The Dual Arrhenius and Michaelis–Menten kinetics model for decomposition of soil organic matter at hourly to seasonal time scales

Decomposition of soil carbon stocks is one of the largest potential biotic feedbacks to climate change. Models of decomposition of soil organic matter and of soil respiration rely on empirical functions that relate variation in temperature and soil water content to rates of microbial metabolism using soil‐C substrates. Here, we describe a unifying modeling framework to combine the effects of temperature, soil water content, and soluble substrate supply on decomposition of soluble soil‐C substrates using simple functions based on process concepts. The model's backbone is the Michaelis–Menten equation, which describes the relationship between reaction velocity and soluble organic‐C and O₂ substrate concentrations at an enzyme's reactive site, which are determined by diffusivity functions based on soil water content. Temperature sensitivity is simulated by allowing the maximum velocity of the reaction (V_max) to vary according to Arrhenius function. The Dual Arrhenius and Michaelis–Menten kinetics (DAMM) model core was able to predict effectively observations from of laboratory enzyme assays of β‐glucosidase and phenol‐oxidase across a range of substrate concentrations and incubation temperatures. The model also functioned as well or better than purely empirical models for simulating hourly and seasonal soil respiration data from a trenched plot in a deciduous forest at the Harvard Forest, in northeastern United States. The DAMM model demonstrates that enzymatic processes can be intrinsically temperature sensitive, but environmental constrains of substrate supply under soil moisture extremes can prevent that response to temperature from being observed. We discuss how DAMM could serve as a core module that is informed by other modules regarding microbial dynamics and supply of soluble‐C substrates from plant inputs and from desorption of physically stabilized soil‐C pools. Most importantly, it presents a way forward from purely empirical representation of temperature and moisture responses and integrates temperature‐sensitive enzymatic processes with constraints of substrate supply.     

Differential physiological responses to environmental change promote woody shrub expansion

Direct and indirect effects of warming are increasingly modifying the carbon-rich vegetation and soils of the Arctic tundra, with important implications for the terrestrial carbon cycle. Understanding the biological and environmental influences on the processes that regulate foliar carbon cycling in tundra species is essential for predicting the future terrestrial carbon balance in this region. To determine the effect of climate change impacts on gas exchange in tundra, we quantified foliar photosynthesis (A_net), respiration in the dark and light (R_D and R_L, determined using the Kok method), photorespiration (PR), carbon gain efficiency (CGE, the ratio of photosynthetic CO₂ uptake to total CO₂ exchange of photosynthesis, PR, and respiration), and leaf traits of three dominant species – Betula nana, a woody shrub; Eriophorum vaginatum, a graminoid; and Rubus chamaemorus, a forb – grown under long-term warming and fertilization treatments since 1989 at Toolik Lake, Alaska. Under warming, B. nana exhibited the highest rates of A_net and strongest light inhibition of respiration, increasing CGE nearly 50% compared with leaves grown in ambient conditions, which corresponded to a 52% increase in relative abundance. Gas exchange did not shift under fertilization in B. nana despite increases in leaf N and P and near-complete dominance at the community scale, suggesting a morphological rather than physiological response. Rubus chamaemorus, exhibited minimal shifts in foliar gas exchange, and responded similarly to B. nana under treatment conditions. By contrast, E. vaginatum, did not significantly alter its gas exchange physiology under treatments and exhibited dramatic decreases in relative cover (warming: −19.7%; fertilization: −79.7%; warming with fertilization: −91.1%). Our findings suggest a foliar physiological advantage in the woody shrub B. nana that is further mediated by warming and increased soil nutrient availability, which may facilitate shrub expansion and in turn alter the terrestrial carbon cycle in future tundra environments.     

Differences in gas exchange contribute to habitat differentiation in Iberian columbines from contrasting light and water environments

During photosynthesis, respiration and transpiration, gas exchange occurs via the stomata and so plants face a trade‐off between maximising photosynthesis while minimising transpiration (expressed as water use efficiency, WUE). The ability to cope with this trade‐off and regulate photosynthetic rate and stomatal conductance may be related to niche differentiation between closely related species. The present study explored this as a possible mechanism for habitat differentiation in Iberian columbines. The roles of irradiance and water stress were assessed to determine niche differentiation among Iberian columbines via distinct gas exchange processes. Photosynthesis–irradiance curves (P–I curves) were obtained for four taxa, and common garden experiments were conducted to examine plant responses to water and irradiance stress, by measuring instantaneous gas exchange and plant performance. Gas exchange was also measured in ten individuals using two to four field populations per taxon. The taxa had different P–I curves and gas exchange in the field. At the species level, water stress and irradiance explained habitat differentiation. Within each species, a combination of irradiance and water stress explained the between‐subspecies habitat differentiation. Despite differences in stomatal conductance and CO₂ assimilation, taxa did not have different WUE under field conditions, which suggests that the environment equally modifies photosynthesis and transpiration. The P–I curves, gas exchange in the field and plant responses to experimental water and irradiance stresses support the hypothesis that habitat differentiation is associated with differences among taxa in tolerance to abiotic stress mediated by distinct gas exchange responses.     

CO2 exchange in an organic field growing barley or grass in eastern Finland

The CO₂ dynamics were measured in an organic soil in eastern Finland during the growing season and wintertime, and the annual CO₂ balance was calculated for plots where barley or grass was grown. During the summer, the CO₂ dynamics were measured by transparent and opaque chambers using a portable infrared gas analyser for the CO₂ analyses. During the winter, the CO₂ release was measured by opaque chambers analysing the samples in the laboratory with a gas chromatograph. Statistical response functions for CO₂ dynamics were constructed to evaluate the annual CO₂ exchange from the climatic data. The net CO₂ exchange was calculated for every hour in the snow‐free season. The carbon balance varied extensively depending on the weather conditions, and type and phenology of vegetation. During the growing season, the grassland was a net source while the barley field was a net sink for CO₂. However, both soils were net sources for CO₂ when autumn, winter and spring were included also. The annual CO₂ emissions from the grassland and barley soil were 750 g CO₂‐C m^?2 and 400 g CO₂‐C m^?2, respectively. The carbon accumulated in root and shoot biomass during the growing season was 330 g m^?2 for grass and 520 g m^?2 for barley. The C in the aboveground plant biomass ranged from 43 to 47% of the carbon fixed in photosynthesis (P_G) and the proportion of C in the root biomass was 10% of the carbon fixed in photosynthesis. The bare soils had 10–60% higher net CO₂ emission than the vegetated soils. These results indicate that the carbon balance of organic soils is affected by the characteristics of the prevailing plant cover. The dry summer of 1997 may have limited the growth of grass in the late summer thus reducing photosynthesis, which could be one reason for the high CO₂ release from this grass field.     

Persistent high temperature and low precipitation reduce peat carbon accumulation

Extreme climate events are predicted to become more frequent and intense. Their ecological impacts, particularly on carbon cycling, can differ in relation to ecosystem sensitivity. Peatlands, being characterized by peat accumulation under waterlogged conditions, can be particularly sensitive to climate extremes if the climate event increases soil oxygenation. However, a mechanistic understanding of peatland responses to persistent climate extremes is still lacking, particularly in terms of aboveground–belowground feedback. Here, we present the results of a transplantation experiment of peat mesocosms from high to low altitude in order to simulate, during 3 years, a mean annual temperature c. 5 °C higher and a mean annual precipitation c. 60% lower. Specifically, we aim at understanding the intensity of changes for a set of biogeochemical processes and their feedback on carbon accumulation. In the transplanted mesocosms, plant productivity showed a species‐specific response depending on plant growth forms, with a significant decrease (c. 60%) in peat moss productivity. Soil respiration almost doubled and Q₁₀ halved in the transplanted mesocosms in combination with an increase in activity of soil enzymes. Spectroscopic characterization of peat chemistry in the transplanted mesocosms confirmed the deepening of soil oxygenation which, in turn, stimulated microbial decomposition. After 3 years, soil carbon stock increased only in the control mesocosms whereas a reduction in mean annual carbon accumulation of c. 30% was observed in the transplanted mesocosms. Based on the above information, a structural equation model was built to provide a mechanistic understanding of the causal connections between peat moisture, vegetation response, soil respiration and carbon accumulation. This study identifies, in the feedback between plant and microbial responses, the primary pathways explaining the reduction in carbon accumulation in response to recurring climate extremes in peat soils.     

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.     

Interactions among shrub cover and the soil microclimate may determine future Arctic carbon budgets

Arctic and Boreal terrestrial ecosystems are important components of the climate system because they contain vast amounts of soil carbon (C). Evidence suggests that deciduous shrubs are increasing in abundance, but the implications for ecosystem C budgets remain uncertain. Using midsummer CO₂ flux data from 21 sites spanning 16° of latitude in the Arctic and Boreal biomes, we show that air temperature explains c. one‐half of the variation in ecosystem respiration (ER) and that ER drives the pattern in net ecosystem CO₂ exchange across ecosystems. Woody sites were slightly stronger C sinks compared with herbaceous communities. However, woody sites with warm soils (> 10 °C) were net sources of CO₂, whereas woody sites with cold soils (< 10 °C) were strong sinks. Our results indicate that transition to a shrub‐dominated Arctic will increase the rate of C cycling, and may lead to net C loss if soil temperatures rise.     

Thermal Acclimation of Respiration and Photosynthesis in the Marine Macroalga Gracilaria lemaneiformis (Gracilariales,Rhodophyta)

The responses of respiration and photosynthesis to temperature fluctuations in marine macroalgae have the potential to significantly affect coastal carbon fluxes and sequestration. In this study, the marine red macroalga Gracilaria lemaneiformis was cultured at three different temperatures (12, 19, and 26°C) and at high‐ and low‐nitrogen (N) availability, to investigate the acclimation potential of respiration and photosynthesis to temperature change. Measurements of respiratory and photosynthetic rates were made at five temperatures (7°C–33°C). An instantaneous change in temperature resulted in a change in the rates of respiration and photosynthesis, and the temperature sensitivities (i.e., the Q₁₀ value) for both the metabolic processes were lower in 26°C‐grown algae than 12°C‐ or 19°C‐grown algae. Both respiration and photosynthesis acclimated to long‐term changes in temperature, irrespective of the N availability under which the algae were grown; respiration displayed strong acclimation, whereas photosynthesis only exhibited a partial acclimation response to changing growth temperatures. The ratio of respiration to gross photosynthesis was higher in 12°C‐grown algae, but displayed little difference between the algae grown at 19°C and 26°C. We propose that it is unlikely that respiration in G. lemaneiformis would increase significantly with global warming, although photosynthesis would increase at moderately elevated temperatures.     

Disentangling effects of air and soil temperature on C allocation in cold environments: A 14C pulse‐labelling study with two plant species

Carbon cycling responses of ecosystems to global warming will likely be stronger in cold ecosystems where many processes are temperature‐limited. Predicting these effects is difficult because air and soil temperatures will not change in concert, and will affect above and belowground processes differently. We disentangled above and belowground temperature effects on plant C allocation and deposition of plant C in soils by independently manipulating air and soil temperatures in microcosms planted with either Leucanthemopsis alpina or Pinus mugo seedlings. Daily average temperatures of 4 or 9°C were applied to shoots and independently to roots, and plants pulse‐labelled with ¹⁴CO₂. We traced soil CO₂ and ¹⁴CO₂ evolution for 4 days, after which microcosms were destructively harvested and ¹⁴C quantified in plant and soil fractions. In microcosms with L. alpina, net ¹⁴C uptake was higher at 9°C than at 4°C soil temperature, and this difference was independent of air temperature. In warmer soils, more C was allocated to roots at greater soil depth, with no effect of air temperature. In P. mugo microcosms, assimilate partitioning to roots increased with air temperature, but only when soils were at 9°C. Higher soil temperatures also increased the mean soil depth at which ¹⁴C was allocated. Our findings highlight the dependence of C uptake, use, and partitioning on both air and soil temperature, with the latter being relatively more important. The strong temperature‐sensitivity of C assimilate use in the roots and rhizosphere supports the hypothesis that cold limitation on C uptake is primarily mediated by reduced sink strength in the roots. We conclude that variations in soil rather than air temperature are going to drive plant responses to warming in cold environments, with potentially large changes in C cycling due to enhanced transfer of plant‐derived C to soils.     

Plant respiration and photosynthesis in global‐scale models: incorporating acclimation to temperature and CO2

To realistically simulate climate feedbacks from the land surface to the atmosphere, models must replicate the responses of plants to environmental changes. Several processes, operating at various scales, cause the responses of photosynthesis and plant respiration to temperature and CO₂ to change over time of exposure to new or changing environmental conditions. Here, we review the latest empirical evidence that short‐term responses of plant carbon exchange rates to temperature and CO₂ are modified by plant photosynthetic and respiratory acclimation as well as biogeochemical feedbacks. We assess the frequency with which these responses have been incorporated into vegetation models, and highlight recently designed algorithms that can facilitate their incorporation. Few models currently include representations of the long‐term plant responses that have been recorded by empirical studies, likely because these responses are still poorly understood at scales relevant for models. Studies show that, at a regional scale, simulated carbon flux between the atmosphere and vegetation can dramatically differ between versions of models that do and do not include acclimation. However, the realism of these results is difficult to evaluate, as algorithm development is still in an early stage, and a limited number of data are available. We provide a series of recommendations that suggest how a combination of empirical and modeling studies can produce mechanistic algorithms that will realistically simulate longer term responses within global‐scale models.     

Responses of soil respiration to elevated CO2, air warming, and changing soil water availability in a model old-field grassland

Responses of soil respiration to atmospheric and climatic change will have profound impacts on ecosystem and global carbon (C) cycling in the future. This study was conducted to examine effects on soil respiration of the concurrent driving factors of elevated atmospheric CO₂ concentration, air warming, and changing precipitation in a constructed old‐field grassland in eastern Tennessee, USA. Model ecosystems of seven old‐field species were established in open‐top chambers and treated with factorial combinations of ambient or elevated (+300 ppm) CO₂ concentration, ambient or elevated (+3 °C) air temperature, and high or low soil moisture content. During the 19‐month experimental period from June 2003 to December 2004, higher CO₂ concentration and soil water availability significantly increased mean soil respiration by 35.8% and 15.7%, respectively. The effects of air warming on soil respiration varied seasonally from small reductions to significant increases to no response, and there was no significant main effect. In the wet side of elevated CO₂ chambers, air warming consistently caused increases in soil respiration, whereas in the other three combinations of CO₂ and water treatments, warming tended to decrease soil respiration over the growing season but increase it over the winter. There were no interactive effects on soil respiration among any two or three treatment factors irrespective of time period. Treatment‐induced changes in soil temperature and moisture together explained 49%, 44%, and 56% of the seasonal variations of soil respiration responses to elevated CO₂, air warming, and changing precipitation, respectively. Additional indirect effects of seasonal dynamics and responses of plant growth on C substrate supply were indicated. Given the importance of indirect effects of the forcing factors and plant community dynamics on soil temperature, moisture, and C substrate, soil respiration response to climatic warming should not be represented in models as a simple temperature response function, and a more mechanistic representation including vegetation dynamics and substrate supply is needed.     

Modelled net carbon gain responses to climate change in boreal trees: Impacts of photosynthetic parameter selection and acclimation

Boreal forests are crucial in regulating global vegetation‐atmosphere feedbacks, but the impact of climate change on boreal tree carbon fluxes is still unclear. Given the sensitivity of global vegetation models to photosynthetic and respiration parameters, we determined how predictions of net carbon gain (C‐gain) respond to variation in these parameters using a stand‐level model (MAESTRA). We also modelled how thermal acclimation of photosynthetic and respiratory temperature sensitivity alters predicted net C‐gain responses to climate change. We modelled net C‐gain of seven common boreal tree species under eight climate scenarios across a latitudinal gradient to capture a range of seasonal temperature conditions. Physiological parameter values were taken from the literature together with different approaches for thermally acclimating photosynthesis and respiration. At high latitudes, net C‐gain was stimulated up to 400% by elevated temperatures and CO₂ in the autumn but suppressed at the lowest latitudes during midsummer under climate scenarios that included warming. Modelled net C‐gain was more sensitive to photosynthetic capacity parameters (V_cmax, J_max, Arrhenius temperature response parameters, and the ratio of J_max to V_cmax) than stomatal conductance or respiration parameters. The effect of photosynthetic thermal acclimation depended on the temperatures where it was applied: acclimation reduced net C‐gain by 10%–15% within the temperature range where the equations were derived but decreased net C‐gain by 175% at temperatures outside this range. Thermal acclimation of respiration had small, but positive, impacts on net C‐gain. We show that model simulations are highly sensitive to variation in photosynthetic parameters and highlight the need to better understand the mechanisms and drivers underlying this variability (e.g., whether variability is environmentally and/or biologically driven) for further model improvement.     

Long‐term balanced fertilization increases the soil microbial functional diversity in a phosphorus‐limited paddy soil

The influence of long‐term chemical fertilization on soil microbial communities has been one of the frontier topics of agricultural and environmental sciences and is critical for linking soil microbial flora with soil functions. In this study, 16S rRNA gene pyrosequencing and a functional gene array, geochip 4.0, were used to investigate the shifts in microbial composition and functional gene structure in paddy soils with different fertilization treatments over a 22‐year period. These included a control without fertilizers; chemical nitrogen fertilizer (N); N and phosphate (NP); N and potassium (NK); and N, P and K (NPK). Based on 16S rRNA gene data, both species evenness and key genera were affected by P fertilization. Functional gene array‐based analysis revealed that long‐term fertilization significantly changed the overall microbial functional structures. Chemical fertilization significantly increased the diversity and abundance of most genes involved in C, N, P and S cycling, especially for the treatments NK and NPK. Significant correlations were found among functional gene structure and abundance, related soil enzymatic activities and rice yield, suggesting that a fertilizer‐induced shift in the microbial community may accelerate the nutrient turnover in soil, which in turn influenced rice growth. The effect of N fertilization on soil microbial functional genes was mitigated by the addition of P fertilizer in this P‐limited paddy soil, suggesting that balanced chemical fertilization is beneficial to the soil microbial community and its functions.     

Carbon cycling in temperate grassland under elevated temperature

An increase in mean soil surface temperature has been observed over the last century, and it is predicted to further increase in the future. The effect of increased temperature on ecosystem carbon fluxes in a permanent temperate grassland was studied in a long‐term (6 years) field experiment, using multiple temperature increments induced by IR lamps. Ecosystem respiration (R‐eco) and net ecosystem exchange (NEE) were measured and modeled by a modified Lloyd and Taylor model including a soil moisture component for R‐eco (average R² of 0.78) and inclusion of a photosynthetic component based on temperature and radiation for NEE (R² = 0.65). Modeled NEE values ranged between 2.3 and 5.3 kg CO₂ m^?2 year^?1, depending on treatment. An increase of 2 or 3°C led to increased carbon losses, lowering the carbon storage potential by around 4 tonnes of C ha^?1 year^?1. The majority of significant NEE differences were found during night‐time compared to daytime. This suggests that during daytime the increased respiration could be offset by an increase in photosynthetic uptake. This was also supported by differences in δ¹³C and δ¹⁸O, indicating prolonged increased photosynthetic activity associated with the higher temperature treatments. However, this increase in photosynthesis was insufficient to counteract the 24 h increase in respiration, explaining the higher CO₂ emissions due to elevated temperature.     

Leaf and Soil Nutrients in a Chronosequence of Second‐Growth Forest in Central Amazonia: Implications for Restoration of Abandoned Lands

Subsistence agriculture, cattle ranching, and periodical land abandonment are common land‐use practices in Amazonia. Because changes in land use affect biogeochemical cycles, secondary forests growing after land abandonment develop at varying speeds and spatial patterns, due in part to varying nutrient dynamics. Leaf and soil nutrient concentrations can provide useful information on nutrient cycling processes and strategies of nutrient use by trees that are suitable for introduction to abandoned areas. To understand nutrient dynamics in secondary forests from different regeneration stages, as well as the importance of pioneer species in the regeneration process, we measured the concentration of macronutrients in leaves of three pioneer tree species (Vismia cayennensis, Cecropia sciadophylla, and Bellucia dichotoma) in central Amazon secondary forests. We also measured macronutrients in the topsoil under the trees. We found that type of prior land use, time since abandonment, and number of fire events were significantly correlated with the concentrations of leaf and soil macronutrients, explaining between 10 and 38% of the variation in macronutrient concentrations. The observed patterns suggest that management practices affect the processes involved in N cycling and availability. Of the three pioneer species, C. sciadophylla showed the highest nutrient resorption efficiency and the highest leaf nutritional quality. We suggest that these two features determine a higher potential of C. sciadophylla for natural regeneration and restoration of abandoned lands.     

Land use of drained peatlands: Greenhouse gas fluxes,plant production,and economics

Drained peatlands are hotspots for greenhouse gas (GHG) emissions, which could be mitigated by rewetting and land use change. We performed an ecological/economic analysis of rewetting drained fertile peatlands in a hemiboreal climate using different land use strategies over 80 years. Vegetation, soil processes, and total GHG emissions were modeled using the CoupModel for four scenarios: (1) business as usual—Norway spruce with average soil water table of ?40 cm; (2) willow with groundwater at ?20 cm; (3) reed canary grass with groundwater at ?10 cm; and (4) a fully rewetted peatland. The predictions were based on previous model calibrations with several high‐resolution datasets consisting of water, heat, carbon, and nitrogen cycling. Spruce growth was calibrated by tree‐ring data that extended the time period covered. The GHG balance of four scenarios, including vegetation and soil, were 4.7, 7.1, 9.1, and 6.2 Mg CO₂eq ha^?1 year^?1, respectively. The total soil emissions (including litter and peat respiration CO₂ + N₂O + CH₄) were 33.1, 19.3, 15.3, and 11.0 Mg CO₂eq ha^?1 year^?1, respectively, of which the peat loss contributed 35%, 24%, and 7% of the soil emissions for the three drained scenarios, respectively. No peat was lost for the wet peatland. It was also found that draining increases vegetation growth, but not as drastically as peat respiration does. The cost–benefit analysis (CBA) is sensitive to time frame, discount rate, and carbon price. Our results indicate that the net benefit was greater with a somewhat higher soil water table and when the peatland was vegetated with willow and reed canary grass (Scenarios 2 and 3). We conclude that saving peat and avoiding methane release using fairly wet conditions can significantly reduce GHG emissions, and that this strategy should be considered for land use planning and policy‐making.     

Model-data synthesis of diurnal and seasonal CO2 fluxes at Niwot Ridge, Colorado

Eddy covariance records hold great promise for understanding the processes controlling the net ecosystem exchange of CO₂ (NEE). However, NEE is the small difference between two large fluxes: photosynthesis and ecosystem respiration. Consequently, separating NEE into its component fluxes, and determining the process‐level controls over these fluxes, is a difficult problem. In this study, we used a model‐data synthesis approach with the Simplified PnET (SIPNET) flux model to extract process‐level information from 5 years of eddy covariance data at an evergreen forest in the Colorado Rocky Mountains. SIPNET runs at a twice‐daily time step, and has two vegetation carbon pools, a single aggregated soil carbon pool, and a soil moisture submodel that models both evaporation and transpiration. By optimizing the model parameters before evaluating model‐data mismatches, we were able to probe the model structure independent of any arbitrary parameter set. In doing so, we were able to learn about the primary controls over NEE in this ecosystem, and in particular the respiration component of NEE. We also used this parameter optimization, coupled with a formal model selection criterion, to investigate the effects of making hypothesis‐driven changes to the model structure. These experiments lent support to the hypotheses that (1) photosynthesis, and possibly foliar respiration, are down‐regulated when the soil is frozen and (2) the metabolic processes of soil microbes vary in the summer and winter, possibly because of the existence of distinct microbial communities at these two times. Finally, we found that including water vapor fluxes, in addition to carbon fluxes, in the parameter optimization did not yield significantly more information about the partitioning of NEE into gross photosynthesis and ecosystem respiration.     

Responses of Ecosystem Carbon Cycling to Climate Change Treatments Along an Elevation Gradient

Global temperature increases and precipitation changes are both expected to alter ecosystem carbon (C) cycling. We tested responses of ecosystem C cycling to simulated climate change using field manipulations of temperature and precipitation across a range of grass-dominated ecosystems along an elevation gradient in northern Arizona. In 2002, we transplanted intact plant?Csoil mesocosms to simulate warming and used passive interceptors and collectors to manipulate precipitation. We measured daytime ecosystem respiration (ER) and net ecosystem C exchange throughout the growing season in 2008 and 2009. Warming generally stimulated ER and photosynthesis, but had variable effects on daytime net C exchange. Increased precipitation stimulated ecosystem C cycling only in the driest ecosystem at the lowest elevation, whereas decreased precipitation showed no effects on ecosystem C cycling across all ecosystems. No significant interaction between temperature and precipitation treatments was observed. Structural equation modeling revealed that in the wetter-than-average year of 2008, changes in ecosystem C cycling were more strongly affected by warming-induced reduction in soil moisture than by altered precipitation. In contrast, during the drier year of 2009, warming induced increase in soil temperature rather than changes in soil moisture determined ecosystem C cycling. Our findings suggest that warming exerted the strongest influence on ecosystem C cycling in both years, by modulating soil moisture in the wet year and soil temperature in the dry year.     

Water regulated effects of photosynthetic substrate supply on soil respiration in a semiarid steppe

Soil respiration is an important part of the global carbon (C) cycle and the largest component of C flux from terrestrial ecosystems to the atmosphere. Here, we investigated possible effects of photosynthetic substrate supply on soil respiration in a semiarid ecosystem. A field experiment combining water addition and shading (low and high shading) treatments was conducted to manipulate photosynthetic substrate supply in a temperate semiarid steppe in two growing seasons. Our result showed that water addition and/or low shading significantly increased net primary productivity (ecosystem‐level photosynthetic substrate supply) and soil respiration in both two growing seasons. However, the effects of high shading on net primary productivity and soil respiration depended on soil water condition, which were negative in wet year (2008) but positive in dry year (2009). On the diel timescale, soil respiration was out of phase with soil temperature and leaf net photosynthesis, but in phase with leaf sugar and starch contents (leaf‐level photosynthetic substrate production). The results indicated that photosynthetic substrate supply was an important factor in regulating soil respiration on both daily and seasonal timescales. Moreover, its effect on soil respiration increased with increasing water availability in this region. The predominant role of C assimilate supply on soil respiration indicates that the predicted positive influence of rising temperature on soil respiration will be simultaneously mediated by substrate supply and water availability in semiarid steppe ecosystems.     

Soil warming alters microbial substrate use in alpine soils

Will warming lead to an increased use of older soil organic carbon (SOC) by microbial communities, thereby inducing C losses from C‐rich alpine soils? We studied soil microbial community composition, activity, and substrate use after 3 and 4 years of soil warming (+4 °C, 2007–2010) at the alpine treeline in Switzerland. The warming experiment was nested in a free air CO₂ enrichment experiment using depleted ¹³CO₂ (δ¹³C = ?30‰, 2001–2009). We traced this depleted ¹³C label in phospholipid fatty acids (PLFA) of the organic layer (0–5 cm soil depth) and in C mineralized from root‐free soils to distinguish substrate ages used by soil microorganisms: fixed before 2001 (‘old’), from 2001 to 2009 (‘new’) or in 2010 (‘recent’). Warming induced a sustained stimulation of soil respiration (+38%) without decline in mineralizable SOC. PLFA concentrations did not reveal changes in microbial community composition due to soil warming, but soil microbial metabolic activity was stimulated (+66%). Warming decreased the amount of new and recent C in the fungal biomarker 18:2ω6,9 and the amount of new C mineralized from root‐free soils, implying a shift in microbial substrate use toward a greater use of old SOC. This shift in substrate use could indicate an imbalance between C inputs and outputs, which could eventually decrease SOC storage in this alpine ecosystem.