Experimental warming and burn severity alter soil CO2 flux and soil functional groups in a recently burned boreal forest

Global warming is projected to be greatest in northern regions, where forest fires are also increasing in frequency. Thus, interactions between fire and temperature on soil respiration at high latitudes should be considered in determining feedbacks to climate. We tested the hypothesis that experimental warming will augment soil CO₂ flux in a recently burned boreal forest by promoting microbial and root growth, but that this increase will be less apparent in more severely burned areas. We used open‐top chambers to raise temperatures 0.4–0.9°C across two levels of burn severity in a fire scar in Alaskan black spruce forest. After 3 consecutive years of warming, soil respiration was measured through a portable gas exchange system. Abundance of active microbes was determined by using Biolog EcoPlates^? for bacteria and ergosterol analysis for fungi. Elevated temperatures increased soil CO₂ flux by 20% and reduced root biomass, but had no effect on bacterial or fungal abundance or soil organic matter (SOM) content. Soil respiration, fungal abundance, SOM, and root biomass decreased with increasing burn severity. There were no significant interactions between temperature and burn severity with respect to any measurement. Higher soil respiration rates in the warmed plots may be because of higher metabolic activity of microbes or roots. All together, we found that postfire soils are a greater source of CO₂ to the atmosphere under elevated temperatures even in severely burned areas, suggesting that global warming may produce a positive feedback to atmospheric CO₂, even in young boreal ecosystems.     

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

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

Rates of net nitrogen mineralization in disturbed and undisturbed soils

Quantification of net nitrogen mineralization (NNM) in soils is indispensable in order to optimize N fertilization of crops. Two long-term laboratory incubation methods were applied to determine rates of net nitrogen mineralization (r_NNM) of soils from two sites of arable land (sandy loam soil, silty loam soil) at four temperature levels (2°C, 8°C, 14°C, 21°C). Since variability within replicates was small, the modified 12-week incubation method of Stanford and Smith (1972) using disturbed soils allowed to establish reliable Arrhenius functions with reasonable expenditure. The fit of the functions derived from the 5-month incubation of 23 undisturbed soil columns (4420 cm³) was worse. This was caused by greater variability and less differentiation between temperature levels. Results of both experiments could be described best by zero-order kinetics. Mean mineralization rates of disturbed samples were approximately twice as high than those of undisturbed samples. The suitability of both methods for the prediction of NNM at site conditions is discussed. Actual respiration (AR) at incubation temperatures and substrate induced respiration (SIR) were measured at the end of the incubation of undisturbed soil columns. The results presented reveal that soil microbial communities develop in a different manner during long-term incubation at different temperatures. This behavior offends the underlying assumption that soil microbes remain in steady-state during incubation and that rising rates are physiological reactions to temperature enhancement. Therefore soil microbial biomass (SMB) dynamics during the experiment has to be accounted for when rates of NNM and Arrhenius functions are established. R Merck Section editor     

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.     

Initial responses of soil CO2 efflux and C, N pools to experimental warming in two contrasting forest ecosystems, Eastern Tibetan Plateau, China

Alpine ecosystems are harsh environments where low temperatures are generally a limiting factor. Predicted global warming is thus expected to have a profound impact on alpine ecosystems in the future. This study was conducted to compare the effect of experimental warming on soils in two contrasting forest ecosystems (a dragon spruce plantation and a natural forest) using the open top chamber (OTC) method in the Eastern Tibetan Plateau of China. The OTC enhanced average daily mean soil temperatures by 0.61°C (plantation) and 0.55°C (natural forest), respectively, throughout the growing season. Conversely, soil volumetric moisture declined by 4.10% in the plantation and by 2.55% in the natural forest. Across all measuring dates, warming increased average soil CO₂ efflux by 10.6% in the plantation and by 15.4% in the natural forest. However, elevated temperatures did not affect the respiration quotient in either forest. Two-stage sulfuric acid hydrolysis was used to quantify labile and recalcitrant C and N fractions in the two contrasting soils. Warming significantly reduced labile C and N fractions in both ecosystems but did not influence the total, recalcitrant and microbial biomass C and N pools. Labile C, N and microbial biomass C showed significant interactions in warming × forest type × season. Irrespective of warming treatments, all measured pools were significantly larger in the natural forest compared to the plantation. Taken together, our results indicate that the lowered soil labile C and N pools may be induced by the increased soil CO₂ efflux. The responses of the natural forest soil were more sensitive to experimental warming than those of the plantation. We conclude that reforestation dramatically lowers soil C and N pools, further affecting the responses of forest soils to future global warming.     

Soil microbial responses to experimental warming and clipping in a tallgrass prairie

Global surface temperature is predicted to increase by 1.4–5.8°C by the end of this century. However, the impacts of this projected warming on soil C balance and the C budget of terrestrial ecosystems are not clear. One major source of uncertainty stems from warming effects on soil microbes, which exert a dominant influence on the net C balance of terrestrial ecosystems by controlling organic matter decomposition and plant nutrient availability. We, therefore, conducted an experiment in a tallgrass prairie ecosystem at the Great Plain Apiaries (near Norman, OK) to study soil microbial responses to temperature elevation of about 2°C through artificial heating in clipped and unclipped field plots. While warming did not induce significant changes in net N mineralization, soil microbial biomass and respiration rate, it tended to reduce extractable inorganic N during the second and third warming years, likely through increasing plant uptake. In addition, microbial substrate utilization patterns and the profiles of microbial phospholipid fatty acids (PLFAs) showed that warming caused a shift in the soil microbial community structure in unclipped subplots, leading to the relative dominance of fungi as evidenced by the increased ratio of fungal to bacterial PLFAs. However, no warming effect on soil microbial community structure was found in clipped subplots where a similar scale of temperature increase occurred. Clipping also significantly reduced soil microbial biomass and respiration rate in both warmed and unwarmed plots. These results indicated that warming‐led enhancement of plant growth rather than the temperature increase itself may primarily regulate soil microbial response. Our observations show that warming may increase the relative contribution of fungi to the soil microbial community, suggesting that shifts in the microbial community structure may constitute a major mechanism underlying warming acclimatization of soil respiration.     

岷江上游华山松林冬季土壤呼吸对模拟增温的短期响应

冬季的土壤呼吸是生态系统呼吸的重要组成部分, 对气候变化的响应可能更为敏感。该文采用红外辐射加热器模拟土壤增温, 研究了岷江上游华山松(Pinus armandii)人工林冬季的土壤呼吸、微生物生物量及无机氮库对模拟增温的响应。结果表明: 在冬季(2009年11月-翌年3月), 模拟增温往往能显著提高土壤呼吸速率, 平均增幅达31.4%; 同样模拟增温使土壤微生物生物量碳、氮分别增加23.2%和22.7%, 而对微生物生物量碳氮比没有影响, 温度升高显著促进了微生物的生长, 但没有改变微生物的群落结构; 增温样地土壤的NO₃ ^--N和NH₄ ⁺-N浓度较对照分别增加了38.5%和12.3%, 增温显著提高了土壤的可溶性无机氮含量。综上所述, 该区针叶林冬季土壤呼吸、微生物生长和养分矿化对未来气候变暖非常敏感。     

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

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

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

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

Microbial community responses reduce soil carbon loss in Tibetan alpine grasslands under short‐term warming

Changes in labile carbon (LC) pools and microbial communities are the primary factors controlling soil heterotrophic respiration (R_h) in warming experiments. Warming is expected to initially increase R_h but studies show this increase may not be continuous or sustained. Specifically, LC and soil microbiome have been shown to contribute to the effect of extended warming on R_h. However, their relative contribution is unclear and this gap in knowledge causes considerable uncertainty in the prediction of carbon cycle feedbacks to climate change. In this study, we used a two‐step incubation approach to reveal the relative contribution of LC limitation and soil microbial community responses in attenuating the effect that extended warming has on R_h. Soil samples from three Tibetan ecosystems—an alpine meadow (AM), alpine steppe (AS), and desert steppe (DS)—were exposed to a temperature gradient of 5–25°C. After an initial incubation period, soils were processed in one of two methods: (a) soils were sterilized then inoculated with parent soil microbes to assess the LC limitation effects, while controlling for microbial community responses; or (b) soil microbes from the incubations were used to inoculate sterilized parent soils to assess the microbial community effects, while controlling for LC limitation. We found both LC limitation and microbial community responses led to significant declines in R_h by 37% and 30%, respectively, but their relative contributions were ecosystem specific. LC limitation alone caused a greater R_h decrease for DS soils than AMs or ASs. Our study demonstrates that soil carbon loss due to R_h in Tibetan alpine soils—especially in copiotrophic soils—will be weakened by microbial community responses under short‐term warming.     

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.     

Effects of long-term experimental warming on plants and soil microbes in a cool temperate semi-natural grassland in Japan

To clarify the effects of long-term warming on ecosystem matter cycling, we conducted an in situ 7-year experimental warming (2009–2015) using infrared heaters in a cool temperate semi-natural grassland in Japan. We measured plant aboveground biomass, soil total C and N, soil inorganic N (NH₄ ⁺-N and NO₃ ^?-N), and soil microbial biomass for 7 years (2009–2015). We also measured heterotrophic respiration for 2 years (2013–2014) and assessed net N mineralization and nitrification in 2015. We found that warming immediately increased plant aboveground biomass, but this effect ceased in 2013. However, the soil microbial biomass was continuously depressed by warming. Soil inorganic N concentrations in warmed plots substantially increased in the later years of the experiment (2013–2015) and the potential net N mineralization rate was also higher than in the earlier years. In contrast, heterotrophic respiration decreased with warming in 2013–2014. Our observations indicate that long-term warming has a contrasting effect on plants and soil microbes. In addition, the warming could have different effects on subterranean C and N cycling. To enhance the accuracy of estimation of future climate change, it is essential to continuously observe the warming effects on ecosystems and to focus on the change in subterranean C and N cycling.     

Offsetting global warming‐induced elevated greenhouse gas emissions from an arable soil by biochar application

Global warming will likely enhance greenhouse gas (GHG) emissions from soils. Due to its slow decomposability, biochar is widely recognized as effective in long‐term soil carbon (C) sequestration and in mitigation of soil GHG emissions. In a long‐term soil warming experiment (+2.5 °C, since July 2008) we studied the effect of applying high‐temperature Miscanthus biochar (0, 30 t/ha, since August 2013) on GHG emissions and their global warming potential (GWP) during 2 years in a temperate agroecosystem. Crop growth, physical and chemical soil properties, temperature sensitivity of soil respiration (R_s), and metabolic quotient (qCO₂) were investigated to yield further information about single effects of soil warming and biochar as well as on their interactions. Soil warming increased total CO₂ emissions by 28% over 2 years. The effect of warming on soil respiration did not level off as has often been observed in less intensively managed ecosystems. However, the temperature sensitivity of soil respiration was not affected by warming. Overall, biochar had no effect on most of the measured parameters, suggesting its high degradation stability and its low influence on microbial C cycling even under elevated soil temperatures. In contrast, biochar × warming interactions led to higher total N₂O emissions, possibly due to accelerated N‐cycling at elevated soil temperature and to biochar‐induced changes in soil properties and environmental conditions. Methane uptake was not affected by soil warming or biochar. The incorporation of biochar‐C into soil was estimated to offset warming‐induced elevated GHG emissions for 25 years. Our results highlight the suitability of biochar for C sequestration in cultivated temperate agricultural soil under a future elevated temperature. However, the increased N₂O emissions under warming limit the GHG mitigation potential of biochar.     

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

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

Differential Nutrient Limitation of Soil Microbial Biomass and Metabolic Quotients (qCO2): Is There a Biological Stoichiometry of Soil Microbes?

Background

Variation in microbial metabolism poses one of the greatest current uncertainties in models of global carbon cycling, and is particularly poorly understood in soils. Biological Stoichiometry theory describes biochemical mechanisms linking metabolic rates with variation in the elemental composition of cells and organisms, and has been widely observed in animals, plants, and plankton. However, this theory has not been widely tested in microbes, which are considered to have fixed ratios of major elements in soils.

Methodology/Principal Findings

To determine whether Biological Stoichiometry underlies patterns of soil microbial metabolism, we compiled published data on microbial biomass carbon (C), nitrogen (N), and phosphorus (P) pools in soils spanning the global range of climate, vegetation, and land use types. We compared element ratios in microbial biomass pools to the metabolic quotient qCO₂ (respiration per unit biomass), where soil C mineralization was simultaneously measured in controlled incubations. Although microbial C, N, and P stoichiometry appeared to follow somewhat constrained allometric relationships at the global scale, we found significant variation in the C∶N∶P ratios of soil microbes across land use and habitat types, and size-dependent scaling of microbial C∶N and C∶P (but not N∶P) ratios. Microbial stoichiometry and metabolic quotients were also weakly correlated as suggested by Biological Stoichiometry theory. Importantly, we found that while soil microbial biomass appeared constrained by soil N availability, microbial metabolic rates (qCO₂) were most strongly associated with inorganic P availability.

Conclusions/Significance

Our findings appear consistent with the model of cellular metabolism described by Biological Stoichiometry theory, where biomass is limited by N needed to build proteins, but rates of protein synthesis are limited by the high P demands of ribosomes. Incorporation of these physiological processes may improve models of carbon cycling and understanding of the effects of nutrient availability on soil C turnover across terrestrial and wetland habitats.     

Differential effects of warming and nitrogen fertilization on soil respiration and microbial dynamics in switchgrass croplands

The mechanistic understanding of warming and nitrogen (N) fertilization, alone or in combination, on microbially mediated decomposition is limited. In this study, soil samples were collected from previously harvested switchgrass (Panicum virgatum L.) plots that had been treated with high N fertilizer (HN: 67 kg N ha^?1) and those that had received no N fertilizer (NN) over a 3‐year period. The samples were incubated for 180 days at 15 °C and 20 °C, during which heterotrophic respiration, δ¹³C of CO₂, microbial biomass (MB), specific soil respiration rate (R_s: respiration per unit of microbial biomass), and exoenzyme activities were quantified at 10 different collections time. Employing switchgrass tissues (referred to as litter) with naturally abundant ¹³C allowed us to partition CO₂ respiration derived from soil and amended litter. Cumulative soil respiration increased significantly by 16.4% and 4.2% under warming and N fertilization, respectively. Respiration derived from soil was elevated significantly with warming, while oxidase, the agent for recalcitrant soil substrate decomposition, was not significantly affected by warming. Warming, however, significantly enhanced MB and R_s indicating a decrease in microbial growth efficiency (MGE). On the contrary, respiration derived from amended litter was elevated with N fertilization, which was consistent with the significantly elevated hydrolase. N fertilization, however, had little effect on MB and R_s, suggesting little change in microbial physiology. Temperature and N fertilization showed minimal interactive effects likely due to little differences in soil N availability between NN and HN samples, which is partly attributable to switchgrass biomass N accumulation (equivalent to ~53% of fertilizer N). Overall, the differential individual effects of warming and N fertilization may be driven by physiological adaptation and stimulated exoenzyme kinetics, respectively. The study shed insights on distinct microbial acquisition of different substrates under global temperature increase and N enrichment.     

Does long-term soil warming affect microbial element limitation? A test by short-term assays of microbial growth responses to labile C,N and P additions

Increasing global temperatures have been reported to accelerate soil carbon (C) cycling, but also to promote nitrogen (N) and phosphorus (P) dynamics in terrestrial ecosystems. However, warming can differentially affect ecosystem C, N and P dynamics, potentially intensifying elemental imbalances between soil resources, plants and soil microorganisms. Here, we investigated the effect of long-term soil warming on microbial resource limitation, based on measurements of microbial growth (¹⁸O incorporation into DNA) and respiration after C, N and P amendments. Soil samples were taken from two soil depths (0–10, 10–20 cm) in control and warmed (>14 years warming, +4°C) plots in the Achenkirch soil warming experiment. Soils were amended with combinations of glucose-C, inorganic/organic N and inorganic/organic P in a full factorial design, followed by incubation at their respective mean field temperatures for 24 h. Soil microbes were generally C-limited, exhibiting 1.8-fold to 8.8-fold increases in microbial growth upon C addition. Warming consistently caused soil microorganisms to shift from being predominately C limited to become C-P co-limited. This P limitation possibly was due to increased abiotic P immobilization in warmed soils. Microbes further showed stronger growth stimulation under combined glucose and inorganic nutrient amendments compared to organic nutrient additions. This may be related to a prolonged lag phase in organic N (glucosamine) mineralization and utilization compared to glucose. Soil respiration strongly positively responded to all kinds of glucose-C amendments, while responses of microbial growth were less pronounced in many of these treatments. This highlights that respiration–though easy and cheap to measure—is not a good substitute of growth when assessing microbial element limitation. Overall, we demonstrate a significant shift in microbial element limitation in warmed soils, from C to C-P co-limitation, with strong repercussions on the linkage between soil C, N and P cycles under long-term warming.     

Microbial physiology and soil CO2 efflux after 9 years of soil warming in a temperate forest – no indications for thermal adaptations

Thermal adaptations of soil microorganisms could mitigate or facilitate global warming effects on soil organic matter (SOM) decomposition and soil CO₂ efflux. We incubated soil from warmed and control subplots of a forest soil warming experiment to assess whether 9 years of soil warming affected the rates and the temperature sensitivity of the soil CO₂ efflux, extracellular enzyme activities, microbial efficiency, and gross N mineralization. Mineral soil (0–10 cm depth) was incubated at temperatures ranging from 3 to 23 °C. No adaptations to long‐term warming were observed regarding the heterotrophic soil CO₂ efflux (R₁₀ warmed: 2.31 ± 0.15 μmol m^?2 s^?1, control: 2.34 ± 0.29 μmol m^?2 s^?1; Q₁₀ warmed: 2.45 ± 0.06, control: 2.45 ± 0.04). Potential enzyme activities increased with incubation temperature, but the temperature sensitivity of the enzymes did not differ between the warmed and the control soils. The ratio of C : N acquiring enzyme activities was significantly higher in the warmed soil. Microbial biomass‐specific respiration rates increased with incubation temperature, but the rates and the temperature sensitivity (Q₁₀ warmed: 2.54 ± 0.23, control 2.75 ± 0.17) did not differ between warmed and control soils. Microbial substrate use efficiency (SUE) declined with increasing incubation temperature in both, warmed and control, soils. SUE and its temperature sensitivity (Q₁₀ warmed: 0.84 ± 0.03, control: 0.88 ± 0.01) did not differ between warmed and control soils either. Gross N mineralization was invariant to incubation temperature and was not affected by long‐term soil warming. Our results indicate that thermal adaptations of the microbial decomposer community are unlikely to occur in C‐rich calcareous temperate forest soils.     

Temperature responses of carbon mineralization in conifer forest soils from different regional climates incubated under standard laboratory conditions

¹⁴C‐labelled straw was mixed with soils collected from seven coniferous forests located on a climatic gradient in Western Europe ranging from boreal to Mediterranean conditions. The soils were incubated in the laboratory at 4°, 10°, 16°, 23° and 30 °C with constant moisture over 550 days. The temperature coefficient (Q₁₀) for straw carbon mineralization decreased with increasing incubation temperatures. This was a characteristic of all the soils with a difference of two Q₁₀ units between the 4–10° and the 23? 30 °C temperature ranges. It was also found that the magnitude of the temperature response function was related to the period of soil incubation. Initial temperature responses of microbial communities were different to those shown after a long period of laboratory incubation and may have reflected shifts in microbial species composition in response to changes in the temperature regime. The rapid exhaustion of the labile fractions of the decomposing material at higher temperatures could also lead to underestimation of the temperature sensitivity of soils unless estimated for carbon pools of similar qualities. Finally, the thermal optima for the organic soil horizons (Of and Oh) were lower than 30 °C even after 550 days of incubation. It was concluded that these responses could not be attributed to microbial physiological adaptations, but rather to the rates at which recalcitrant microbial secondary products were formed at higher temperatures. The implication of these variable temperature responses of soil materials is discussed in relation to modelling potential effects of global warming.     

Soil biological activity and their seasonal variations in response to long-term application of organic and inorganic fertilizers

The objectives of this study were to explore the effects of long-term and continued application of fertilizers and manures on microbial biomass, soil biological activity and their seasonal variations in surface and subsurface soils in relation to soil fertility. For this, soils were sampled in spring, summer and autumn from Shenyang Long-term Experimental Station, northeastern China. The results showed that soil total nitrogen (N), organic carbon (C), basal respiration, microbial biomass and enzymatic activity increased in manure-amended surface soils, but decreased with soil depth. Long-term application of inorganic fertilizers significantly decreased soil pH value, sucrase activity and microbial biomass C, but increased soil metabolic quotient (qCO₂). However, no significant effect of inorganic fertilizers on soil total N, urease activity and microbial biomass N was observed in comparison with CK0 (neither tillage nor fertilization) and CK (no fertilizers). There was no significant difference between CK0 and CK in soil total N, organic C and microbial activity in surface soil layer (0–20 cm), but these parameters in subsurface soil layer (20–40 cm) were higher in CK than in CK0. Moreover, seasonal changes were observed in terms of soil nutrient contents, enzymatic activity, microbial biomass and soil respiration. There were significant correlations between soil microbial biomass C and N, between organic C and sucrase activity and between total N and urease activity, respectively. It is recommended that combined use of organic manure with inorganic fertilizers should be considered to maintain higher microbial biomass, soil biological activity and soil fertility. Considering considerably high nutrients reserve and microbial activity in subsurface layers of soil and wind-erosion-caused nutrient loss in spring in north China, we also propose that low tillage should be considered to make use of nutrients in soils.