邢唷��>� ��欹�%` ��wbjbjN郚� <€,�,�4�&U�� 2 BNBNBN8zN�Q|2 崠v嶲�FR"hRhRhR_Z�[DI[$T�V�V�V�V�V�V�$�hk��z�� 膆Z^_Z膆膆z�hRhR�麜&xxx膆X �hR hRT�x膆T�xxr竵T�J p�hR俀恆�#惿BNr��钄l!�l崠�VG�鈜�G�p�p�G� 剛dm[_�xgbT籨 m[m[m[z�z�竪dm[m[m[崠膆膆膆膆2 Dv �Z�#�/d2 v Z�/2 2 2 ��Soil microbial respiration in arctic soil does not acclimate to temperature Authors: Iain P. Hartley1,*, David W. Hopkins1,2, Mark H. Garnett3, Martin Sommerkorn4 and Philip A. Wookey1 Affiliations: 1 School of Biological and Environmental Sciences, 我要吃瓜, Stirling, FK9 4LA, UK 2 Scottish Crop Research Institute, Invergowrie, Dundee, DD2 5DA, UK 3 NERC Radiocarbon Laboratory, Scottish Enterprise Technology Park, Rankine Avenue, East Kilbride, Glasgow G75 0QF 4 Macaulay Institute, Craigiebuckler, Aberdeen, AB15 8QH, UK E-mails: Iain P. Hartley: HYPERLINK "mailto:i.p.hartley@stir.ac.uk" i.p.hartley@stir.ac.uk David W. Hopkins: HYPERLINK "mailto:d.w.hopkins@scri.ac.uk" d.w.hopkins@scri.ac.uk Mark H. Garnett: HYPERLINK "mailto:M.Garnett@nercrcl.gla.ac.uk" M.Garnett@nercrcl.gla.ac.uk Martin Sommerkorn: HYPERLINK "mailto:M.Sommerkorn@macaulay.ac.uk" M.Sommerkorn@macaulay.ac.uk Philip A. Wookey: HYPERLINK "mailto:philip.wookey@stir.ac.uk" philip.wookey@stir.ac.uk Running title: Thermal acclimation of microbial respiration Keywords: Acclimation, adaptation, arctic, carbon cycling, climate change, CO2, respiration, microbial community, soil, temperature Article type: Letter Number of words in abstract: 169 Number of words in article: 4497 Number of references: 50 Number of figures: 2 Number of tables: 0 Correspondence author (*): Iain P. Hartley, School of Biological and Environmental Sciences, 我要吃瓜, Stirling, FK9 4LA, UK; E-mail: HYPERLINK "mailto:i.p.hartley@stir.ac.uk" i.p.hartley@stir.ac.uk Tel: +44 1786 467757; Fax: +44 1786 467843 Abstract Warming-induced release of CO2 from the large carbon (C) stores present 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 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 respiration observed in warming experiments, then it must also subsequently increase rates post cooling. We demonstrate that thermal acclimation did not occur. Rather, over the following 90 days, cooling resulted in a further reduction in respiration which was only reversed by extended reexposure 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. Key words: Adaptation, acclimation, arctic, carbon cycling, climate change, CO2, respiration, microbial community, soil, temperature INTRODUCTION Rising global temperatures are likely to increase the rate of soil organic matter decomposition resulting in a substantial release of CO2 (Raich & Schlesinger 1992; Kirschbaum 1995), and this phenomenon has the potential to accelerate climate change by up to 40% (Cox et al. 2000). In fact, the importance of soil C-cycling is recognized in the updated IPCC scenarios (IPCC 2007). However, increasingly, ecologists are recognizing that in order to predict long-term trends in ecosystem C fluxes and biological feedbacks, greater emphasis needs to be placed on measuring potential acclimation and adaptation responses (Oechel et al. 2000; Enquist 2007). Critically, acclimation has the potential to reduce the projected soil-C losses associated with global warming (Luo et al. 2001). Respiratory thermal acclimation has been defined as 搕he subsequent adjustment in the rate of respiration to compensate for an initial change in temperature� (Atkin & Tjoelker 2003). When many plant species are exposed to higher temperatures for a prolonged period of time, physiological acclimation results in a reduction in respiration rates allowing for the maintenance of a positive C balance (Atkin & Tjoelker 2003). Similarly, thermal acclimation of respiration has been demonstrated for both ectomycorrhizal (Malcolm et al. 2008) and arbuscular mycorrhizal fungi in soils (Heinemeyer et al. 2006), and the fungal symbiont in lichens (Lange & Green 2005). Further, although cooling reduces respiration rates, prolonged exposure often results in a subsequent increase in plant respiration rates, allowing for the maintenance of critical metabolic processes (Armstrong et al. 2006). Many physiological modifications have been observed in microbial communities present at low temperatures which allow for continued growth (D扐mico et al. 2006), and this may suggest that there is potential for up-regulation of activity following extended exposure to the cold. In soils, although increased rates of respiration have been observed in many warming experiments (Rustad et al. 2001), the magnitude of the initial positive response to temperature often declines over time (Rustad et al. 2001; Eliasson et al. 2005). Because alterations in microbial community structure accompany soil warming in both the field (Zhang et al. 2005) and the laboratory (Zogg et al. 1997; Andrews et al. 2000; Pettersson & B邋th 2003; Pietik鋓nen et al. 2005), as well as in response to seasonal changes in temperature (Schadt et al. 2003; Lipson & Schmidt 2004; Wallenstein et al. 2007), the reduction in the initial positive response of soil respiration to warming may be the result of acclimation of microbial respiration (Luo et al. 2001; Balser et al. 2006; Luo 2007; Wan et al. 2007). Investigating temperature responses of soil respiration and microbial activity is complicated by the fact that the effect of experimental soil warming is confounded by the depletion of the most readily-decomposable soil C fractions. This could equally explain the reduction in respiration rates observed in long-term studies (Rustad et al. 2001; Eliasson et al. 2005). Consequently, the main evidence for thermal acclimation of soil microbial respiration remains questionable (Kirschbaum 2004; Eliasson et al. 2005; Knorr et al. 2005; Hartley et al., 2007b). Identifying the potential for thermal acclimation of microbial respiration in arctic regions is particularly important due to the high rates of global warming already being experienced at high latitudes (ACIA 2005), the general sensitivity of communities close to environmental extremes to changing conditions, and the large amounts of C stored in these systems (Post et al. 1982). In addition, substantial changes in microbial communities have been observed between seasons in tundra soils (Schadt et al. 2003; Lipson & Schmidt 2004; Wallenstein et al. 2007) raising the possibility of acclimation of microbial respiration in these systems. Accurate predictions of the long-term rates of C and nitrogen cycling in arctic soils, which in turn may determine total ecosystem C storage (Hobbie et al. 2000), plant productivity (van Wijk et al. 2005) and species composition (Weintraub & Schimel 2005), require a much greater understanding of microbial acclimation responses. Here we present the results from one of the first studies to investigate the effect of an extended period of cooling on microbial respiration, utilizing organic soils taken from a sub-arctic tundra heath system in northern Sweden. If thermal acclimation is responsible for the down-regulation of microbial activity observed at high temperatures, then microbial activity must be gradually up-regulated when temperatures are reduced. This is because, as a compensatory response, acclimation must be reversible; otherwise temporary exposure to higher temperatures would result in a permanent down-regulation of respiration, preventing the recovery of rates even when temperature have declined, for example between summer and winter. In support of this logic, changes in soil microbial community structure have been observed both when soil temperatures increase (Andrews et al. 2000; Lipson & Schmidt 2004) and decrease (Schadt et al. 2003; Monson et al. 2006), and the thermal optimum for the activity of key C-cycling enzymes has been to shown increase and decrease with seasonal changes in temperature (Fenner et al. 2005). Furthermore, thermal acclimation of plant respiration, in response to seasonal and experimental changes in temperature, is dynamic and reversible, occurring both in response to warming and cooling (Atkin & Tjoelker 2003; Atkin et al. 2005; Zaragoza-Castells et al. 2008). Therefore, the use of experimental cooling allowed us to minimize the confounding factor of warming-induced substrate depletion (substrate depletion will occur at a slightly faster rate in the control soils, but total carbon losses should be sufficiently small to avoid confounding the results) whilst still determining whether soil microbial respiration acclimates to temperature. We demonstrate that (i) soil microbial respiration does not acclimate to temperature, (ii) the short-term temperature sensitivity of respiration is unaltered by the prevailing temperature regime, and (iii) when soil temperatures were reduced for an extended period of time, changes in the microbial community resulted in a further decrease in the baseline rate of respiration, lowering rates of CO2 production beyond the instantaneous response to temperature. METHODS Soil sampling and incubation On 13th September 2006, twenty-six soil cores (68爉m diameter and 100爉m deep) were removed from an area of tundra heath above the tree-line (at an altitude of approximately 750爉), about 200爇m north of the Arctic Circle, near Abisko, northern Sweden (68o18�07拻N, 18o51�16拻E). The mean annual temperature at this site is 1oC with mean January and July temperatures of -12 and 11oC, respectively (van Wijk et al. 2005). The dominant plant species are ericaceous shrubs, mainly of the genera Vaccinum and Empetrum, with some dwarf birch (Betula nana L.) also present. The soils have an organic horizon of between approximately 5 and 20燾m deep (mean depth = 11燾m), overlying well-drained medium to coarse-grained till deposits with some large boulders and intermittent pockets of mineral soil. In this study, only the organic horizon was sampled. This soil is well-suited for investigating the long-term response of soil microbial respiration to changing temperatures because it contains a large amount of C, but does not experience waterlogging (except briefly during spring melt), and field conditions can thus be well replicated in the laboratory. Further, issues such as the mineral protection of SOM changing with temperature are avoided (Rasmussen et al. 2006). The soils were transported to the 我要吃瓜 using cooled air cargo. The water content of the soil was raised to water holding capacity (WHC) and samples were placed in an incubator (MIR-153, SANYO, Loughborough, UK) at 10oC (�1oC) for 110 days to allow respiration rates to stabilize as the most labile C pool was depleted and for the microbial community to adjust to this temperature. Sixteen cores were then transferred to a separate incubator (same make and model) set at 2oC (�1oC). Of these 16 cores, 10 were then maintained at 2oC for 90 days (high-low treatment), and the other 6 cores were returned to the 10oC incubator after 60 days at 2oC (the high-low-high treatment). The remaining 10 cores were maintained at 10oC for the whole 200-day incubation (constant high treatment). Soil samples were maintained at WHC throughout by frequent addition of distilled water. Data loggers (Tinytag� Plus, Gemini Data Loggers Ltd., Chichester, UK) connected to thermistor probes (PB-5001, Gemini Data Loggers Ltd., Chichester, UK) confirmed that the temperatures in the incubators remained stable. The incubation temperatures used are within the range regularly experienced by the soil during the growing season, and soil temperatures were not reduced below 0oC to avoid changes in substrate availability caused by the alterations in the proportion of liquid water present (Mikan et al. 2002; Monson et al. 2006) and freeze-thaw effects. Respiration measurements Respiration measurements were carried out using an infra-red gas analyzer (EGM-4, PP Systems, Hitchen, UK) connected to an incubation chamber (700 ml Lock & Lock� container, Hana Cobi Plastic Co Ltd., Seoul, Korea) in a closed loop configuration. The rate of CO2 accumulation in the headspace was logged every 1.6 seconds until a 35爌pm increase in CO2 concentration had occurred. Therefore, measurements were made close to ambient CO2 concentrations. Respiration rates were expressed as m餲�C�g C1�h1. Finally, at the end of the incubation, the short-term temperature sensitivity of respiration (between 2 and 10oC) in six replicates taken from the high-low and constant high treatments was measured. The samples were transferred to an incubator at 2oC, and one day later respiration rates were measured. The incubator temperature was then raised to 6oC and subsequently 10oC, before being reduced back to 6oC and then 2oC. The soils were maintained at each new temperature for approximately 24 hours. Mean respiration rates were calculated at each temperature to allow changes in baseline rates of respiration over the five-day experiment to be included in the Q10 calculation (Fang et al. 2005). Changes in baseline rates of respiration could have been caused by changes in soil moisture (although samples were watered each day), or growth of microbial biomass in the previously cooled soils (Monson et al. 2006). The aim of this temperature manipulation was to determine whether the direct or instantaneous response of respiration to temperature had been altered by the cooling treatment and, therefore, we wanted to account for any changes in baseline rates. Respiration rates were natural log transformed and plotted against temperature. Linear regressions were then used to calculate the slope (K) of the relationship and Q10 values calculated using Equation 1. Q10=爀10K Equation 1 Substrate-induced respiration At the end of the experiment, soil from all 26 samples was sieved through a 2爉m mesh, large root fragments were removed and sub-samples dried for moisture and C content (loss on ignition) determination. After all samples had been incubated at 10oC over-night, a solution containing 15 mg of glucose per gram of soil C was added to a 5爂 (fresh wt.) sub-sample of each soil, with the corresponding volume (1 cm3) of distilled water added to a further 5 g sub-sample. Total CO2 production after 24 hours at 10oC was measured using gas chromatography (Model 90-P, Varian Aerograph, Palo Alto, CA, USA). The difference between the two treatments was considered to represent substrate-induced respiration (SIR), which is considered to be proportional to the size of microbial biomass (Anderson & Domsch 1978). Statistics Statistical analyses were carried out using SPSS (SPSS Science, version 15, Birmingham, UK). Before cooling, one-way ANOVAs were used to determine whether there were any significant differences between the respiration rates of the soils in the different temperature treatment groups. Post-cooling, for the high-low and high-low-high samples, linear regressions were used to determine whether the respiration rates changed significantly over the following 60 days. After the high-low-high samples were returned to 10oC, repeated measures ANOVAs and paired ttests were used to determine whether there were significant differences between dates, both immediately before and after the cooling treatment was applied, and between the high-low-high and constant high treatments. At the end of the incubation, independent samples t-tests were used to determine whether the short-term temperature sensitivity of respiration differed significantly between the high-low and constant high soils, and paired t-tests were used to determine whether respiration rates differed between the increasing and decreasing phase of the manipulation. An independent samples t-test was used to determine whether the rate of SIR differed between samples that were at 10oC at the end of the experiment (as there was no significant difference between the two treatments, constant high and high-low-high soils were grouped together) compared with the soils that were at 2oC at the end of the incubation (the high-low soils). RESULTS Respiration rates Before cooling, there were no significant differences in respiration rates measured at 10oC between the soils in the three temperature treatments (P�=�0.622; Fig. 1a). On day 110, the high-low and high-low-high cores were cooled from 10oC to 2oC and the following day the respiration rates had declined by about 67%. Over the following 60 days, rather than an increase in the rate of respiration indicative of acclimation, respiration rates declined significantly by on average 28% (Fig. 1b). The effect of temperature manipulation on the rate of respiration can be expressed using Q10 functions (Equation 1): SHAPE \* MERGEFORMAT Equation 1 Where RT is the respiration rate at temperature (T), R0 is the respiration rate at 0oC and Q10 is the proportion change in the rate of respiration given a 10oC change in temperature. The equations corresponding to the mean effect of cooling for 1 and 60 days across both the high-low and high-low-high soils are as follows: SHAPE \* MERGEFORMAT Day 1 SHAPE \* MERGEFORMAT Day 60 The reduction in the baseline rate of respiration caused by the cooling treatment has increased the apparent temperature sensitivity of respiration by ~50% (i.e. Q10 values have increased from 4.01 to 6.06). However, in the high-low treatment, about 50 days after cooling, respiration rates stabilized with there being no significant subsequent change in rates between days 157 and 200 (linear regression: P�=�0.404; Fig. 1). In contrast, over the entire incubation period, the respiration rate of the constant high cores did not change significantly (linear regression: P�=�0.359) indicating that the gradual reduction in respiration rates only occurred when soil temperatures were reduced. These results demonstrate that sustained exposure to low temperatures amplified the negative effect of cooling on soil respiration rates. On day 171, the high-low-high cores were returned to 10oC and respiration rates increased by approximately 72%. However, this rate was significantly less than that measured on day 109, immediately before the temperature reduction (paired ttest: P�=�0.037; Fig.�1c). This indicated that the reduction in respiration rates observed at 2oC was still apparent when samples were returned to 10oC. Over the following 28 days (i.e. days 172-200) the respiration rate increased by approximately 22% with the rate measured on day 193 differing significantly from the rate measured on day 172 (P�=�0.028; Fig.�1c). Further, the increase in respiration rates during this period only occurred in the high-low-high samples and not in the constant high samples (P�=�0.026; Fig. 1c). Thus, extended exposure to 10oC was required for the respiration rates to recover to their pre-cooling levels. Temperature sensitivity of respiration At the end of the 200-day incubation period, the response of the constant high and high-low samples to short-term changes in temperature was investigated. Overall, respiration rates were highly temperature sensitive, but there was no significant difference between treatments (Fig.�2; P�=�0.149) suggesting that extended exposure to 2oC had not resulted in microbial respiration becoming more (or less) temperature sensitive. However, the response of respiration to the increasing phase of the temperature manipulation was significantly higher in the high-low soils than in the constant high soils (high-low: Q10�=�4.736�0.248; constant爃igh: Q10�=�3.959�0.189; P�=�0.032). This appeared to have been caused by a significant increase in the baseline rate of respiration in the high-low soils as demonstrated by significantly (or marginally significantly) higher rates of respiration on the declining phase of the temperature manipulation (Fig. 2; 6oC: P�=�0.053, 2oC: P�=�0.001). No corresponding significant increase in the rate of respiration was observed in the constant-high treatment. The Q10 values calculated for the declining phase of the manipulation were similar and not significantly different (high-low: Q10�=�3.859�0.214; constant爃igh: Q10�=�3.655�0.197; P�=�0.497). Substrate-induced respiration A significantly greater rate of SIR (measured at 10oC in all cases) was observed in the soil samples that were at 10oC at the end of the experiment compared to those that were at 2oC (t-test: P�=�0.027; 75.3 vs. 66.7 m餲�C�g1 soil C�h1). DISCUSSION Thermal acclimation Our soil-cooling experiment produced no evidence that microbial respiration acclimates to temperature. The length of incubation carried out in our experiment should have allowed for thermal acclimation of microbial respiration to occur given that changes in microbial communities have been observed between seasons in tundra soils (Schadt et al. 2003; Lipson & Schmidt 2004; Wallenstein et al. 2007), and in response to temperature changes in laboratory experiments of a similar duration (Pettersson & B邋th 2003). Therefore, our results provide support for the modeling studies (Kirschbaum 2004; Eliasson et al. 2005; Knorr et al. 2005) that have proposed that the decline in the initial positive response of soil respiration to increased temperatures in long-term warming studies is due to substrate depletion and not acclimation of microbial respiration. Unlike plants it appears that the respiration of free-living, heterotrophic soil microbes does not acclimate to temperature. This is perhaps not surprising given the fundamental differences that exist between autotrophic and heterotrophic organisms. Whilst physiological acclimation serves to maintain a positive C balance in plants when shifted to a higher growth temperature (Atkin & Tjoelker 2003), it is unclear what advantage microbes would gain from reduced activity once temperature constraints have been relaxed. Thermal acclimation has been observed in mycorrhizal fungi (Heinemeyer et al. 2006; Malcolm et al. 2008) and the fungal component of lichens (Lange & Green 2005), but the activity of these microbes is tightly linked to, and controlled by (Heinemeyer et al. 2006), the rate of photosynthesis in their symbiotic partners. As such, these organisms are not representative of free-living heterotrophic microbes in soils. Previously, it has been shown that the temperature sensitivity of microbial activity may increase in microbial communities adapted to low temperatures (Monson et al. 2006), and that it may be the temperature response rather than the baseline rate of respiration that changes when systems acclimate to temperature (Luo et al. 2001; Wan et al. 2007). However, we found little evidence for the microbial respiration being more temperature sensitive in the cooled soils. The apparent down-regulation of the temperature response, that was observed in previous studies (Luo et al. 2001; Wan et al. 2007), was based on changes in seasonal Q10s in intact plant-soil systems. These results could have been caused by seasonal changes in the contributions of roots versus soil microbes to total belowground respiration. Hartley et al. (2007a) demonstrated that rhizosphere respiration responded less to soil warming than microbial respiration in bare soil. As the contribution of the more temperature insensitive flux, rhizosphere respiration, is likely to be greatest during mid season, a time when soil temperatures are likely to be highest, this could explain the apparent reduction in the temperature sensitivity of respiration in warmed plots (i.e. differences between warmed and ambient plots are expected to be lowest during the time of year when rhizosphere respiration contributes the most to belowground respiration). Our results indicate that it is unlikely that the development of a microbial community which responds little to changes in temperature can explain the lower seasonal Q10s measured in the warmed plots in previous studies (Luo et al. 2001; Wan et al. 2007). In our study, by carrying out our measurements in the absence of a rhizosphere, we avoided the possibility of microbial responses being mediated through changes in plant activity. Adaptation enhancing a positive feedback Our study goes further than demonstrating that thermal acclimation does not occur in these sub-arctic soils. Exposure to low temperatures for an extended period reduced the rate of respiration beyond the initial short-term response (Fig. 1b) and, similarly, extended exposure to moderate temperatures resulted in an increase in activity beyond the instantaneous response to temperature (Fig. 1c). Further, as the rate of SIR (measured at 10oC in all cases) was significantly lower in the cooled soils, it appears the microbial community had been affected. Whether the lower SIR rate in the cooled soil was due to a reduction in microbial biomass per se or reflected a shift in microbial community structure is debatable. However, the results from our study suggest that the microbial community was altered by the cooling and that this resulted in a further reduction in respiration rates. Therefore, at the low to moderate temperatures experienced in many soils, such as the arctic soil investigated here, when global warming increases soil temperatures it seems probable that C losses will be enhanced by changes in microbial community functioning. In support of this suggestion, a soil-warming study demonstrated that, during winter months, microbial activity in warmed plots was higher than in control plots even when measurements were made at a common temperature; it was concluded that warming had produced a more active microbial community (Hartley et al. 2007a). Further, it has been demonstrated that the temperature optimum for the activity of key microbial enzymes in organic soils may shift with time of year (Fenner et al. 2005), and that thermal tolerances of bacterial community activity gradually change in response to temperature manipulations (Pettersson & B邋th 2003). Rather than a compensatory response, it appears that, in the longer term, changes in the microbial community may result in a further increase in activity as temperatures rise. Therefore, soil-C losses from cold environments, and during winter periods, are likely to be enhanced by climate change due to changes in soil microbial communities amplifying the instantaneous response to temperature. Here we return to the issue of terminology; the changes in the microbial community which resulted in the decreasing rate of respiration for the 60-day period after cooling, and the increase in the rate of respiration following warming of the high-low-high soils, should be termed adaptation as it almost certainly contains a genetic component. We reiterate that the term acclimation is probably never appropriate when referring to a change occurring at the level of the whole community. If a compensatory response is observed then perhaps the term 揷ompensatory adaptation� would be more appropriate. Previously, studies which have modeled mineralization kinetics based on the results of incubation studies have suggested that substrate pool sizes may increase at higher temperatures (MacDonald et al. 1995; Waldrop & Firestone 2004; Rasmussen et al. 2006). Molecules that decompose in reactions with large activation energies are likely to decompose especially slowly at low temperatures (Davidson & Janssens 2006; Hartley & Ineson 2008), but may become more available at increased temperatures, potentially explaining the increased pool sizes and shifts in substrate utilization patterns observed in these studies (e.g. Waldrop & Firestone 2004). Within this context, in the study presented here, the gradual reduction in respiration rates post-cooling may reflect a loss of the most labile pool of substrates which are most available to microbes at low temperatures. This may in turn have induced the changes in the microbial community that occurred (reflected by the reduction in SIR). On return to the warmer temperature, thermal constraints on substrate availability may have been relaxed and the microbes again adapted to their prevailing environment. This is just one potential explanation for the reduction in respiration rates that occurred post-cooling and the changes in the microbial community. However, it is clear that thermal acclimation of microbial respiration did not occur, and adaptive responses of soil microbes to increasing temperatures may accelerate decomposition rates, at least at the low to moderate temperatures experienced in many soils. Timescale of the response of microbial respiration to warming In light of the findings of this study we can perhaps consider three separate processes which may determine the rate of soil C losses from arctic soils over different timescales. Firstly, in agreement with the study of Mikan et al. (2002), we found a strong instantaneous response of microbial respiration to changes in temperature (Fig.�2). When changes in the baseline rate of respiration were accounted for it appeared that the temperature sensitivity of respiration was not affected by the thermal regime the microbes had experienced. Secondly, cooling reduced the baseline rate of respiration as the microbial community was altered by the new temperature, and this medium-term response to the temperature manipulation was reversible. It should be mentioned that there was some evidence of a faster response of the microbial community to the warming than the cooling treatment. It took almost 60 days for the full cooling effect to occur whilst rates had fully recovered within 30 days of warming in the high-low-high samples. In addition, there was some evidence of an almost immediate, partial up-regulation of the baseline rate of respiration in the high-low soils during the short-term temperature manipulation. Therefore, at a timescale of about 1 month, respiration rates are likely to increase in warmer arctic soils as changes in the microbial community result in an increase in the baseline rate. Thirdly, at the decadal time scale, there may be a change in both total SOM stocks as warming stimulates C loss, and also a change in the composition of SOM as substrate pools with shorter turnover times are preferentially lost (舋ren & Bosatta, 2002; Kirschbaum 2004; Eliasson et al. 2005; Knorr et al. 2005). These changes will result in a subsequent decline in the rates of microbial respiration. Finally, in situ, if higher decomposition rates increase soil nutrient availability (Schmidt et al. 2002; Pregitzer et al. 2008), increased plant productivity may partly or fully offset these C losses, and so determine the extent to which rates of microbial respiration decline. However, further research is required to estimate the importance of this potential feedback. CONCLUSION Compensatory thermal acclimation of soil microbial respiration did not occur in our experiment. Rather, the effect of temperature on microbial community functioning increased respiration rates beyond the instantaneous effect of temperature. This response may enhance substantially soil-C losses, at least at low to moderate temperatures. Taking into account the rapid rate of climate change predicted for high-latitude ecosystems, and the high temperature sensitivity of decomposition measured at low temperatures, the large C stores in arctic and alpine soils may be especially vulnerable. Given that they contain over 20% of soil C, increased decomposition in these ecosystems has the potential to accelerate climate change. Finally, our study highlights the need to consider not only the instantaneous responses of processes to changes in abiotic factors, but also any adaptive responses that may subsequently occur at the community or ecosystem level. 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Soil microbial responses to experimental warming and clipping in a tallgrass prairie. Glob. Change Biol., 11, 266-277. FIGURE LEGENDS Figure 1 The mean soil respiration rates in the three different temperature treatments (constant high �% , high-low " " " �%" " " , high-low-high �% ). Error bars represent �1SE (constant high and high-low: n�=�10; high-low-high: n�=�6). The main panel (a) shows the whole of the incubation period during which respiration measurements were made. The timing of the reduction in temperature from 10oC to 2oC in the high-low and high-low-high treatments is indicated as is the subsequent return to 10oC in the high-low-high treatment. Panels (b) and (c) highlight the periods of key interest. Panel (b) shows the decline in the rate of respiration at 2oC over the first 60 days at the lower incubation temperature in the high-low and high-low-high treatments. Linear regressions are fitted to each temperature treatment separately although there is no significant difference between the two fitted lines (high-low (dotted line): y�=�0.0112x�+�4.00, R2�=�0.817; high-low-high (dashed line): y�=�0.0135x�+�4.36, R2�=�0.815). Panel (c) shows the rate of respiration at 10oC in the high-low-high and constant high samples immediately after the high-low-high samples were returned to 10oC. The horizontal dashed line indicates the mean rate of respiration in the high-low-high samples on day 109 immediately before the high-low-high samples were transferred to 2oC. Initially the rate of respiration in the high-low-high samples was significantly less than on day 109 (paired t-test: P�=�0.037) and significantly lower than in the constant high treatment (ttest: P�=�0.044), but these differences were subsequently lost as the respiration rates in the high-low-high samples increased. A significant interaction term between time and temperature treatment (repeated measures ANOVA; P�=�0.026) indicated that the increase in respiration rates only occurred in the high-low-high samples. Figure 2 The response of respiration to the short-term changes in temperature in the high-low and constant high samples. Mean respiration rates on both the increasing and decreasing phase of the temperature manipulation are shown. Error bars represent +1SE (n�=�6). 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