-1 in the autumn and 0.087 mg CO2 m-2s-1 in the winter. Soil temperature, which accounts for most of the temporal variability in CO2 efflux, is by far the most influential factor controlling soil respiration rate and its temporal variation. The CO2 efflux in the slash pine plantation is highly spatially variable and effluxes from the soil under palmetto is significantly higher than that from the open floor. The CO2 efflux generally increases with increase in soil fine root biomass, litter and humus amount on the forest floor but is inversely related to the amount of organic matter in the mineral soil. The spatial variation in CO2 efflux can be well characterised by a simple multiple regression model incorporating live and dead biomass and soil total porosity as predictor variables. Understorey plants, mostly Serenoa repens, are an important component of the C cycle and the major contributor to the spatial heterogeneity of soil CO2 efflux. The influence of understorey plants on soil respiration is probably via two approaches: increasing litterfall and root metabolism, both consequently stimulating microbial activity in the mineral soil.     

Increased nitrate availability in the soil of a mixed mature temperate forest subjected to elevated CO2 concentration (canopy FACE)

In a mature temperate forest in Hofstetten, Switzerland, deciduous tree canopies were subjected to a free‐air CO₂ enrichment (FACE) for a period of 8 years. The effect of this treatment on the availability of nitrogen (N) in the soil was assessed along three transects across the experimental area, one under Fagus sylvatica, one under Quercus robur and Q. petraea and one under Carpinus betulus. Nitrate, ammonium and dissolved organic N (DON) were analysed in soil solution obtained with suction cups. Nitrate and ammonium were also captured in buried ion‐exchange resin bags. These parameters were related to the local intensity of the FACE treatment as measured from the ¹³C depletion of dissolved inorganic carbon in the soil solution. Over the 8 years of experiment, the CO₂ enrichment reduced DON concentrations, did not affect ammonium, but induced higher nitrate concentrations, both in soil solution and resin bags. In the nitrate captured in the resin bags, the natural abundance of the isotope ¹⁵N increased strongly. This indicates that the CO₂ enrichment accelerated net nitrification, probably as an effect of the higher soil moisture resulting from the reduced transpiration of the CO₂‐enriched trees. It is also possible that N mineralization was enhanced by root exudates (priming effect) or that the uptake of inorganic N by these trees decreased slightly as the result of a reduced N demand for fine‐root growth. In this mature deciduous forest, we did not observe any progressive N limitation due to elevated atmospheric CO₂ concentrations; on the contrary, we observed an enhanced N availability over the 8 years of our measurements. This may, together with the global warming projected, exacerbate problems related to N saturation and nitrate leaching, although it is uncertain how long the observed trends will last in the future.     

Ecosystem fluxes of hydrogen in a mid‐latitude forest driven by soil microorganisms and plants

Molecular hydrogen (H₂) is an atmospheric trace gas with a large microbe‐mediated soil sink, yet cycling of this compound throughout ecosystems is poorly understood. Measurements of the sources and sinks of H₂ in various ecosystems are sparse, resulting in large uncertainties in the global H₂ budget. Constraining the H₂ cycle is critical to understanding its role in atmospheric chemistry and climate. We measured H₂ fluxes at high frequency in a temperate mixed deciduous forest for 15 months using a tower‐based flux‐gradient approach to determine both the soil‐atmosphere and the net ecosystem flux of H₂. We found that Harvard Forest is a net H₂ sink (?1.4 ± 1.1 kg H₂ ha^?1) with soils as the dominant H₂ sink (?2.0 ± 1.0 kg H₂ ha^?1) and aboveground canopy emissions as the dominant H₂ source (+0.6 ± 0.8 kg H₂ ha^?1). Aboveground emissions of H₂ were an unexpected and substantial component of the ecosystem H₂ flux, reducing net ecosystem uptake by 30% of that calculated from soil uptake alone. Soil uptake was highly seasonal (July maximum, February minimum), positively correlated with soil temperature and negatively correlated with environmental variables relevant to diffusion into soils (i.e., soil moisture, snow depth, snow density). Soil microbial H₂ uptake was correlated with rhizosphere respiration rates (r = 0.8, P < 0.001), and H₂ metabolism yielded up to 2% of the energy gleaned by microbes from carbon substrate respiration. Here, we elucidate key processes controlling the biosphere–atmosphere exchange of H₂ and raise new questions regarding the role of aboveground biomass as a source of atmospheric H₂ and mechanisms linking soil H₂ and carbon cycling. Results from this study should be incorporated into modeling efforts to predict the response of the H₂ soil sink to changes in anthropogenic H₂ emissions and shifting soil conditions with climate and land‐use change.     

CO2 efflux from a Mediterranean semi-arid forest soil. I. Seasonality and effects of stoniness

We studied the seasonality of total soil CO₂efflux and labeled C-CO₂ released from ¹⁴Clabeled straw incubated in the H horizon of asemi-arid Mediterranean forest soil. Fieldmeasurements were carried out over 520 days in aseries of reconstructed soil profiles with and withouta gravel layer below the H horizon. We monitored soilclimate and related this to soil CO₂ efflux.Seasonal variations in soil CO₂ efflux in asemiarid Mediterranean forest were mainly related tochanges in soil temperature. In spite of drought, highrespiration rates were observed in mid summer. Highsoil CO₂ efflux in hot and dry episodes wasattributed to increases in soil biological activity.The minimum soil CO₂ efflux occurred in latesummer also under dry conditions, probably related toa decrease in soil biological activity in deephorizons. Biological activity in organic layers waslimited by water potential () in summer and bytemperature in winter. Rewetting a dry soil resultedin large increases in soil CO₂ efflux only at hightemperatures. These large increases represented asignificant contribution to the decomposition oforganic matter in the uppermost horizons. Soilbiological activity in the uppermost horizons was moresensitive to changes in soil and hence tosummer rainstorms than the bulk soil microbialactivity. The presence of a layer of gravel improvedboth moisture and temperature conditions for thedecomposition of organic matter. As a result, soilCO₂ efflux increased in soils containing rockfragments. These effects were especially large for theorganic layers.