Sustained effects of atmospheric [CO2] and nitrogen availability on forest soil CO2 efflux

Soil CO₂ efflux (F_soil) is the largest source of carbon from forests and reflects primary productivity as well as how carbon is allocated within forest ecosystems. Through early stages of stand development, both elevated [CO₂] and availability of soil nitrogen (N; sum of mineralization, deposition, and fixation) have been shown to increase gross primary productivity, but the long‐term effects of these factors on F_soil are less clear. Expanding on previous studies at the Duke Free‐Air CO₂ Enrichment (FACE) site, we quantified the effects of elevated [CO₂] and N fertilization on F_soil using daily measurements from automated chambers over 10 years. Consistent with previous results, compared to ambient unfertilized plots, annual F_soil increased under elevated [CO₂] (ca. 17%) and decreased with N (ca. 21%). N fertilization under elevated [CO₂] reduced F_soil to values similar to untreated plots. Over the study period, base respiration rates increased with leaf productivity, but declined after productivity saturated. Despite treatment‐induced differences in aboveground biomass, soil temperature and water content were similar among treatments. Interannually, low soil water content decreased annual F_soil from potential values – estimated based on temperature alone assuming nonlimiting soil water content – by ca. 0.7% per 1.0% reduction in relative extractable water. This effect was only slightly ameliorated by elevated [CO₂]. Variability in soil N availability among plots accounted for the spatial variability in F_soil, showing a decrease of ca. 114 g C m^?2 yr^?1 per 1 g m^?2 increase in soil N availability, with consistently higher F_soil in elevated [CO₂] plots ca. 127 g C per 100 ppm [CO₂] over the +200 ppm enrichment. Altogether, reflecting increased belowground carbon partitioning in response to greater plant nutritional needs, the effects of elevated [CO₂] and N fertilization on F_soil in this stand are sustained beyond the early stages of stand development and through stabilization of annual foliage production.     

Natural variations in snow cover do not affect the annual soil CO2 efflux from a mid‐elevation temperate forest

Climate change might alter annual snowfall patterns and modify the duration and magnitude of snow cover in temperate regions with resultant impacts on soil microclimate and soil CO₂ efflux (F_soil). We used a 5‐year time series of F_soil measurements from a mid‐elevation forest to assess the effects of naturally changing snow cover. Snow cover varied considerably in duration (105–154 days) and depth (mean snow depth 19–59 cm). Periodically shallow snow cover (<10 cm) caused soil freezing or increased variation in soil temperature. This was mostly not reflected in F_soil which tended to decrease gradually throughout winter. Progressively decreasing C substrate availability (identified by substrate induced respiration) likely over‐rid the effects of slowly changing soil temperatures and determined the overall course of F_soil. Cumulative CO₂ efflux from beneath snow cover varied between 0.46 and 0.95 t C ha^?1 yr^?1 and amounted to between 6 and 12% of the annual efflux. When compared over a fixed interval (the longest period of snow cover during the 5 years), the cumulative CO₂ efflux ranged between 0.77 and 1.18 t C ha^?1 or between 11 and 15% of the annual soil CO₂ efflux. The relative contribution (15%) was highest during the year with the shortest winter. Variations in snow cover were not reflected in the annual CO₂ efflux (7.44–8.41 t C ha^?1) which did not differ significantly between years and did not correlate with any snow parameter. Regional climate at our site was characterized by relatively high amounts of precipitation. Therefore, snow did not play a role in terms of water supply during the warm season and primarily affected cold season processes. The role of changing snow cover therefore seems rather marginal when compared to potential climate change effects on F_soil during the warm season.     

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.     

Shrub expansion stimulates soil C and N storage along a coastal soil chronosequence

Expansion of woody vegetation in grasslands is a worldwide phenomenon with implications for C and N cycling at local, regional and global scales. Although woody encroachment is often accompanied by increased annual net primary production (ANPP) and increased inputs of litter, mesic ecosystems may become sources for C after woody encroachment because stimulation of soil CO₂ efflux releases stored soil carbon. Our objective was to determine if young, sandy soils on a barrier island became a sink for C after encroachment of the nitrogen‐fixing shrub Morella cerifera, or if associated stimulation of soil CO₂ efflux mitigated increased litterfall. We monitored variations in litterfall in shrub thickets across a chronosequence of shrub expansion and compared those data to previous measurements of ANPP in adjacent grasslands. In the final year, we quantified standing litter C and N pools in shrub thickets and soil organic matter (SOM), soil organic carbon (SOC), soil total nitrogen (TN) and soil CO₂ efflux in shrub thickets and adjacent grasslands. Heavy litterfall resulted in a dense litter layer storing an average of 809 g C m^?2 and 36 g N m^?2. Although soil CO₂ efflux was stimulated by shrub encroachment in younger soils, soil CO₂ efflux did not vary between shrub thickets and grasslands in the oldest soils and increases in CO₂ efflux in shrub thickets did not offset contributions of increased litterfall to SOC. SOC was 3.6–9.8 times higher beneath shrub thickets than in grassland soils and soil TN was 2.5–7.7 times higher under shrub thickets. Accumulation rates of soil and litter C were highest in the youngest thicket at 101 g m^?2 yr^?1 and declined with increasing thicket age. Expansion of shrubs on barrier islands, which have low levels of soil carbon and high potential for ANPP, has the potential to significantly increase ecosystem C sequestration.     

Effects of soil phosphorus availability,temperature and moisture on soil respiration in Eucalyptus pauciflora forest

Rates of soil respiration (CO₂ efflux) were measured for a year in a mature Eucalyptus pauciflora forest in unfertilized and phosphorus-fertilized plots. Soil CO₂ efflux showed a distinct seasonal trend, and average daily rates ranged from 124 to 574 mg CO₂ m^–2 hr^–1. Temperature and moisture are the main variables that cause variation in soil CO₂ efflux; hence their effects were investigated over a year so as to then differentiate the treatment effect of phosphorus (P) nutrition.Soil temperature had the greatest effect on CO₂ efflux and exhibited a highly significant logarithmic relationship (r² = 0.81). Periods of low soil and litter moisture occurred during summer when temperatures were greater than 10 °C, and this resulted in depression of soil CO₂ efflux. During winter, when temperatures were less than 10 °C, soil and litter moisture were consistently high and thus their variation had little effect on soil CO₂ efflux. A multiple regression model including soil temperature, and soil and litter moisture accounted for 97% of the variance in rates of CO₂ efflux, and thus can be used to predict soil CO₂ efflux at this site with high accuracy. Total annual efflux of carbon from soil was estimated to be 7.11 t C ha^–1 yr^–1. The model was used to predict changes in this annual flux if temperature and moisture conditions were altered. The extent to which coefficients of the model differ among sites and forest types requires testing.Increased soil P availability resulted in a large increase in stem growth of trees but a reduction in the rate of soil CO₂ efflux by approximately 8%. This reduction is suggested to be due to lower root activity resulting from reduced allocation of assimilate belowground. Root activity changed when P was added to microsites within plots, and via the whole tree root system at the plot level. These relationships of belowground carbon fluxes with temperature, moisture and nutrient availability provide essential information for understanding and predicting potential changes in forest ecosystems in response to land use management or climate change.     

C and N allocation in soil under ryegrass and alfalfa estimated by 13C and 15N labelling

Background and Aims

Below-ground translocated carbon (C) released as rhizodeposits is an important driver for microbial mobilization of nitrogen (N) for plants. We investigated how a limited substrate supply due to reduced photoassimilation alters the allocation of recently assimilated C in plant and soil pools under legume and non-legume species.

Methods

A non-legume (Lolium perenne) and a legume (Medicago sativa) were labelled with ¹⁵N before the plants were clipped or shaded, and labelled twice with ¹³CO₂ thereafter. Ten days after clipping and shading, the ¹⁵N and ¹³C in shoots, roots, soil, dissolved organic nitrogen (DON) and carbon (DOC) and in microbial biomass, as well as the ¹³C in soil CO₂ were analyzed.

Results

After clipping, about 50 % more ¹³C was allocated to regrowing shoots, resulting in a lower translocation to roots compared to the unclipped control. Clipping also reduced the total soil CO₂ efflux under both species and the ¹³C recovery of soil CO₂ under L. perenne. The ¹⁵N recovery increased in the shoots of M. sativa after clipping, because storage compounds were remobilized from the roots and/or the N uptake from the soil increased. After shading, the assimilated ¹³C was preferentially retained in the shoots of both species. This caused a decreased ¹³C recovery in the roots of M. sativa. Similarly, the total soil CO₂ efflux under M. sativa decreased more than 50 % after shading. The ¹⁵N recovery in plant and soil pools showed that shading has no effect on the N uptake and N remobilization for L. perenne, but, the ¹⁵N recovery increased in the shoot of M. sativa.

Conclusions

The experiment showed that the dominating effect on C and N allocation after clipping is the need of C and N for shoot regrowth, whereas the dominating effect after shading is the reduced substrate supply for growth and respiration. Only slight differences could be observed between L. perenne and M. sativa in the C and N distribution after clipping or shading.     

Contrasting soil respiration in young and old-growth ponderosa pine forests

Three years of fully automated and manual measurements of soil CO₂ efflux, soil moisture and temperature were used to explore the diel, seasonal and inter‐annual patterns of soil efflux in an old‐growth (250‐year‐old, O site) and recently regenerating (14‐year‐old, Y site) ponderosa pine forest in central Oregon. The data were used in conjunction with empirical models to determine which variables could be used to predict soil efflux in forests of contrasting ages and disturbance histories. Both stands experienced similar meteorological conditions with moderately cold wet winters and hot dry summers. Soil CO₂ efflux at both sites showed large inter‐annual variability that could be attributed to soil moisture availability in the deeper soil horizons (O site) and the quantity of summer rainfall (Y site). Seasonal patterns of soil CO₂ efflux at the O site showed a strong positive correlation between diel mean soil CO₂ efflux and soil temperature at 64 cm depth whereas diel mean soil efflux at the Y site declined before maximum soil temperature occurred during summer drought. The use of diel mean soil temperature and soil water potential inferred from predawn foliage water potential measurements could account for 80% of the variance of diel mean soil efflux across 3 years at both sites, however, the functional shape of the soil water potential constraint was site‐specific. Based on the similarity of the decomposition rates of litter and fine roots between sites, but greater productivity and amount of fine litter detritus available for decomposition at the O site, we would expect higher rates of soil CO₂ efflux at the O site. However, annual rates were only higher at the O site in one of the 3 years (597 ± 45 vs. 427 ± 80 g C m^?2). Seasonal patterns of soil efflux at both sites showed influences of soil water limitations that were also reflected in patterns of canopy stomatal conductance, suggesting strong linkages between above and below ground processes.     

Role of soil animals in C and N mineralisation

Addition of single species of soil animals to animal-free microcosms often increases total heterotrophic respiration, but sometimes additions of microarthropods have been reported not to increase or even decrease CO₂ evolution rates. Most studies indicate that addition of soil animals increases net N mineralisation. In a study with F/H layer materials from a spruce stand in central Sweden kept at two temperatures (5 and 15°C) and three moisture levels (15, 30 and 60% of WHC), addition of a mixed fauna of soil arthropods, mainly microarthropods, could not be shown to change the CO₂ evolution rates in comparison with materials where arthropods were absent. However, addition of the arthropods significantly increased net N mineralisation for each of the temperature and moisture combinations. The increase due to the arthropods was dependent on soil temperature but not on soil moisture. Because the total net N mineralisation decreased with decreasing soil moisture, the soil arthropods had a much larger relative effect on net N mineralisation under dry than under moist conditions. It is concluded that soil arthropods are important in maintaining net N mineralisation under dry conditions when the microflora is largely inactive. The microbial/faunal release of mineral N is discussed in relation to the CN of the substrate.     

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.     

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.     

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.     

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.     

青藏高原高寒草甸土壤CO2排放对模拟氮沉降的早期响应

研究大气氮沉降输入对青藏高原高寒草甸土壤-大气界面CO₂交换通量的影响,对于准确评价全球变化背景下区域碳平衡至关重要。通过构建多形态、低剂量的增氮控制试验,利用静态箱-气相色谱法测定土壤CO₂排放通量,同时测定相关土壤变量和地上生物量,分析高寒草甸土壤CO₂排放特征及其主要驱动因子。研究结果表明:低、高剂量氮输入倾向于消耗土壤水分,而中剂量氮输入有利于土壤水分的保持;施氮初期总体上增加了土壤无机氮含量,铵态氮累积效应更为显著;施氮显著增加地上生物量和土壤CO₂排放通量,铵态氮的促进效应显著高于硝态氮。另外,土壤CO₂排放通量主要受土壤温度驱动,其次为地上生物量和铵态氮储量。上述结果反映了氮沉降输入短期内可能刺激了植物生长和土壤微生物活性,加剧了土壤-大气界面CO₂排放。     

Soil pCO2, soil respiration,and root activity in CO2-fumigated and nitrogen-fertilized ponderosa pine

The purpose of this paper is to describe the effects of CO₂ and N treatments on soil pCO₂, calculated CO₂ efflux, root biomass and soil carbon in open-top chambers planted with Pinus ponderosa seedlings. Based upon the literature, it was hypothesized that both elevated CO₂ and N would cause increased root biomass which would in turn cause increases in both total soil CO₂ efflux and microbial respiration. This hypothesis was only supported in part: both CO₂ and N treatments caused significant increases in root biomass, soil pCO₂, and calculated CO₂ efflux, but there were no differences in soil microbial respiration measured in the laboratory. Both correlative and quantitative comparisons of CO₂ efflux rates indicated that microbial respiration contributes little to total soil CO₂ efflux in the field. Measurements of soil pCO₂ and calculated CO₂ efflux provided inexpensive, non-invasive, and relatively sensitive indices of belowground response to CO₂ and N treatments.     

High arctic heath soil respiration and biogeochemical dynamics during summer and autumn freeze‐in – effects of long‐term enhanced water and nutrient supply

In High Arctic NE Greenland, temperature and precipitation are predicted to increase during this century, however, relatively little information is available on the role of increased water supply on soil CO ₂ efflux in dry, high arctic ecosystems. We measured soil respiration (R_soil) in summer and autumn of 2009 in combination with microbial biomass and nutrient availability during autumn freeze‐in at a dry, open heath in Zackenberg, NE Greenland. This tundra site has been subject to fully factorial manipulation consisting of increased soil water supply for 14 years, and occasional nitrogen (N) addition in pulses. Summer watering enhanced R_soil during summer, but decreased R_soil in the following autumn. We speculate that this is due to intensified depletion of recently fixed plant carbon by soil organisms. Hence, autumn soil microbial activity seems tightly linked to growing season plant production through plant‐associated carbon pools. Nitrogen addition alone consistently increased R_soil, but when water and nitrogen were added in combination, autumn R_soil declined similarly to when water was added alone. Despite several freeze‐thaw events, the microbial biomass carbon (C) remained constant until finally being reduced by ~60% in late September. In spite of significantly reduced microbial biomass C and phosphorus (P), microbial N did not change. This suggests N released from dead microbes was quickly assimilated by surviving microbes. We observed no change in soil organic matter content after 14 years of environmental manipulations, suggesting high ecosystem resistance to environmental changes.     

Soil‐surface CO2 efflux and its spatial and temporal variations in a young ponderosa pine plantation in northern California

Soil‐surface CO₂ efflux and its spatial and temporal variations were examined in an 8‐y‐old ponderosa pine plantation in the Sierra Nevada Mountains in California from June 1998 to August 1999. Continuous measurements of soil CO₂ efflux, soil temperatures and moisture were conducted on two 20 × 20 m sampling plots. Microbial biomass, fine root biomass, and the physical and chemical properties of the soil were also measured at each of the 18 sampling locations on the plots. It was found that the mean soil CO₂ efflux in the plantation was 4.43 µmol m^?2 s^?1 in the growing season and 3.12 µmol m^?2 s^?1 in the nongrowing season. These values are in the upper part of the range of published soil‐surface CO₂ efflux data. The annual maximum and minimum CO₂ efflux were 5.87 and 1.67 µmol m^?2 s^?1, respectively, with the maximum occurring between the end of May and early June and the minimum in December. The diurnal fluctuation of CO₂ efflux was relatively small (< 20%) with the minimum appearing around 09.00 hours and the maximum around 14.00 hours. Using daytime measurements of soil CO₂ efflux tends to overestimate the daily mean soil CO₂ efflux by 4–6%. The measurements taken between 09.00 and 11.00 hours (local time) seem to better represent the daily mean with a reduced sampling error of 0.9–1.5%. The spatial variation of soil CO₂ efflux among the 18 sampling points was high, with a coefficient of variation of approximately 30%. Most (84%) of the spatial variation was explained by fine root biomass, microbial biomass, and soil physical and chemical properties. Although soil temperature and moisture explained most of the temporal variations (76–95%) of soil CO₂ efflux, the two variables together explained less than 34% of the spatial variation. Microbial biomass, fine root biomass, soil nitrogen content, organic matter content, and magnesium content were significantly and positively correlated with soil CO₂ efflux, whereas bulk density and pH value were negatively correlated with CO₂ efflux. The relationship between soil CO₂ efflux and soil temperature was significantly controlled by soil moisture with a Q₁₀ value of 1.4 when soil moisture was <14% and 1.8 when soil moisture was >14%. Understanding the spatial and temporal variations is essential to accurately assessment of carbon budget at whole ecosystem and landscape scales. Thus, this study bears important implications for the study of large‐scale ecosystem dynamics, particularly in response to climatic variations and management regimes.     

Soil C and N responses to woody plant expansion in a mesic grassland

Changes in land management and reductions in fire frequency have contributed to increased cover of woody species in grasslands worldwide. These shifts in plant community composition have the potential to alter ecosystem function, particularly through changes in soil processes and properties. In semi-arid grasslands, the invasion of shrubs and trees is often accompanied by increases in soil resources and more rapid N and C cycling. We assessed the effects of shrub encroachment in a mesic grassland in Kansas (USA) on soil CO₂ flux, extractable inorganic N, and N mineralization beneath shrub communities (Cornus drummondii) and surrounding undisturbed grassland sites. In this study, a shift in plant community composition from grassland to shrubland resulted in a 16% decrease in annual soil CO₂ flux(4.78 kg CO₂ m^–2 year^–1 for shrub dominated sites versus 5.84 kg CO₂ m^–2 year^–1 for grassland sites) with no differences in total soil C or N or inorganic N. There was considerable variability in N mineralization rates within sites, which resulted in no overall difference in cumulative N mineralized during this study (4.09 g N m^–2 for grassland sites and 3.03 g N m^–2 for shrub islands). These results indicate that shrub encroachment into mesic grasslands does not significantly alter N availability (at least initially), but does alter C cycling by decreasing soil CO₂ flux.     

Effects of experimental drought on soil respiration and radiocarbon efflux from a temperate forest soil

Soil moisture affects microbial decay of SOM and rhizosphere respiration (RR) in temperate forest soils, but isolating the response of soil respiration (SR) to summer drought and subsequent wetting is difficult because moisture changes are often confounded with temperature variation. We distinguished between temperature and moisture effects by simulation of prolonged soil droughts in a mixed deciduous forest at the Harvard Forest, Massachusetts. Roofs constructed over triplicate 5 × 5 m² plots excluded throughfall water during the summers of 2001 (168 mm) and 2002 (344 mm), while adjacent control plots received ambient throughfall and the same natural temperature regime. In 2003, throughfall was not excluded to assess the response of SR under natural weather conditions after two prolonged summer droughts. Throughfall exclusion significantly decreased mean SR rate by 53 mg C m^?2 h^?1 over 84 days in 2001, and by 68 mg C m^?2 h^?1 over 126 days in 2002, representing 10–30% of annual SR in this forest and 35–75% of annual net ecosystem exchange (NEE) of C. The differences in SR were best explained by differences in gravimetric water content in the Oi horizon (r²=0.69) and the Oe/Oa horizon (r²=0.60). Volumetric water content of the A horizon was not significantly affected by throughfall exclusion. The radiocarbon signature of soil CO₂ efflux and of CO₂ respired during incubations of O horizon, A horizon and living roots allowed partitioning of SR into contributions from young C substrate (including RR) and from decomposition of older SOM. RR (root respiration and microbial respiration of young substrates in the rhizosphere) made up 43–71% of the total C respired in the control plots and 41–80% in the exclusion plots, and tended to increase with drought. An exception to this trend was an interesting increase in CO₂ efflux of radiocarbon‐rich substrates during a period of abundant growth of mushrooms. Our results suggest that prolonged summer droughts decrease primarily heterotrophic respiration in the O horizon, which could cause increases in the storage of soil organic carbon in this forest. However, the C stored during two summers of simulated drought was only partly released as increased respiration during the following summer of natural throughfall. We do not know if this soil C sink during drought is transient or long lasting. In any case, differential decomposition of the O horizon caused by interannual variation of precipitation probably contributes significantly to observed interannual variation of NEE in temperate forests.     

Soil respiration under climate change: prolonged summer drought offsets soil warming effects

Climate change may considerably impact the carbon (C) dynamics and C stocks of forest soils. To assess the combined effects of warming and reduced precipitation on soil CO₂ efflux, we conducted a two‐way factorial manipulation experiment (4 °C soil warming + throughfall exclusion) in a temperate spruce forest from 2008 until 2010. Soil was warmed by heating cables throughout the growing seasons. Soil drought was simulated by throughfall exclusions with three 100 m² roofs during 25 days in July/August 2008 and 2009. Soil warming permanently increased the CO₂ efflux from soil, whereas throughfall exclusion led to a sharp decrease in soil CO₂ efflux (45% and 50% reduction during roof installation in 2008 and 2009, respectively). In 2008, CO₂ efflux did not recover after natural rewetting and remained lowered until autumn. In 2009, CO₂ efflux recovered shortly after rewetting, but relapsed again for several weeks. Drought offset the increase in soil CO₂ efflux by warming in 2008 (growing season CO₂ efflux in t C ha^?1: control: 7.1 ± 1.0; warmed: 9.5 ± 1.7; warmed + roof: 7.4 ± 0.3; roof: 5.9 ± 0.4) and in 2009 (control: 7.6 ± 0.8; warmed + roof: 8.3 ± 1.0). Throughfall exclusion mainly affected the organic layer and the top 5 cm of the mineral soil. Radiocarbon data suggest that heterotrophic and autotrophic respiration were affected to the same extent by soil warming and drying. Microbial biomass in the mineral soil (0–5 cm) was not affected by the treatments. Our results suggest that warming causes significant C losses from the soil as long as precipitation patterns remain steady at our site. If summer droughts become more severe in the future, warming induced C losses will likely be offset by reduced soil CO₂ efflux during and after summer drought.     

Annual burning of a tallgrass prairie inhibits C and N cycling in soil,increasing recalcitrant pyrogenic organic matter storage while reducing N availability

Grassland ecosystems store an estimated 30% of the world's total soil C and are frequently disturbed by wildfires or fire management. Aboveground litter decomposition is one of the main processes that form soil organic matter (SOM). However, during a fire biomass is removed or partially combusted and litter inputs to the soil are substituted with inputs of pyrogenic organic matter (py‐OM). Py‐OM accounts for a more recalcitrant plant input to SOM than fresh litter, and the historical frequency of burning may alter C and N retention of both fresh litter and py‐OM inputs to the soil. We compared the fate of these two forms of plant material by incubating ¹³C‐ and ¹⁵N‐labeled Andropogon gerardii litter and py‐OM at both an annually burned and an infrequently burned tallgrass prairie site for 11 months. We traced litter and py‐OM C and N into uncomplexed and organo‐mineral SOM fractions and CO₂ fluxes and determined how fire history affects the fate of these two forms of aboveground biomass. Evidence from CO₂ fluxes and SOM C:N ratios indicates that the litter was microbially transformed during decomposition while, besides an initial labile fraction, py‐OM added to SOM largely untransformed by soil microbes. Additionally, at the N‐limited annually burned site, litter N was tightly conserved. Together, these results demonstrate how, although py‐OM may contribute to C and N sequestration in the soil due to its resistance to microbial degradation, a long history of annual removal of fresh litter and input of py‐OM infers N limitation due to the inhibition of microbial decomposition of aboveground plant inputs to the soil. These results provide new insight into how fire may impact plant inputs to the soil, and the effects of py‐OM on SOM formation and ecosystem C and N cycling.