Does declining carbon‐use efficiency explain thermal acclimation of soil respiration with warming?

Enhanced soil respiration in response to global warming may substantially increase atmospheric CO₂ concentrations above the anthropogenic contribution, depending on the mechanisms underlying the temperature sensitivity of soil respiration. Here, we compared short‐term and seasonal responses of soil respiration to a shifting thermal environment and variable substrate availability via laboratory incubations. To analyze the data from incubations, we implemented a novel process‐based model of soil respiration in a hierarchical Bayesian framework. Our process model combined a Michaelis–Menten‐type equation of substrate availability and microbial biomass with an Arrhenius‐type nonlinear temperature response function. We tested the competing hypotheses that apparent thermal acclimation of soil respiration can be explained by depletion of labile substrates in warmed soils, or that physiological acclimation reduces respiration rates. We demonstrated that short‐term apparent acclimation can be induced by substrate depletion, but that decreasing microbial biomass carbon (MBC) is also important, and lower MBC at warmer temperatures is likely due to decreased carbon‐use efficiency (CUE). Observed seasonal acclimation of soil respiration was associated with higher CUE and lower basal respiration for summer‐ vs. winter‐collected soils. Whether the observed short‐term decrease in CUE or the seasonal acclimation of CUE with increased temperatures dominates the response to long‐term warming will have important consequences for soil organic carbon storage.     

Soil microbial respiration in arctic soil does not acclimate to temperature

Warming-induced release of CO2 from the large carbon (C) stores in arctic soils could accelerate climate change. However, declines in the response of soil respiration to warming in long-term experiments suggest that microbial activity acclimates to temperature, greatly reducing the potential for enhanced C losses. As reduced respiration rates with time could be equally caused by substrate depletion, evidence for thermal acclimation remains controversial. To overcome this problem, we carried out a cooling experiment with soils from arctic Sweden. If acclimation causes the reduction in soil respiration observed after experimental warming, then it should subsequently lead to an increase in respiration rates after cooling. We demonstrate that thermal acclimation did not occur following cooling. Rather, during the 90 days after cooling, a further reduction in the soil respiration rate was observed, which was only reversed by extended re-exposure to warmer temperatures. We conclude that over the time scale of a few weeks to months, warming-induced changes in the microbial community in arctic soils will amplify the instantaneous increase in the rates of CO2 production and thus enhance C losses potentially accelerating the rate of 21st century climate change.     

Apparent thermal acclimation of soil heterotrophic respiration mainly mediated by substrate availability

Multiple lines of existing evidence suggest that increasing CO₂ emission from soils in response to rising temperature could accelerate global warming. However, in experimental studies, the initial positive response of soil heterotrophic respiration (R_H) to warming often weakens over time (referred to apparent thermal acclimation). If the decreased R_H is driven by thermal adaptation of soil microbial community, the potential for soil carbon (C) losses would be reduced substantially. In the meanwhile, the response could equally be caused by substrate depletion, and would then reflect the gradual loss of soil C. To address uncertainties regarding the causes of apparent thermal acclimation, we carried out sterilization and inoculation experiments using the soil samples from an alpine meadow with 6 years of warming and nitrogen (N) addition. We demonstrate that substrate depletion, rather than microbial adaptation, determined the response of R_H to long-term warming. Furthermore, N addition appeared to alleviate the apparent acclimation of R_H to warming. Our study provides strong empirical support for substrate availability being the cause of the apparent acclimation of soil microbial respiration to temperature. Thus, these mechanistic insights could facilitate efforts of biogeochemical modeling to accurately project soil C stocks in the future climate.     

Geothermally warmed soils reveal persistent increases in the respiratory costs of soil microbes contributing to substantial C losses

Increasing temperatures can accelerate soil organic matter decomposition and release large amounts of CO₂ to the atmosphere, potentially inducing positive warming feedbacks. Alterations to the temperature sensitivity and physiological functioning of soil microorganisms may play a key role in these carbon (C) losses. Geothermally active areas in Iceland provide stable and continuous soil temperature gradients to test this hypothesis, encompassing the full range of warming scenarios projected by the Intergovernmental Panel on Climate Change for the northern region. We took soils from these geothermal sites 7 years after the onset of warming and incubated them at varying temperatures and substrate availability conditions to detect persistent alterations of microbial physiology to long-term warming. Seven years of continuous warming ranging from 1.8 to 15.9 °C triggered a 8.6–58.0% decrease on the C concentrations in the topsoil (0–10 cm) of these sub-arctic silt-loam Andosols. The sensitivity of microbial respiration to temperature (Q₁₀) was not altered. However, soil microbes showed a persistent increase in their microbial metabolic quotients (microbial respiration per unit of microbial biomass) and a subsequent diminished C retention in biomass. After an initial depletion of labile soil C upon soil warming, increasing energy costs of metabolic maintenance and resource acquisition led to a weaker capacity of C stabilization in the microbial biomass of warmer soils. This mechanism contributes to our understanding of the acclimated response of soil respiration to in situ soil warming at the ecosystem level, despite a lack of acclimation at the physiological level. Persistent increases in the respiratory costs of soil microbes in response to warming constitute a fundamental process that should be incorporated into climate change-C cycling models.     

Substrate availability and not thermal acclimation controls microbial temperature sensitivity response to long-term warming

Microbes are responsible for cycling carbon (C) through soils, and predicted changes in soil C stocks under climate change are highly sensitive to shifts in the mechanisms assumed to control the microbial physiological response to warming. Two mechanisms have been suggested to explain the long-term warming impact on microbial physiology: microbial thermal acclimation and changes in the quantity and quality of substrates available for microbial metabolism. Yet studies disentangling these two mechanisms are lacking. To resolve the drivers of changes in microbial physiology in response to long-term warming, we sampled soils from 13- and 28-year-old soil warming experiments in different seasons. We performed short-term laboratory incubations across a range of temperatures to measure the relationships between temperature sensitivity of physiology (growth, respiration, carbon use efficiency, and extracellular enzyme activity) and the chemical composition of soil organic matter. We observed apparent thermal acclimation of microbial respiration, but only in summer, when warming had exacerbated the seasonally-induced, already small dissolved organic matter pools. Irrespective of warming, greater quantity and quality of soil carbon increased the extracellular enzymatic pool and its temperature sensitivity. We propose that fresh litter input into the system seasonally cancels apparent thermal acclimation of C-cycling processes to decadal warming. Our findings reveal that long-term warming has indirectly affected microbial physiology via reduced C availability in this system, implying that earth system models including these negative feedbacks may be best suited to describe long-term warming effects on these soils.     

Effects of sieving, drying and rewetting upon soil bacterial community structure and respiration rates

Soil microcosm studies often require some form of soil homogenisation, such as sieving, to provide a representative sample. Frequently, soils are also homogenised following drying and are then rewetted, yet little research has been done to understand how these methods impact upon microbial communities. Here we compared the molecular diversity and functional responses of intact cores from a Scottish grassland soil with homogenised samples prepared by drying, sieving and rewetting or freshly sieving wet soils. Results showed that there was no significant difference in total soil CO₂-C efflux between the freshly sieved and intact core treatments, however, respiration was significantly higher in the dried and rewetted microcosms. Molecular fingerprinting (T-RFLP) of bacterial communities at two different time-points showed that both homogenisation methods significantly altered bacterial community structure with the largest differences being observed after drying and rewetting. Assessments of responsive taxa in each treatment showed that intact cores were dominated by Acidobacterial peaks whereas an increased relative abundance of Alphaproteobacterial terminal restriction fragments were apparent in both homogenised treatments. However, the shift in community structure was not as large in the freshly sieved soil. Our findings suggest that if soil homogenisation must be performed, then fresh sieving of wet soil is preferable to drying and rewetting in approximating the bacterial diversity and functioning of intact cores.     

Thermal adaptation of soil microbial respiration to elevated temperature

In the short‐term heterotrophic soil respiration is strongly and positively related to temperature. In the long‐term, its response to temperature is uncertain. One reason for this is because in field experiments increases in respiration due to warming are relatively short‐lived. The explanations proposed for this ephemeral response include depletion of fast‐cycling, soil carbon pools and thermal adaptation of microbial respiration. Using a > 15 year soil warming experiment in a mid‐latitude forest, we show that the apparent ‘acclimation’ of soil respiration at the ecosystem scale results from combined effects of reductions in soil carbon pools and microbial biomass, and thermal adaptation of microbial respiration. Mass‐specific respiration rates were lower when seasonal temperatures were higher, suggesting that rate reductions under experimental warming likely occurred through temperature‐induced changes in the microbial community. Our results imply that stimulatory effects of global temperature rise on soil respiration rates may be lower than currently predicted.     

Contrasting above‐ and belowground organic matter decomposition and carbon and nitrogen dynamics in response to warming in High Arctic tundra

Tundra regions are projected to warm rapidly during the coming decades. The tundra biome holds the largest terrestrial carbon pool, largely contained in frozen permafrost soils. With warming, these permafrost soils may thaw and become available for microbial decomposition, potentially providing a positive feedback to global warming. Warming may directly stimulate microbial metabolism but may also indirectly stimulate organic matter turnover through increased plant productivity by soil priming from root exudates and accelerated litter turnover rates. Here, we assess the impacts of experimental warming on turnover rates of leaf litter, active layer soil and thawed permafrost sediment in two high‐arctic tundra heath sites in NE‐Greenland, either dominated by evergreen or deciduous shrubs. We incubated shrub leaf litter on the surface of control and warmed plots for 1 and 2 years. Active layer soil was collected from the plots to assess the effects of 8 years of field warming on soil carbon stocks. Finally, we incubated open cores filled with newly thawed permafrost soil for 2 years in the active layer of the same plots. After field incubation, we measured basal respiration rates of recovered thawed permafrost cores in the lab. Warming significantly reduced litter mass loss by 26% after 1 year incubation, but differences in litter mass loss among treatments disappeared after 2 years incubation. Warming also reduced litter nitrogen mineralization and decreased the litter carbon to nitrogen ratio. Active layer soil carbon stocks were reduced 15% by warming, while soil dissolved nitrogen was reduced by half in warmed plots. Warming had a positive legacy effect on carbon turnover rates in thawed permafrost cores, with 10% higher respiration rates measured in cores from warmed plots. These results demonstrate that warming may have contrasting effects on above‐ and belowground tundra carbon turnover, possibly governed by microbial resource availability.     

Low soil moisture suppresses the thermal compensatory response of microbial respiration

The thermal compensatory response of microbial respiration contributes to a decrease in warming-induced enhancement of soil respiration over time, which could weaken the positive feedback between the carbon cycle and climate warming. Climate warming is also predicted to cause a worldwide decrease in soil moisture, which has an effect on the microbial metabolism of soil carbon. However, whether and how changes in moisture affect the thermal compensatory response of microbial respiration are unexplored. Here, using soils from an 8-year warming experiment in an alpine grassland, we assayed the thermal response of microbial respiration rates at different soil moisture levels. The results showed that relatively low soil moisture suppressed the thermal compensatory response of microbial respiration, leading to an enhanced response to warming. A subsequent moisture incubation experiment involving off-plot soils also showed that the response of microbial respiration to 100 d warming shifted from a slight compensatory response to an enhanced response with decreasing incubation moisture. Further analysis revealed that such respiration regulation by moisture was associated with shifts in enzymatic activities and carbon use efficiency. Our findings suggest that future drought induced by climate warming might weaken the thermal compensatory capacity of microbial respiration, with important consequences for carbon–climate feedback.     

Sucrose application,soil microbial respiration and evolved carbon dioxide isotope enrichment under contrasting land uses

Heterotrophic decomposition of organic matter dictates that substrate supply rate, including energy and nutrients, can limit soil microbial activity. In New Zealand, soils are naturally deficient in nitrogen and phosphorus. Fertiliser application is a part of pastoral agriculture, the countrys most widespread land use. We postulated that organic soils under grazed pasture and pristine forest would be at the extremes of substrate quality and supply rate, and thus potential microbial response to food opportunities. Soil microbial responses to the addition of fresh energy (sucrose) were determined by laboratory experiments with root-free samples and intact cores including roots. Responses were quantified by respiration and respired carbon (C) isotope (¹³C) enrichment measurements. A supra-trace sucrose dose (0.002 mol kg^–1 (soil)) caused the forest soils microbial respiration rate to nearly double within 2 h. The peak response took 20 h, and saturation occurred beyond a sucrose dose of 0.05 mol kg^–1 (soil). Intact soil cores from the forest had similar respiration rates and responses. For root-free soil samples from the grazed pasture, respiration response to sucrose was nearly immediate, dose dependent, and there was up to a 9-fold increase in the rate. Intact cores from the pasture had much higher respiration rates, but a similar response to sucrose. For both soils, the similarity of sucrose application effects on respiration and relative ¹³C enrichment of the respired carbon was striking.     

Increasing microbial carbon use efficiency with warming predicts soil heterotrophic respiration globally

The degree to which climate warming will stimulate soil organic carbon (SOC) losses via heterotrophic respiration remains uncertain, in part because different or even opposite microbial physiology and temperature relationships have been proposed in SOC models. We incorporated competing microbial carbon use efficiency (CUE)–mean annual temperature (MAT) and enzyme kinetic–MAT relationships into SOC models, and compared the simulated mass‐specific soil heterotrophic respiration rates with multiple published datasets of measured respiration. The measured data included 110 dryland soils globally distributed and two continental to global‐scale cross‐biome datasets. Model–data comparisons suggested that a positive CUE–MAT relationship best predicts the measured mass‐specific soil heterotrophic respiration rates in soils distributed globally. These results are robust when considering models of increasing complexity and competing mechanisms driving soil heterotrophic respiration–MAT relationships (e.g., carbon substrate availability). Our findings suggest that a warmer climate selects for microbial communities with higher CUE, as opposed to the often hypothesized reductions in CUE by warming based on soil laboratory assays. Our results help to build the impetus for, and confidence in, including microbial mechanisms in soil biogeochemical models used to forecast changes in global soil carbon stocks in response to warming.     

Soil respiration under prolonged soil warming: are rate reductions caused by acclimation or substrate loss?

The world's soils contain a large amount of carbon so that even a fractionally small loss or gain could have a quantitatively important feedback effect on net CO₂ emissions to the atmosphere. It is therefore important to fully understand the temperature dependence of soil‐carbon decomposition. Evidence from various observations can be used to quantify the temperature dependence of carbon efflux, but it is important to ensure that confounding factors, such as changing water relations or availability of readily decomposable substrate, are fully considered in inferring an underlying temperature response from observed response patterns. A number of recent findings from soil‐warming experiments have led to the suggestion that stimulation of soil‐carbon efflux by increasing temperature is only transitory before acclimation takes place and carbon efflux rates return to similar rates as before the increase in temperature. It is shown here that this response pattern can be explained through a simple two‐pool soil‐carbon model with no acclimation response needing to be invoked. The temporal pattern is, instead, due to depletion of readily decomposable substrate. It shows that findings of reduced respiration rate in soil‐warming experiments are consistent with unchanged high temperature sensitivity of organic carbon decomposition and affirms that there is, indeed, a danger of positive feedback between global warming and the release of soil organic carbon that can lead to further warming.     

Thermal acclimation in widespread heterotrophic soil microbes

Respiration by plants and microorganisms is primarily responsible for mediating carbon exchanges between the biosphere and atmosphere. Climate warming has the potential to influence the activity of these organisms, regulating exchanges between carbon pools. Physiological ‘down‐regulation’ of warm‐adapted species (acclimation) could ameliorate the predicted respiratory losses of soil carbon under climate change scenarios, but unlike plants and symbiotic microbes, the existence of this phenomenon in heterotrophic soil microbes remains controversial. Previous studies using complex soil microbial communities are unable to distinguish physiological acclimation from other community‐scale adjustments. We explored the temperature‐sensitivity of individual saprotrophic basidiomycete fungi growing in agar, showing definitively that these widespread heterotrophic fungi can acclimate to temperature. In almost all cases, the warm‐acclimated individuals had lower growth and respiration rates at intermediate temperatures than cold‐acclimated isolates. Inclusion of such microbial physiological responses to warming is essential to enhance the robustness of global climate‐ecosystem carbon models.     

Interactive effects of wildfire and permafrost on microbial communities and soil processes in an Alaskan black spruce forest

Boreal forests contain significant quantities of soil carbon that may be oxidized to CO₂ given future increases in climate warming and wildfire behavior. At the ecosystem scale, decomposition and heterotrophic respiration are strongly controlled by temperature and moisture, but we questioned whether changes in microbial biomass, activity, or community structure induced by fire might also affect these processes. We particularly wanted to understand whether postfire reductions in microbial biomass could affect rates of decomposition. Additionally, we compared the short‐term effects of wildfire to the long‐term effects of climate warming and permafrost decline. We compared soil microbial communities between control and recently burned soils that were located in areas with and without permafrost near Delta Junction, AK. In addition to soil physical variables, we quantified changes in microbial biomass, fungal biomass, fungal community composition, and C cycling processes (phenol oxidase enzyme activity, lignin decomposition, and microbial respiration). Five years following fire, organic surface horizons had lower microbial biomass, fungal biomass, and dissolved organic carbon (DOC) concentrations compared with control soils. Reductions in soil fungi were associated with reductions in phenol oxidase activity and lignin decomposition. Effects of wildfire on microbial biomass and activity in the mineral soil were minor. Microbial community composition was affected by wildfire, but the effect was greater in nonpermafrost soils. Although the presence of permafrost increased soil moisture contents, effects on microbial biomass and activity were limited to mineral soils that showed lower fungal biomass but higher activity compared with soils without permafrost. Fungal abundance and moisture were strong predictors of phenol oxidase enzyme activity in soil. Phenol oxidase enzyme activity, in turn, was linearly related to both ¹³C lignin decomposition and microbial respiration in incubation studies. Taken together, these results indicate that reductions in fungal biomass in postfire soils and lower soil moisture in nonpermafrost soils reduced the potential of soil heterotrophs to decompose soil carbon. Although in the field increased rates of microbial respiration can be observed in postfire soils due to warmer soil conditions, reductions in fungal biomass and activity may limit rates of decomposition.     

Adenylate control contributes to thermal acclimation of sugar maple fine‐root respiration in experimentally warmed soil

We investigated the occurrence of and mechanisms responsible for acclimation of fine‐root respiration of mature sugar maple (Acer saccharum) after 3+ years of experimental soil warming (+4 to 5 °C) in a factorial combination with soil moisture addition. Potential mechanisms for thermal respiratory acclimation included changes in enzymatic capacity, as indicated by root N concentration; substrate limitation, assessed by examining nonstructural carbohydrates and effects of exogenous sugar additions; and adenylate control, examined as responses of root respiration to a respiratory uncoupling agent. Partial acclimation of fine‐root respiration occurred in response to soil warming, causing specific root respiration to increase to a much lesser degree (14% to 26%) than would be expected for a 4 to 5 °C temperature increase (approximately 55%). Acclimation was greatest when ambient soil temperature was warmer or soil moisture availability was low. We found no evidence that enzyme or substrate limitation caused acclimation but did find evidence supporting adenylate control. The uncoupling agent caused a 1.4 times greater stimulation of respiration in roots from warmed soil. Sugar maple fine‐root respiration in warmed soil was at least partially constrained by adenylate use, helping constrain respiration to that needed to support work being performed by the roots.     

Investigating biological control over soil carbon temperature sensitivity

Understanding the temperature sensitivity of soil respiration is critical for predicting the response of ecosystems to climate change, yet the microbial communities responsible are rarely considered explicitly in studies or models. In this study, we assessed total microbial community composition, quantified bacterial respiration temperature response, and investigated the temperature dependence of bacterial carbon substrate utilization in tropical, temperate, and taiga soils (from Puerto Rico, California, and Alaska). Microbial community composition was characterized using phospholipid fatty acid analysis. Bacterial community respiration on a standardized set of substrates was ascertained using the BiOLOG^™ substrate utilization assay incubated at four temperatures: 4, 12, 28, and 40 °C. First, we found that microbial communities from the three latitudes were compositionally distinct and that the bacterial component of the three communities had markedly different respiration temperature–response curves corresponding with their experienced temperature regimes. We use these data to highlight limitations of widely used temperature–response equations and investigate temperature-dependent patterns of substrate utilization. We found that temperature response, in terms of both respiration rates and substrate use, varied for these bacterial communities independent of substrate quality or quantity interactions such as labile depletion. In contrast to the common assumption of heterotrophic microbial ubiquity, we found that bacterial community differences from these diverse systems appeared to determine both rates of respiration and patterns of carbon substrate usage. We suggest that microbial community composition-specific responses to changing climate may be important in predicting the long-term role of ecosystems in atmospheric CO₂ dynamics.     

How interactions between microbial resource demands,soil organic matter stoichiometry,and substrate reactivity determine the direction and magnitude of soil respiratory responses to warming

Recent empirical and theoretical advances inform us about multiple drivers of soil organic matter (SOM) decomposition and microbial responses to warming. Absent from our conceptual framework of how soil respiration will respond to warming are adequate links between microbial resource demands, kinetic theory, and substrate stoichiometry. Here, we describe two important concepts either insufficiently explored in current investigations of SOM responses to temperature, or not yet addressed. First, we describe the complete range of responses for how warming may change microbial resource demands, physiology, community structure, and total biomass. Second, we describe how any relationship between SOM activation energy of decay and carbon (C) and nitrogen (N) stoichiometry can alter the relative availability of C and N as temperature changes. Changing availabilities of C and N liberated from their organic precursors can feedback to microbial resource demands, which in turn influence the aggregated respiratory response to temperature we observe. An unsuspecting biogeochemist focused primarily on temperature sensitivity of substrate decay thus cannot make accurate projections of heterotrophic CO₂ losses from diverse organic matter reservoirs in a warming world. We establish the linkages between enzyme kinetics, SOM characteristics, and potential for microbial adaptation critical for making such projections. By examining how changing microbial needs interact with inherent SOM structure and composition, and thus reactivity, we demonstrate the means by which increasing temperature could result in increasing, unchanging, or even decreasing respiration rates observed in soils. We use this exercise to highlight ideas for future research that will develop our abilities to predict SOM feedbacks to climate.     

Microbial activity and soil respiration under nitrogen addition in Alaskan boreal forest

Climate warming could increase rates of soil organic matter turnover and nutrient mineralization, particularly in northern high‐latitude ecosystems. However, the effects of increasing nutrient availability on microbial processes in these ecosystems are poorly understood. To determine how soil microbes respond to nutrient enrichment, we measured microbial biomass, extracellular enzyme activities, soil respiration, and the community composition of active fungi in nitrogen (N) fertilized soils of a boreal forest in central Alaska. We predicted that N addition would suppress fungal activity relative to bacteria, but stimulate carbon (C)‐degrading enzyme activities and soil respiration. Instead, we found no evidence for a suppression of fungal activity, although fungal sporocarp production declined significantly, and the relative abundance of two fungal taxa changed dramatically with N fertilization. Microbial biomass as measured by chloroform fumigation did not respond to fertilization, nor did the ratio of fungi : bacteria as measured by quantitative polymerase chain reaction. However, microbial biomass C : N ratios narrowed significantly from 16.0 ± 1.4 to 5.2 ± 0.3 with fertilization. N fertilization significantly increased the activity of a cellulose‐degrading enzyme and suppressed the activities of protein‐ and chitin‐degrading enzymes but had no effect on soil respiration rates or ¹⁴C signatures. These results indicate that N fertilization alters microbial community composition and allocation to extracellular enzyme production without affecting soil respiration. Thus, our results do not provide evidence for strong microbial feedbacks to the boreal C cycle under climate warming or N addition. However, organic N cycling may decline due to a reduction in the activity of enzymes that target nitrogenous compounds.     

The effect of soil warming on bulk soil vs. rhizosphere respiration

There has been considerable debate on whether root/rhizosphere respiration or bulk soil respiration is more sensitive to long-term temperature changes. We investigated the response of belowground respiration to soil warming by 3 °C above ambient in bare soil plots and plots planted with wheat and maize. Initially, belowground respiration responded more to the soil warming in bare soil plots than in planted plots. However, as the growing season progressed, a greater soil-warming response developed in the planted plots as the contribution of root/rhizosphere respiration to belowground respiration declined. A negative correlation was observed between the contribution of root/rhizosphere respiration to total belowground respiration and the magnitude of the soil-warming response indicating that bulk soil respiration is more temperature sensitive than root/rhizosphere respiration. The dependence of root/rhizosphere respiration on substrate provision from photosynthesis is the most probable explanation for the observed lower temperature sensitivity of root/rhizosphere respiration. At harvest in late September, final crop biomass did not differ between the two soil temperature treatments in either the maize or wheat plots. Postharvest, flux measurements during the winter months indicated that the response of belowground respiration to the soil-warming treatment increased in magnitude (response equated to a Q ₁₀ value of 5.7 compared with ∼2.3 during the growing season). However, it appeared that this response was partly caused by a strong indirect effect of soil warming. When measurements were made at a common temperature, belowground respiration remained higher in the warmed subplots suggesting soil warming had maintained a more active microbial community through the winter months. It is proposed that any changes in winter temperatures, resulting from global warming, could alter the sink strength of terrestrial ecosystems considerably.