Thermodynamic theory explains the temperature optima of soil microbial processes and high Q10 values at low temperatures

Our current understanding of the temperature response of biological processes in soil is based on the Arrhenius equation. This predicts an exponential increase in rate as temperature rises, whereas in the laboratory and in the field, there is always a clearly identifiable temperature optimum for all microbial processes. In the laboratory, this has been explained by denaturation of enzymes at higher temperatures, and in the field, the availability of substrates and water is often cited as critical factors. Recently, we have shown that temperature optima for enzymes and microbial growth occur in the absence of denaturation and that this is a consequence of the unusual heat capacity changes associated with enzymes. We have called this macromolecular rate theory – MMRT (Hobbs et al., 2013 , ACS Chem. Biol. 8:2388). Here, we apply MMRT to a wide range of literature data on the response of soil microbial processes to temperature with a focus on respiration but also including different soil enzyme activities, nitrogen and methane cycling. Our theory agrees closely with a wide range of experimental data and predicts temperature optima for these microbial processes. MMRT also predicted high relative temperature sensitivity (as assessed by Q₁₀ calculations) at low temperatures and that Q₁₀ declined as temperature increases in agreement with data synthesis from the literature. Declining Q₁₀ and temperature optima in soils are coherently explained by MMRT which is based on thermodynamics and heat capacity changes for enzyme‐catalysed rates. MMRT also provides a new perspective, and makes new predictions, regarding the absolute temperature sensitivity of ecosystems – a fundamental component of models for climate change.     

Rapid laboratory measurement of the temperature dependence of soil respiration and application to changes in three diverse soils through the year

Determining the temperature dependence of soil respiration is needed to test predictive models such as Arrhenius-like functions and macro-molecular rate theory (MMRT). We tested a method for rapid measurement of respiration using a temperature gradient block, cooled at one end (~2 °C) and heated at the other (~50 °C) that accommodated 44 tubes containing soil incubated at roughly 1 °C increments. Gas samples were taken after 5 h incubation and analysed for CO₂. The temperature gradient block allowed rapid assessment of temperature dependence of soil respiration with the precision needed to test models and explore existing theories of how temperature and moisture interact to control biochemical processes. Temperature response curves were well fitted by MMRT and allowed calculation of the temperature at which absolute temperature sensitivity was maximal (T_inf). We measured temperature response of three soils at seven moisture contents and showed that the absolute rate and sensitivity of respiration was partly dependent on adjusted moisture content. This result implied that comparisons between soils need to be made at a common moisture content. We also measured potential changes in the temperature dependence (and sensitivity) of respiration for three different soils collected at one site throughout a year. T_inf ranged from 43 to 51 °C for the three soils. T_inf and temperature sensitivity were not dependent on soil type collected but was partly dependent on time of year of collection. Temporal changes in temperature response suggested that the microbial communities may tune their metabolisms in response to changes in soil temperatures.     

The Response of Soil Processes to Climate Change: Results from Manipulation Studies of Shrublands Across an Environmental Gradient

Predicted changes in climate may affect key soil processes such as respiration and net nitrogen (N) mineralization and thus key ecosystem functions such as carbon (C) storage and nutrient availability. To identify the sensitivity of shrubland soils to predicted climate changes, we have carried out experimental manipulations involving ecosystem warming and prolonged summer drought in ericaceous shrublands across a European climate gradient. We used retractable covers to create artificial nighttime warming and prolonged summer drought to 20-m² experimental plots. Combining the data from across the environmental gradient with the results from the manipulation experiments provides evidence for strong climate controls on soil respiration, net N mineralization and nitrification, and litter decomposition. Trends of 0%–19% increases of soil respiration in response to warming and decreases of 3%–29% in response to drought were observed. Across the environmental gradient and below soil temperatures of 20°C at a depth of 5–10 cm, a mean Q₁₀ of 4.1 in respiration rates was observed although this varied from 2.4 to 7.0 between sites. Highest Q₁₀ values were observed in Spain and the UK and were therefore not correlated with soil temperature. A trend of increased accumulated surface litter mass loss was observed with experimental warming (2%– 22%) but there was no consistent response to experimental drought. In contrast to soil respiration and decomposition, variability in net N mineralization was best explained by soil moisture rather than temperature. When water was neither limiting or in excess, a Q₁₀ of 1.5 was observed for net N mineralization rates. These data suggest that key soil processes will be differentially affected by predicted changes in rainfall pattern and temperature and the net effect on ecosystem functioning will be difficult to predict without a greater understanding of the controls underlying the sensitivity of soils to climate variables.     

Biogeographic variation in temperature sensitivity of decomposition in forest soils

Determining soil carbon (C) responses to rising temperature is critical for projections of the feedbacks between terrestrial ecosystems, C cycle, and climate change. However, the direction and magnitude of this feedback remain highly uncertain due largely to our limited understanding of the spatial heterogeneity of soil C decomposition and its temperature sensitivity. Here we quantified C decomposition and its response to temperature change with an incubation study of soils from 203 sites across tropical to boreal forests in China spanning a wide range of latitudes (18°16′ to 51°37′N) and longitudes (81°01′ to 129°28′E). Mean annual temperature (MAT) and mean annual precipitation primarily explained the biogeographic variation in the decomposition rate and temperature sensitivity of soils: soil C decomposition rate decreased from warm and wet forests to cold and dry forests, while Q_10‐MAT (standardized to the MAT of each site) values displayed the opposite pattern. In contrast, biological factors (i.e. plant productivity and soil bacterial diversity) and soil factors (e.g. clay, pH, and C availability of microbial biomass C and dissolved organic C) played relatively small roles in the biogeographic patterns. Moreover, no significant relationship was found between Q_10‐MAT and soil C quality, challenging the current C quality–temperature hypothesis. Using a single, fixed Q_10‐MAT value (the mean across all forests), as is usually done in model predictions, would bias the estimated soil CO₂ emissions at a temperature increase of 3.0°C. This would lead to overestimation of emissions in warm biomes, underestimation in cold biomes, and likely significant overestimation of overall C release from soil to the atmosphere. Our results highlight that climate‐related biogeographic variation in soil C responses to temperature needs to be included in next‐generation C cycle models to improve predictions of C‐climate feedbacks.     

A meta‐analysis of temperature sensitivity as a microbial trait

Traits‐based approaches in microbial ecology provide a valuable way to abstract organismal interaction with the environment and to generate hypotheses about community function. Using macromolecular rate theory (MMRT), we recently identified that temperature sensitivity can be characterized as a distinct microbial trait. As temperature is fundamental in controlling biological reactions, variation in temperature sensitivity across communities, organisms, and processes has the potential to vastly improve understanding of microbial response to climate change. These microbial temperature sensitivity traits include the heat capacity (), temperature optimum (T_opt), and point of maximum temperature sensitivity (TS_max), each of which provide unique insights about organismal response to changes in temperature. In this meta‐analysis, we analyzed the distribution of these temperature sensitivity traits from bacteria, fungi, and mixed communities across a variety of biological systems (e.g., soils, oceans, foods, wastewater treatment plants) in order to identify commonalities in temperature responses across these diverse organisms and reaction rates. Our analysis of temperature sensitivity traits from over 350 temperature response curves reveals a wide distribution of temperature sensitivity traits, with T_opt and TS_max well within biological relevant temperatures. We find that traits vary significantly depending on organism type, microbial diversity, source environment, and biological process, with higher temperature sensitivity found in fungi than bacteria and in less diverse systems. Carbon dioxide production was found to be less temperature sensitive than denitrification, suggesting that changes in temperature will have a potentially larger impact on nitrogen‐related processes. As climate changes, these results have important implications for basic understanding of the temperature sensitivity of biological reactions and for ecological understanding of species’ trait distributions, as well as for improved treatment of temperature sensitivity in models.     

Vegetation type determines heterotrophic respiration in subalpine Australian ecosystems

Soils are the largest store of carbon in the biosphere and cool‐cold climate ecosystems are notable for their carbon‐rich soils. Characterizing effects of future climates on soil‐stored C is critical to elucidating feedbacks to changes in the atmospheric pool of CO₂. Subalpine vegetation in south‐eastern Australia is characterized by changes over short distances (scales of tens to hundreds of metres) in community phenotype (woodland, shrubland, grassland) and in species composition. Despite common geology and only slight changes in landscape position, we measured striking differences in a range of soil properties and rates of respiration among three of the most common vegetation communities in subalpine Australian ecosystems. Rates of heterotrophic respiration in bulk soil were fastest in the woodland community with a shrub understorey, slowest in the grassland, and intermediate in woodland with grass understorey. Respiration rates in surface soils were 2.3 times those at depth in soils from woodland with shrub understorey. Surface soil respiration in woodlands with grass understorey and in grasslands was about 3.5 times that at greater depth. Both Arrhenius and simple exponential models fitted the data well. Temperature sensitivity (Q₁₀) varied and depended on the model used as well as community type and soil depth – highlighting difficulties associated with calculating and interpreting Q₁₀. Distributions of communities in these subalpine areas are dynamic and respond over relatively short time‐frames (decades) to changes in fire regime and, possibly, to changes in climate. Shifts in boundaries among communities and possible changes in species composition as a result of both direct and indirect (e.g. via fire regime) climatic effects will significantly alter rates of respiration through plant‐mediated changes in soil chemistry. Models of future carbon cycles need to take into account changes in soil chemistry and rates of respiration driven by changes in vegetation as well as those that are temperature‐ and moisture‐driven.     

Regional variation in the temperature sensitivity of soil organic matter decomposition in China's forests and grasslands

How to assess the temperature sensitivity (Q₁₀) of soil organic matter (SOM) decomposition and its regional variation with high accuracy is one of the largest uncertainties in determining the intensity and direction of the global carbon (C) cycle in response to climate change. In this study, we collected a series of soils from 22 forest sites and 30 grassland sites across China to explore regional variation in Q₁₀ and its underlying mechanisms. We conducted a novel incubation experiment with periodically changing temperature (5–30 °C), while continuously measuring soil microbial respiration rates. The results showed that Q₁₀ varied significantly across different ecosystems, ranging from 1.16 to 3.19 (mean 1.63). Q₁₀ was ordered as follows: alpine grasslands (2.01) > temperate grasslands (1.81) > tropical forests (1.59) > temperate forests (1.55) > subtropical forests (1.52). The Q₁₀ of grasslands (1.90) was significantly higher than that of forests (1.54). Furthermore, Q₁₀ significantly increased with increasing altitude and decreased with increasing longitude. Environmental variables and substrate properties together explained 52% of total variation in Q₁₀ across all sites. Overall, pH and soil electrical conductivity primarily explained spatial variation in Q₁₀. The general negative relationships between Q₁₀ and substrate quality among all ecosystem types supported the C quality temperature (CQT) hypothesis at a large scale, which indicated that soils with low quality should have higher temperature sensitivity. Furthermore, alpine grasslands, which had the highest Q₁₀, were predicted to be more sensitive to climate change under the scenario of global warming.     

Temperature Sensitivity of Soil Organic Carbon Mineralization along an Elevation Gradient in the Wuyi Mountains,China

Soil organic carbon (SOC) actively participates in the global carbon (C) cycle. Despite much research, however, our understanding of the temperature sensitivity of soil organic carbon (SOC) mineralization is still very limited. To investigate the responses of SOC mineralization to temperature, we sampled surface soils (0–10 cm) from evergreen broad-leaf forest (EBF), coniferous forest (CF), sub-alpine dwarf forest (SDF), and alpine meadow (AM) along an elevational gradient in the Wuyi Mountains, China. The soil samples were incubated at 5, 15, 25, and 35°C with constant soil moisture for 360 days. The temperature sensitivity of SOC mineralization (Q₁₀) was calculated by comparing the time needed to mineralize the same amount of C at any two adjacent incubation temperatures. Results showed that the rates of SOC mineralization and the cumulative SOC mineralized during the entire incubation significantly increased with increasing incubation temperatures across the four sites. With the increasing extent of SOC being mineralized (increasing incubation time), the Q₁₀ values increased. Moreover, we found that both the elevational gradient and incubation temperature intervals significantly impacted Q₁₀ values. Q₁₀ values of the labile and recalcitrant organic C linearly increased with elevation. For the 5–15, 15–25, and 25–35°C intervals, surprisingly, the overall Q₁₀ values for the labile C did not decrease as the recalcitrant C did. Generally, our results suggest that subtropical forest soils may release more carbon than expected in a warmer climate.     

Drivers of temperature sensitivity of decomposition of soil organic matter along a mountain altitudinal gradient in the Western Carpathians

Mountain forest soils contain an important stock of carbon. Their altitudinal gradient can serve as a model for research on the potential risk of increased emission of carbon dioxide to the atmosphere, in a positive feedback of global warming. Using soil samples collected at three elevations (600, 900, and 1200 m a.s.l.) from five separate slopes of the Carpathian Mountains (Poland), we studied the effects of soil physical, chemical and microbial properties controlling the temperature sensitivity (Q₁₀ values) of organic matter decomposition in forest soils. Data of soil basal respiration rate measured in laboratory conditions at six different temperatures (5, 10, 15, 20, 25 and 30 °C) were fitted to a Gaussian function. The modelled soil respiration rates differed between altitudes at temperature exceeding 15 °C, and the respiration rate of soil from 1200 m a.s.l. was higher than in soils from the two lower elevations. Based on the modelled respiration values, we calculated Q₁₀ values in the low (Q₁₀L, 0–10 °C), medium (Q₁₀M, 10–20 °C) and high (Q₁₀H, 20–30 °C) temperature ranges. The Q₁₀ values did not differ between elevations. Q₁₀L and Q₁₀M were negatively related only with the C:N ratio. Temperature sensitivity of decomposition of soil organic matter was not affected by bacterial activity and functional diversity (assessed using Biolog^® ECO plates), microbial biomass or community structure (inferred from phospholipid fatty acid assays). Our findings support a kinetics-based theory of the higher temperature sensitivity of more chemically recalcitrant soil organic matter, put forward by other authors.     

Sensitivity of organic matter decomposition to warming varies with its quality

The relationship between organic matter (OM) lability and temperature sensitivity is disputed, with recent observations suggesting that responses of relatively more resistant OM to increased temperature could be greater than, equivalent to, or less than responses of relatively more labile OM. This lack of clear understanding limits the ability to forecast carbon (C) cycle responses to temperature changes. Here, we derive a novel approach (denoted Q_10?q) that accounts for changes in OM quality during decomposition and use it to analyze data from three independent sources. Results from new laboratory soil incubations (labile Q_10?q=2.1 ± 0.2; more resistant Q_10?q=3.8 ± 0.3) and reanalysis of data from other soil incubations reported in the literature (labile Q_10?q=2.3; more resistant Q_10?q=3.3) demonstrate that temperature sensitivity of soil OM decomposition increases with decreasing soil OM lability. Analysis of data from a cross‐site, field litter bag decomposition study (labile Q_10?q=3.3 ± 0.2; resistant Q_10?q=4.9 ± 0.2) shows that litter OM follows the same pattern, with greater temperature sensitivity for more resistant litter OM. Furthermore, the initial response of cultivated soils, presumably containing less labile soil OM (Q_10?q=2.4 ± 0.3) was greater than that for undisturbed grassland soils (Q_10?q=1.7 ± 0.1). Soil C losses estimated using this approach will differ from previous estimates as a function of the magnitude of the temperature increase and the proportion of whole soil OM comprised of compounds sensitive to temperature over that temperature range. It is likely that increased temperature has already prompted release of significant amounts of C to the atmosphere as CO₂. Our results indicate that future losses of litter and soil C may be even greater than previously supposed.     

Carbon quality and soil microbial property control the latitudinal pattern in temperature sensitivity of soil microbial respiration across Chinese forest ecosystems

Understanding the temperature sensitivity (Q₁₀) of soil organic C (SOC) decomposition is critical to quantifying the climate–carbon cycle feedback and predicting the response of ecosystems to climate change. However, the driving factors of the spatial variation in Q₁₀ at a continental scale are fully unidentified. In this study, we conducted a novel incubation experiment with periodically varying temperature based on the mean annual temperature of the soil origin sites. A total of 140 soil samples were collected from 22 sites along a 3,800 km long north–south transect of forests in China, and the Q₁₀ of soil microbial respiration and corresponding environmental variables were measured. Results showed that changes in the Q₁₀ values were nonlinear with latitude, particularly showing low Q₁₀ values in subtropical forests and high Q₁₀ values in temperate forests. The soil C:N ratio was positively related to the Q₁₀ values, and coniferous forest soils with low SOC quality had higher Q₁₀ values than broadleaved forest soils with high SOC quality, which supported the “C quality temperature” hypothesis. Out of the spatial variations in Q₁₀ across all ecosystems, gram‐negative bacteria exhibited the most importance in regulating the variation in Q₁₀ and contributed 25.1%, followed by the C:N ratio (C quality), fungi, and the fungi:bacteria ratio. However, the dominant factors that regulate the regional variations in Q₁₀ differed among the tropical, subtropical, and temperate forest ecosystems. Overall, our findings highlight the importance of C quality and microbial controls over Q₁₀ value in China's forest ecosystems. Meanwhile, C dynamics in temperate forests under a global warming scenario can be robustly predicted through the incorporation of substrate quality and microbial property into models.     

Temperature and vegetation effects on soil organic carbon quality along a forested mean annual temperature gradient in North America

Both climate and plant species are hypothesized to influence soil organic carbon (SOC) quality, but accurate prediction of how SOC process rates respond to global change will require an improved understanding of how SOC quality varies with mean annual temperature (MAT) and forest type. We investigated SOC quality in paired hardwood and pine stands growing in coarse textured soils located along a 22 °C gradient in MAT. To do this, we conducted 80‐day incubation experiments at 10 and 30 °C to quantify SOC decomposition rates, which we used to kinetically define SOC quality. We used these experiments to test the hypotheses that SOC quality decreases with MAT, and that SOC quality is higher under pine than hardwood tree species. We found that both SOC quantity and quality decreased with increasing MAT. During the 30 °C incubation, temperature sensitivity (Q₁₀) values were strongly and positively related to SOC decomposition rates, indicating that substrate supply can influence temperature responsiveness of SOC decomposition rates. For a limited number of dates, Q₁₀ was negatively related to MAT. Soil chemical properties could not explain observed patterns in soil quality. Soil pH and cation exchange capacity (CEC) both declined with increasing MAT, and soil C quality was positively related to pH but negatively related to CEC. Clay mineralogy of soils also could not explain patterns of SOC quality as complex (2 : 1), high CEC clay minerals occurred in cold climate soils while warm climate soils were dominated by simpler (1 : 1), low CEC clay minerals. While hardwood sites contained more SOC than pine sites, with differences declining with MAT, clay content was also higher in hardwood soils. In contrast, there was no difference in SOC quality between pine and hardwood soils. Overall, these findings indicate that SOC quantity and quality may both decrease in response to global warming, despite long‐term changes in soil chemistry and mineralogy that favor decomposition.     

Exponential growth of “snow molds” at sub-zero temperatures: an explanation for high beneath-snow respiration rates and Q 10 values

Numerous studies have demonstrated exceptionally high temperature sensitivity of the beneath-snow respiratory flux in cold-winter ecosystems. The most common, but still untested, explanation for this high sensitivity is a physical one based on the observation that water availability in soils increases exponentially as soils warm from −3 to 0°C. Here, we present evidence for a biological hypothesis to explain exponential kinetics and high Q ₁₀ values as beneath-snow soils warm from −3 to 0°C during the early spring in a high-elevation subalpine forest. First, we show that some of the dominant organisms of the beneath-snow microbial community, “snow molds”, exhibit robust exponential growth at temperatures from −3 to −0.3°C. Second, Q ₁₀ values based on growth rates across the temperature range of −2 to −0.3°C for these snow molds vary from 22 to 330. Third, we derive an analytical equation that combines the relative contributions of microbial growth and microbial metabolism to the temperature sensitivity of respiration. Finally, we use this equation to show that with only moderate snow mold growth (several generations), the combined sensitivities of growth and metabolism to small changes in beneath-snow soil temperature, create a double exponential in the Q ₁₀ function that may explain the extremely high (~1 × 10⁶) Q ₁₀ values observed in past studies. Our biological explanation for high Q ₁₀ levels is supported by several independent studies that have demonstrated build up of microbial biomass under the snow as temperatures warm from −2 to 0°C.     

Effects of soil moisture on the temperature sensitivity of heterotrophic respiration vary seasonally in an old‐field climate change experiment

Microbial decomposition of soil organic matter produces a major flux of CO₂ from terrestrial ecosystems and can act as a feedback to climate change. Although climate‐carbon models suggest that warming will accelerate the release of CO₂ from soils, the magnitude of this feedback is uncertain, mostly due to uncertainty in the temperature sensitivity of soil organic matter decomposition. We examined how warming and altered precipitation affected the rate and temperature sensitivity of heterotrophic respiration (R_h) at the Boston‐Area Climate Experiment, in Massachusetts, USA. We measured R_h inside deep collars that excluded plant roots and litter inputs. In this mesic ecosystem, R_h responded strongly to precipitation. Drought reduced R_h, both annually and during the growing season. Warming increased R_h only in early spring. During the summer, when R_h was highest, we found evidence of threshold, hysteretic responses to soil moisture: R_h decreased sharply when volumetric soil moisture dropped below ~15% or exceeded ~26%, but R_h increased more gradually when soil moisture rose from the lower threshold. The effect of climate treatments on the temperature sensitivity of R_h depended on the season. Apparent Q₁₀ decreased with high warming (~3.5 °C) in spring and fall. Presumably due to limiting soil moisture, warming and precipitation treatments did not affect apparent Q₁₀ in summer. Drought decreased apparent Q₁₀ in fall compared to ambient and wet precipitation treatments. To our knowledge, this is the first field study to examine the response of R_h and its temperature sensitivity to the combined effects of warming and altered precipitation. Our results highlight the complex responses of R_h to soil moisture, and to our knowledge identify for the first time the seasonal variation in the temperature sensitivity of microbial respiration in the field. We emphasize the importance of adequately simulating responses such as these when modeling trajectories of soil carbon stocks under climate change scenarios.     

Macromolecular rate theory explains the temperature dependence of membrane conductance kinetics

The factor Q₁₀ is used in neuroscience to adjust reaction rates of voltage-activated membrane conductances to different temperatures and is widely assumed to be constant. By performing an analysis of published data of the reaction rates of sodium, potassium, and calcium membrane conductances, we demonstrate that 1) Q₁₀ is temperature dependent, 2) this relationship is similar across conductances, and 3) there is a strong effect at low temperatures (<15°C). We show that macromolecular rate theory (MMRT) explains this temperature dependency. MMRT predicts the existence of optimal temperatures at which reaction rates decrease as temperature increases, a phenomenon that we also found in the published data sets. We tested the consequences of using MMRT-adjusted reaction rates in the Hodgkin-Huxley model of the squid’s giant axon. The MMRT-adjusted model reproduces the temperature dependence of the rising and falling times of the action potential. Furthermore, the model also reproduces these properties for different squid species that live in different climates. In a second example, we compare spiking patterns of biophysical models based on human pyramidal neurons from the Allen Cell Types database at room and physiological temperatures. The original models, calibrated at 34°C, failed to generate realistic spikes at room temperature in more than half of the tested models, while the MMRT produces realistic spiking in all conditions. In another example, we show that using the MMRT correction in hippocampal pyramidal cell models results in 100% differences in voltage responses. Finally, we show that the shape of the Q₁₀ function results in systematic errors in predicting reaction rates. We propose that the optimal temperature could be a thermodynamical barrier to avoid over excitation in neurons. While this study is centered on membrane conductances, our results have important consequences for all biochemical reactions involved in cell signaling.     

Reduction of the temperature sensitivity of soil organic matter decomposition with sustained temperature increase

The degree to which microbial communities adjust their decomposition of soil carbon over time in response to long-term increases in temperature is one of the key uncertainties in our modeling of the responses of terrestrial ecosystems to warming. To better understand changes in temperature sensitivity of soil microbial communities to long-term increases in soil temperature, we incubated 27 soils for one year with both short-term and long-term manipulations of temperature. In response to increasing temperature short-term from 20 to 30 °C, respiration rates increased more than threefold on average across soils. Yet, in response to long-term increases in temperature, respiration rates increased approximately half as much as they did to short-term increases in temperature. Short-term Q₁₀ of recalcitrant C correlated positively with long-term Q₁₀ measured between 10 and 20 °C, yet there was no relationship between short-term Q₁₀ and long-term Q₁₀ between 20 and 30 °C. In all, under laboratory conditions, it is clear that there is reduction in the temperature sensitivity of decomposition to long-term increases in temperature that disassociate short- and long-term responses of microbial decomposition to temperature. Determining the fate of soil organic matter to increased temperature will not only require further research on the controls and mechanisms of these patterns, but also require models to incorporate responses to both short-term and long-term increases in temperature.     

Different temperature sensitivity and kinetics of soil enzymes indicate seasonal shifts in C,N and P nutrient stoichiometry in acid forest soil

Acid forest soils in the Bohemian Forest in Central Europe are biogeochemically imbalanced in organic C, N and P processing. We hypothesized that these imbalances can be due to different temperature sensitivities of soil enzyme activities and their affinities to substrate in litter and organic soil horizons. We measured potential activities of five main soil enzymes (β-glucosidase, cellobiohydrolase, Leu-aminopeptidase, Ala-aminopeptidase, and phosphatase) responsible for organic carbon, nitrogen and phosphorus acquisition. We also modeled potential in situ enzyme activities and nutrient release based on continuous in situ temperature measurements. We determined basic kinetic parameters (K_m, V_max), enzyme efficiencies (k_cat) and temperature sensitivities (E_a and Q₁₀) according to Michaelis–Menten kinetic and modified Arrhenius models. Our results showed significant differences in substrate affinities between the litter and organic soil horizons. Higher aminopeptidase affinity (lower K_m) in the litter soil horizon can lead to leaching of peptidic compounds to lower soil horizons. β-Glucosidase and phosphatase showed high temperature response following the Arrhenius model. However, both aminopeptidases showed no or even decreased activity with increasing temperature. The aminopeptidase temperature insensitivity means that peptidic compounds are degraded at the same or even lower rate in warmer and colder periods of the year in acid forest soils. This imbalance results in different release of available nutrients from plant litter and soil organic matter which may affect bacterial and fungal community composition and nutrient leaching from these ecosystems.     

封育对羊草草地土壤碳矿化激发效应和温度敏感性的影响

土壤碳矿化(或土壤异养呼吸)的温度敏感性和激发效应是深入揭示土壤呼吸控制机理及其对未来气候变化响应与适应的重要研究方向。该文以自由放牧(FG0)、封育11年(FG11)、封育31年(FG31)的羊草(Leymus chinensis)草地为研究对象, 通过0、5、10、15、20、25 ℃培养, 探讨了封育对羊草草地土壤碳矿化激发效应和温度敏感性的影响。结果表明: 封育年限、添加葡萄糖、培养温度和培养时间对土壤碳矿化速率均具有显著的影响, 不同因素间存在显著的交互效应(p < 0.000 1)。FG0的羊草草地土壤碳矿化累积量显著高于FG11和FG31的, 在添加葡萄糖处理下也呈现相同的趋势。长期封育降低了羊草草地土壤碳矿化的激发效应。在添加葡萄糖后, 培养前7天的土壤碳矿化的激发效应随温度增加而增加, 增加2.28-9.01倍; 在整个56天培养期间, 激发效应介于2.21-5.10倍, 最高值出现在10或15 ℃。土壤碳矿化速率可用经典的指数方程来表示, FG0草地的土壤碳矿化的温度敏感性指数(Q₁₀)大于长期封育草地(FG11和FG31); 与未添加处理相比, 添加葡萄糖显著增加了土壤碳矿化速率的温度敏感性, 即在添加葡萄糖后土壤微生物呼吸受温度的影响更大。长期封育会降低羊草草地土壤的碳矿化速率、温度敏感性和激发效应, 从而降低土壤碳周转速率和释放速率, 使内蒙古地区长期封育草地仍然具有碳固持能力。