全球变暖背景下土壤微生物呼吸的热适应性:证据、机理和争议

土壤微生物呼吸的热适应性被认为是决定陆地生态系统对全球变暖反馈作用的潜在重要机制,可能显著改变未来的气候变化趋势,然而学术界对于这一机制是否真实存在尚有分歧。阐述了土壤微生物呼吸的热适应性概念,从证据、机理和争议3方面对已有研究进展进行了综述和分析。土壤微生物呼吸的热适应性是微生物在群落尺度上对温度变化的适应性,具有坚实的生物学与生态学理论基础,研究者们运用各类指标已在许多实验中证实土壤微生物物种及群落的呼吸过程能够在高温环境产生适应性变化。土壤微生物呼吸的热适应性机理涉及生物膜结构变化、酶活性变化、微生物碳分配比例变化和微生物群落结构变化等方面。关于土壤微生物呼吸热适应性的争议可能是由研究方法、微生物物种及环境条件的差异引起的。根据对已有研究的分析,认为土壤微生物呼吸的热适应性是真实存在的,未来的研究可进一步探索土壤微生物呼吸的热适应性机理,深入研究环境和全球变化对土壤微生物呼吸的热适应性影响,定量评估土壤微生物呼吸的热适应性对陆地生态系统反馈过程的影响。     

Carbon losses due to soil warming: Do autotrophic and heterotrophic soil respiration respond equally?

Global warming has the potential to increase soil respiration (R_S), one of the major fluxes in the global carbon (C) cycle. R_S consists of an autotrophic (R_A) and a heterotrophic (R_H) component. We combined a soil warming experiment with a trenching experiment to assess how R_S, R_A, and R_H are affected. The experiment was conducted in a mature forest dominated by Norway spruce. The site is located in the Austrian Alps on dolomitic bedrock. We warmed the soil of undisturbed and trenched plots by means of heating cables 4 °C above ambient during the snow‐free seasons of 2005 and 2006. Soil warming increased the CO₂ efflux from control plots (R_S) by ∼45% during 2005 and ∼47% during 2006. The CO₂ efflux from trenched plots (R_H) increased by ∼39% during 2005 and ∼45% during 2006. Similar responses of R_S and R_H indicated that the autotrophic and heterotrophic components of R_S responded equally to the temperature increase. Thirty‐five to forty percent or 1 t C ha⁻¹ yr⁻¹ of the overall annual increase in R_S (2.8 t C ha⁻¹ yr⁻¹) was autotrophic. The remaining, heterotrophic part of soil respiration (1.8 t C ha⁻¹ yr⁻¹), represented the warming‐induced C loss from the soil. The autotrophic component showed a distinct seasonal pattern. Contribution of R_A to R_S was highest during summer. Seasonally derived Q₁₀ values reflected this pattern and were correspondingly high (5.3–9.3). The autotrophic CO₂ efflux increase due to the 4 °C warming implied a Q₁₀ of 2.9. Hence, seasonally derived Q₁₀ of R_A did not solely reflect the seasonal soil temperature development.     

Reduced carbon use efficiency and increased microbial turnover with soil warming

Global soil carbon (C) stocks are expected to decline with warming, and changes in microbial processes are key to this projection. However, warming responses of critical microbial parameters such as carbon use efficiency (CUE) and biomass turnover (rB) are not well understood. Here, we determine these parameters using a probabilistic inversion approach that integrates a microbial‐enzyme model with 22 years of carbon cycling measurements at Harvard Forest. We find that increasing temperature reduces CUE but increases rB, and that two decades of soil warming increases the temperature sensitivities of CUE and rB. These temperature sensitivities, which are derived from decades‐long field observations, contrast with values obtained from short‐term laboratory experiments. We also show that long‐term soil C flux and pool changes in response to warming are more dependent on the temperature sensitivity of CUE than that of rB. Using the inversion‐derived parameters, we project that chronic soil warming at Harvard Forest over six decades will result in soil C gain of <1.0% on average (1st and 3rd quartiles: 3.0% loss and 10.5% gain) in the surface mineral horizon. Our results demonstrate that estimates of temperature sensitivity of microbial CUE and rB can be obtained and evaluated rigorously by integrating multidecadal datasets. This approach can potentially be applied in broader spatiotemporal scales to improve long‐term projections of soil C feedbacks to climate warming.     

土壤呼吸对温度升高的适应

土壤呼吸是陆地生态系统碳循环的重要环节之一 ,其对温度升高的敏感程度在相当大的程度上决定着全球气候变化与碳循环之间的反馈关系。土壤呼吸对温度升高的适应是个比较普遍的现象 ,其表现形式主要为随着温度的持续升高和升温时间的延长 ,土壤呼吸对温度升高反应的敏感程度下降。产生这一现象的机制包括影响因子主导地位的转移和温度以外其他因子的协同变化。土壤呼吸对温度升高的适应可以视为碳循环对全球变暖的负反馈效应 ,它可能会在一定程度上缓和陆地生态系统对全球气候系统之间的耦合作用 ,并且导致土壤呼吸对全球温度升高响应的时空差异。由于目前生态系统模型多数没有考虑土壤呼吸的对温度升高的适应性 ,而采用统一的 Q1 0 值 ,其对未来土壤呼吸和未来气候变化幅度的预测可能存在偏差     

Allometric constraints on,and trade‐offs in,belowground carbon allocation and their control of soil respiration across global forest ecosystems

To fully understand how soil respiration is partitioned among its component fluxes and responds to climate, it is essential to relate it to belowground carbon allocation, the ultimate carbon source for soil respiration. This remains one of the largest gaps in knowledge of terrestrial carbon cycling. Here, we synthesize data on gross and net primary production and their components, and soil respiration and its components, from a global forest database, to determine mechanisms governing belowground carbon allocation and their relationship with soil respiration partitioning and soil respiration responses to climatic factors across global forest ecosystems. Our results revealed that there are three independent mechanisms controlling belowground carbon allocation and which influence soil respiration and its partitioning: an allometric constraint; a fine‐root production vs. root respiration trade‐off; and an above‐ vs. belowground trade‐off in plant carbon. Global patterns in soil respiration and its partitioning are constrained primarily by the allometric allocation, which explains some of the previously ambiguous results reported in the literature. Responses of soil respiration and its components to mean annual temperature, precipitation, and nitrogen deposition can be mediated by changes in belowground carbon allocation. Soil respiration responds to mean annual temperature overwhelmingly through an increasing belowground carbon input as a result of extending total day length of growing season, but not by temperature‐driven acceleration of soil carbon decomposition, which argues against the possibility of a strong positive feedback between global warming and soil carbon loss. Different nitrogen loads can trigger distinct belowground carbon allocation mechanisms, which are responsible for different responses of soil respiration to nitrogen addition that have been observed. These results provide new insights into belowground carbon allocation, partitioning of soil respiration, and its responses to climate in forest ecosystems and are, therefore, valuable for terrestrial carbon simulations and projections.     

Thermal adaptation of heterotrophic soil respiration in laboratory microcosms

Respiration of heterotrophic microorganisms decomposing soil organic carbon releases carbon dioxide from soils to the atmosphere. In the short term, soil microbial respiration is strongly dependent on temperature. In the long term, the response of heterotrophic soil respiration to temperature is uncertain. However, following established evolutionary trade‐offs, mass‐specific respiration (R_mass) rates of heterotrophic soil microbes should decrease in response to sustained increases in temperature (and vice‐versa). Using a laboratory microcosm approach, we tested the potential for the R_mass of the microbial biomass in six different soils to adapt to three, experimentally imposed, thermal regimes (constant 10, 20 or 30 °C). To determine R_mass rates of the heterotrophic soil microbial biomass across the temperature range of the imposed thermal regimes, we periodically assayed soil subsamples using similar approaches to those used in plant, animal and microbial thermal adaptation studies. As would be expected given trade‐offs between maximum catalytic rates and the stability of the binding structure of enzymes, after 77 days of incubation R_mass rates across the range of assay temperatures were greatest for the 10 °C experimentally incubated soils and lowest for the 30 °C soils, with the 20 °C incubated soils intermediate. The relative magnitude of the difference in R_mass rates between the different incubation temperature treatments was unaffected by assay temperature, suggesting that maximum activities and not Q₁₀ were the characteristics involved in thermal adaptation. The time taken for changes in R_mass to manifest (77 days) suggests they likely resulted from population or species shifts during the experimental incubations; we discuss alternate mechanistic explanations for those results we observed. A future research priority is to evaluate the role that thermal adaptation plays in regulating heterotrophic respiration rates from field soils in response to changing temperature, whether seasonally or through climate change.     

Thermal adaptation of microbial respiration persists throughout long-term soil carbon decomposition

Soil microbial respiration is expected to show adaptations to changing temperatures, greatly weakening the magnitude of feedback over time, as shown in labile carbon substrates. However, whether such thermal adaptation persists during long-term soil carbon decomposition as carbon substrates decrease in decomposability remains unknown. Here, we conducted a 6-year incubation experiment in natural and arable soils with distinct properties under three temperatures (10, 20 and 30°C). Mass-specific microbial respiration was consistently lower under higher long-term incubation temperatures, suggesting the occurrence and persistence of microbial thermal adaptation in long-term soil carbon decomposition. Furthermore, changes in microbial community composition and function largely explained the persistence of microbial respiratory thermal adaptation. If such thermal adaptation generally occurs in large low-decomposability carbon pools, warming-induced soil carbon losses may be lower than previously predicted and thus may not contribute as much as expected to greenhouse warming.     

Rhizospheric and heterotrophic components of soil respiration in six Chinese temperate forests

Partitioning soil respiration (R_S) into heterotrophic (R_H) and rhizospheric (R_R) components is an important step for understanding and modeling carbon cycling in forest ecosystems, but few studies on R_R and R_H exist in Chinese temperate forests. In this study, we used a trenching plot approach to partition R_S in six temperate forests in northeastern China. Our specific objectives were to (1) examine seasonal patterns of soil surface CO₂ fluxes from trenched (R_T) and untrenched plots (R_UT) of these forests; (2) quantify annual fluxes of R_S components and their relative contributions in the forest ecosystems; and (3) examine effects of plot trenching on measurements of R_S and related environmental factors. The R_T maximized in early growing season, but the difference between R_UT and R_T peaked in later summer. The annual fluxes of R_H and R_R varied with forest types. The estimated values of R_H for the Korean pine (Pinus koraiensis Sieb. et Zucc.), Dahurian larch (Larix gmelinii Rupr.), aspen‐birch (Populous davidiana Dode and Betula platyphylla Suk.), hardwood (Fraxinus mandshurica Rupr., Juglans mandshurica Maxim. and Phellodendron amurense Rupr.), Mongolian oak (Quercus mongolica Fisch.) and mixed deciduous (no dominant tree species) forests averaged 89, 196, 187, 245, 261 and 301 g C m⁻² yr⁻¹, respectively; those of R_R averaged 424, 209, 628, 538, 524 and 483 g C m⁻² yr⁻¹, correspondingly; calculated contribution of R_R to R_S (RC) varied from 52% in the larch forest to 83% in the pine forest. The annual flux of R_R was strongly correlated to biomass of roots <0.5 cm in diameter, while that of R_H was weakly correlated to soil organic carbon concentration at A horizon. We concluded that vegetation type and associated carbon metabolisms of temperate forests should be considered in assessing and modeling R_S components. The significant impacts of changed soil physical environments and substrate availability by plot trenching should be appropriately tackled in analyzing and interpreting measurements of R_S components.     

Ocean warming enhances respiration and carbon demand of coastal microbial plankton

The increase of anthropogenic CO₂ during this century is expected to cause warming of large regions of the ocean. Microbes lead the biological role in the CO₂ balance of marine ecosystems, their activity is known to be influenced by temperature, and it is important to constrain and quantify these effects on bacterial carbon use. Furthermore, if warming were to enhance the carbon demand (production + respiration) of planktonic microbes but would maintain their efficiency low (as it generally is), then most carbon consumed would end up respired. We designed a strategy in which we measured bacterial production and respiration throughout a seasonal cycle in a coastal Mediterranean site, and determined experimentally the effects of ca. 2.5 °C on these processes. We show that warming will increase nearly 20% the total carbon demand of coastal microbial plankton without any effect on their (commonly low) growth efficiency, which could generate a positive feedback between coastal warming and CO₂ production.     

The response of heterotrophic CO2 flux to soil warming

In a forest ecosystem at steady state, net carbon (C) assimilation by plants and C loss through soil and litter decomposition by heterotrophic organisms are balanced. However, a perturbation to the system, such as increased mean soil temperature, will lead to faster decay, enhancing CO₂ release from decomposers, and thus upsetting the balance. Recent in situ experiments have indicated that the stimulation of soil respiration following a step increase in annual average soil temperature declines over time. One possible explanation for this decline may be changes in substrate availability. This hypothesis is examined by using the ecosystem model G'DAY, which simulates C and nitrogen (N) dynamics in plants and soil. We applied the model to observations from a soil‐warming experiment in a Norway spruce (Picea abies (L.) Karst.) stand by simulating a step increase of soil temperature. The model provided a good qualitative reproduction of the observed reduction of heterotrophic respiration (R_h) under sustained warming. The simulations showed how the combined effects of faster turnover and reduced substrate availability lead to a transient increase of R_h. The simulated annual increase in R_h from soil was 60% in the first year after perturbation but decreased to 30% after a decade. One conclusion from the analysis of the simulations is that R_h can decrease even though the temperature response function for decomposition remains unchanged. G'DAY suggests that acclimation of R_h to soil warming is partly an effect of substrate depletion of labile C pools during the first decade of warming as a result of accelerated rates of mineralization. The response is attributed mainly to changing levels of C in pools with short time constants, reflecting the importance of high‐quality soil C fractions. Changes of the structure or physiology of the decomposer community were not invoked. Therefore, it becomes a question of definition whether the simulated dynamics of the declining response of CO₂ release to the warming should be named acclimation or seen as a natural part of the system dynamics.     

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.     

No overall stimulation of soil respiration under mature deciduous forest trees after 7 years of CO2 enrichment

The anthropogenic rise in atmospheric CO₂ is expected to impact carbon (C) fluxes not only at ecosystem level but also at the global scale by altering C cycle processes in soils. At the Swiss Canopy Crane (SCC), we examined how 7 years of free air CO₂ enrichment (FACE) affected soil CO₂ dynamics in a ca. 100‐year‐old mixed deciduous forest. The use of ¹³C‐depleted CO₂ for canopy enrichment allowed us to trace the flow of recently fixed C. In the 7th year of growth at ～550 ppm CO₂, soil respiratory CO₂ consisted of 39% labelled C. During the growing season, soil air CO₂ concentration was significantly enhanced under CO₂‐exposed trees. However, elevated CO₂ failed to stimulate cumulative soil respiration (R_s) over the growing season. We found periodic reductions as well as increases in instantaneous rates of R_s in response to elevated CO₂, depending on soil temperature and soil volumetric water content (VWC; significant three‐way interaction). During wet periods, soil water savings under CO₂‐enriched trees led to excessive VWC (>45%) that suppressed R_s. Elevated CO₂ stimulated R_s only when VWC was ≤40% and concurrent soil temperature was high (>15 °C). Seasonal Q₁₀ estimates of R_s were significantly lower under elevated (Q₁₀=3.30) compared with ambient CO₂ (Q₁₀=3.97). However, this effect disappeared when three consecutive sampling dates of extremely high VWC were disregarded. This suggests that elevated CO₂ affected Q₁₀ mainly indirectly through changes in VWC. Fine root respiration did not differ significantly between treatments but soil microbial biomass (C_mic) increased by 14% under elevated CO₂ (marginally significant). Our findings do not indicate enhanced soil C emissions in such stands under future atmospheric CO₂. It remains to be shown whether C losses via leaching of dissolved organic or inorganic C (DOC, DIC) help to balance the C budget in this forest.     

Contrasting responses of heterotrophic and autotrophic respiration to experimental warming in a winter annual‐dominated prairie

Understanding how soil respiration (Rs) and its source components respond to climate warming is crucial to improve model prediction of climate‐carbon (C) feedback. We conducted a manipulation experiment by warming and clipping in a prairie dominated by invasive winter annual Bromus japonicas in Southern Great Plains, USA. Infrared radiators were used to simulate climate warming by 3 °C and clipping was used to mimic yearly hay mowing. Heterotrophic respiration (Rh) was measured inside deep collars (70 cm deep) that excluded root growth, while total soil respiration (Rs) was measured inside surface collars (2–3 cm deep). Autotrophic respiration (Ra) was calculated by subtracting Rh from Rs. During 3 years of experiment from January 2010 to December 2012, warming had no significant effect on Rs. The neutral response of Rs to warming was due to compensatory effects of warming on Rh and Ra. Warming significantly (P < 0.05) stimulated Rh but decreased Ra. Clipping only marginally (P < 0.1) increased Ra in 2010 but had no effect on Rh. There were no significant interactive effects of warming and clipping on Rs or its components. Warming stimulated annual Rh by 22.0%, but decreased annual Ra by 29.0% across the 3 years. The decreased Ra was primarily associated with the warming‐induced decline of the winter annual productivity. Across the 3 years, warming increased Rh/Rs by 29.1% but clipping did not affect Rh/Rs. Our study highlights that climate warming may have contrasting effects on Rh and Ra in association with responses of plant productivity to warming.     

Decreased mass specific respiration under experimental warming is robust to the microbial biomass method employed

Hartley et al. question whether reduction in R _mass, under experimental warming, arises because of the biomass method. We show the method they treat as independent yields the same result. We describe why the substrate-depletion hypothesis may not solely explain observed responses, and urge caution in interpretation of the seasonal data.     

模拟增温对土壤呼吸影响机制的研究进展与展望

土壤呼吸是土壤碳库向大气碳库输入的主要途径,而温度升高会影响土壤呼吸从而改变全球碳平衡.据预测在21世纪末,全球平均地表温度将升高0.3～4.8 ℃,因此野外自然条件下的模拟增温试验对土壤呼吸的影响是全球变化研究的热点之一.本文综述了不同时空格局下土壤呼吸对模拟增温的响应特征,指出短期增温能提高土壤呼吸,而长期增温下无统一规律,并且不同生态系统之间也存在差异;重点讨论了模拟增温对土壤呼吸的影响机制,指出增温能直接影响土壤呼吸,同时增温也能通过影响土壤水分、盐分、土壤理化性质等环境因子以及光合作用、凋落物等生物因子对土壤呼吸产生间接影响;另外,分析了土壤呼吸对增温产生适应性的形成机制,主要包括微生物、根、酶的温度适应性、水分限制、氮素过量以及呼吸底物限制.在此基础上对今后的研究方向加以展望:加强根际微生态系统的研究;重点研究不对称增温下土壤呼吸的特征及机制;关注典型物候期和不同季节典型天气土壤呼吸的测定;构建土壤呼吸响应模拟增温试验的研究网络,进行联网试验.     

Responses of dryland soil respiration and soil carbon pool size to abrupt vs. gradual and individual vs. combined changes in soil temperature, precipitation, and atmospheric [CO2]: a simulation analysis

With the large extent and great amount of soil carbon (C) storage, drylands play an important role in terrestrial C balance and feedbacks to climate change. Yet, how dryland soils respond to gradual and concomitant changes in multiple global change drivers [e.g., temperature (T_s), precipitation (Ppt), and atmospheric [CO₂] (CO₂)] has rarely been studied. We used a process‐based ecosystem model patch arid land simulator to simulate dryland soil respiration (R_s) and C pool size (C_s) changes to abrupt vs. gradual and single vs. combined alterations in T_s, Ppt and CO₂ at multiple treatment levels. Results showed that abrupt perturbations generally resulted in larger R_s and had longer differentiated impacts than did gradual perturbations. R_s was stimulated by increases in T_s, Ppt, and CO₂ in a nonlinear fashion (e.g., parabolically or asymptotically) but suppressed by Ppt reduction. Warming mainly stimulated heterotrophic R_s (i.e., R_h) whereas Ppt and CO₂ influenced autotrophic R_s (i.e., R_a). The combined effects of warming, Ppt, and CO₂ were nonadditive of primary single‐factor effects as a result of substantial interactions among these factors. Warming amplified the effects of both Ppt addition and CO₂ elevation whereas Ppt addition and CO₂ elevation counteracted with each other. Precipitation reduction either magnified or suppressed warming and CO₂ effects, depending on the magnitude of factor's alteration and the components of R_s (R_a or R_h) being examined. Overall, Ppt had dominant influence on dryland R_s and C_s over T_s and CO₂. Increasing Ppt individually or in combination with T_s and CO₂ benefited soil C sequestration. We therefore suggested that global change experimental studies for dryland ecosystems should focus more on the effects of precipitation regime changes and the combined effects of Ppt with other global change factors (e.g., T_s, CO₂, and N deposition).     

Organic carbon cycling in Taylor Valley, Antarctica: quantifying soil reservoirs and soil respiration

Organic carbon reservoirs and respiration rates in soils have been calculated for most major biomes on Earth revealing patterns related to temperature, precipitation, and location. Yet data from one of the Earth's coldest, driest, and most southerly soil ecosystems, that of the McMurdo Dry Valleys of Antarctica, are currently not a part of this global database. In this paper, we present the first regional calculations of the soil organic carbon reservoirs in a dry valley ecosystem (Taylor Valley) and report measurements of CO₂ efflux from Antarctic soils. Our analyses indicate that, despite the absence of visible accumulations of organic matter in most of Taylor Valley's arid soils, this soil environment contained a significant percentage (up to 72%) of the seasonally unfrozen organic carbon reservoir in the terrestrial ecosystem. Field measurements of soil CO₂‐efflux in Taylor Valley soils were used to evaluate biotic respiration and averaged 0.10 ± 0.08 μmol CO₂ m^?2 s^?1. Laboratory soil microcosms suggested that this respiration rate was sensitive to increases in temperature, moisture, and carbon addition. Finally, a steady‐state calculation of the mean residence time for organic carbon in Taylor Valley soils was 23 years. Because this value contradicts all that is currently known about carbon cycling rates in the dry valleys, we suggest that the dry valley soil carbon dynamics is not steady state. Instead, we suggest that the dynamic is complex, with at least two (short‐ and long‐term) organic carbon reservoirs. We also suggest that organic carbon in the dry valley soil environment may be more important, and play a more active role in long‐term ecosystem processes, than previously believed.     

Separating root and soil microbial contributions to soil respiration: A review of methods and observations

Forest soil respiration is the sum of heterotrophic (microbes, soil fauna) and autotrophic (root) respiration. The contribution of each group needs to be understood to evaluate implications of environmental change on soil carbon cycling and sequestration. Three primary methods have been used to distinguish hetero- versus autotrophic soil respiration including: integration of components contributing to in situ forest soil CO₂ efflux (i.e., litter, roots, soil), comparison of soils with and without root exclusion, and application of stable or radioactive isotope methods. Each approach has advantages and disadvantages, but isotope based methods provide quantitative answers with the least amount of disturbance to the soil and roots. Published data from all methods indicate that root/rhizosphere respiration can account for as little as 10 percent to greater than 90 percent of total in situ soil respiration depending on vegetation type and season of the year. Studies which have integrated percent root contribution to total soil respiration throughout an entire year or growing season show mean values of 45.8 and 60.4 percent for forest and nonforest vegetation, respectively. Such average annual values must be extrapolated with caution, however, because the root contribution to total soil respiration is commonly higher during the growing season and lower during the dormant periods of the year.     

Differential responses of carbon‐degrading enzyme activities to warming: Implications for soil respiration

Extracellular enzymes catalyze rate‐limiting steps in soil organic matter decomposition, and their activities (EEAs) play a key role in determining soil respiration (SR). Both EEAs and SR are highly sensitive to temperature, but their responses to climate warming remain poorly understood. Here, we present a meta‐analysis on the response of soil cellulase and ligninase activities and SR to warming, synthesizing data from 56 studies. We found that warming significantly enhanced ligninase activity by 21.4% but had no effect on cellulase activity. Increases in ligninase activity were positively correlated with changes in SR, while no such relationship was found for cellulase. The warming response of ligninase activity was more closely related to the responses of SR than a wide range of environmental and experimental methodological factors. Furthermore, warming effects on ligninase activity increased with experiment duration. These results suggest that soil microorganisms sustain long‐term increases in SR with warming by gradually increasing the degradation of the recalcitrant carbon pool.     

森林土壤融化期异养呼吸和微生物碳变化特征

采用室内土柱培养的方法,研究在不同湿度(55%和80%WFPS,土壤充水孔隙度)和不同氮素供给(NH_4Cl和KNO_3,4.5 g N/m~2)条件下,外源碳添加(葡萄糖,6.4 g C/m~2)对温带成熟阔叶红松混交林和次生白桦林土壤融化过程微生物呼吸和微生物碳的激发效应。结果表明:在整个融化培养期间,次生白桦林土壤对照CO_2累积排放量显著高于阔叶红松混交林土壤。随着土壤湿度的增加,次生白桦林土壤对照CO_2累积排放量和微生物代谢熵(q_(CO_2))显著降低,而阔叶红松混交林土壤两者显著地增加(P0.05)。两种林分土壤由葡萄糖(Glu)引起的CO_2累积排放量(9.61—13.49 g C/m~2)显著大于实验施加的葡萄糖含碳量(6.4g C/m~2),同时由Glu引起的土壤微生物碳增量为3.65—27.18 g C/m~2,而施加Glu对土壤DOC含量影响较小。因此,这种由施加Glu引起的额外碳释放可能来源于土壤固有有机碳分解。融化培养结束时,阔叶红松混交林土壤未施氮处理由Glu引起的CO_2累积排放量在两种湿度条件下均显著大于次生白桦林土壤(P0.001);随着湿度的增加,两种林分土壤Glu引起的CO_2累积排放量显著增大(P0.001)。单施KNO_3显著地增加两种湿度的次生白桦林土壤Glu引起的CO_2累积排放量(P0.01)。单施KNO_3显著地增加了两种湿度次生白桦林土壤Glu引起的微生物碳(P0.001),单施NH_4Cl显著地增加低湿度阔叶红松混交林土壤Glu引起的微生物碳(P0.001)。结合前期报道的未冻结实验结果,发现冻结过程显著地影响外源Glu对温带森林土壤微生物呼吸和微生物碳的刺激效应(P0.05),并且无论冻结与否,温带森林土壤微生物呼吸和微生物碳对外源Glu的响应均与植被类型、土壤湿度、外源氮供给及其形态存在显著的相关性。