The response of heterotrophic CO2 flux to soil warming

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

Soil CO2 efflux in a boreal pine forest under atmospheric CO2 enrichment and air warming

The response of forest soil CO₂ efflux to the elevation of two climatic factors, the atmospheric concentration of CO₂ (↑CO₂ of 700 μmol mol⁻¹) and air temperature (↑ T with average annual increase of 5°C), and their combination (↑CO₂+↑ T ) was investigated in a 4-year, full-factorial field experiment consisting of closed chambers built around 20-year-old Scots pines ( Pinus sylvestris L.) in the boreal zone of Finland. Mean soil CO₂ efflux in May–October increased with elevated CO₂ by 23–37%, with elevated temperature by 27–43%, and with the combined treatment by 35–59%. Temperature elevation was a significant factor in the combined 4-year efflux data, whereas the effect of elevated CO₂ was not as evident. Elevated temperature had the most pronounced impact early and late in the season, while the influence of elevated CO₂ alone was especially notable late in the season. Needle area was found to be a significant predictor of soil CO₂ efflux, particularly in August, a month of high root growth, thus supporting the assumption of a close link between whole-tree physiology and soil CO₂ emissions. The decrease in the temperature sensitivity of soil CO₂ efflux observed in the elevated temperature treatments in the second year nevertheless suggests the existence of soil response mechanisms that may be independent of the assimilating component of the forest ecosystem. In conclusion, elevated atmospheric CO₂ and air temperature consistently increased forest soil CO₂ efflux over the 4-year period, their combined effect being additive, with no apparent interaction.     

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.     

The response of soil CO2 flux to changes in atmospheric CO2, nitrogen supply and plant diversity

We measured soil CO₂ flux over 19 sampling periods that spanned two growing seasons in a grassland Free Air Carbon dioxide Enrichment (FACE) experiment that factorially manipulated three major anthropogenic global changes: atmospheric carbon dioxide (CO₂) concentration, nitrogen (N) supply, and plant species richness. On average, over two growing seasons, elevated atmospheric CO₂ and N fertilization increased soil CO₂ flux by 0.57 µmol m^?2 s^?1 (13% increase) and 0.37 µmol m^?2 s^?1 (8% increase) above average control soil CO₂ flux, respectively. Decreases in planted diversity from 16 to 9, 4 and 1 species decreased soil CO₂ flux by 0.23, 0.41 and 1.09 µmol m^?2 s^?1 (5%, 8% and 21% decreases), respectively. There were no statistically significant pairwise interactions among the three treatments. During 19 sampling periods that spanned two growing seasons, elevated atmospheric CO₂ increased soil CO₂ flux most when soil moisture was low and soils were warm. Effects on soil CO₂ flux due to fertilization with N and decreases in diversity were greatest at the times of the year when soils were warm, although there were no significant correlations between these effects and soil moisture. Of the treatments, only the N and diversity treatments were correlated over time; neither were correlated with the CO₂ effect. Models of soil CO₂ flux will need to incorporate ecosystem CO₂ and N availability, as well as ecosystem plant diversity, and incorporate different environmental factors when determining the magnitude of the CO₂, N and diversity effects on soil CO₂ flux.     

Controls on microbial CO2 production: a comparison of surface and subsurface soil horizons

Although a significant amount of the organic C stored in soil resides in subsurface horizons, the dynamics of subsurface C stores are not well understood. The objective of this study was to determine if changes in soil moisture, temperature, and nutrient levels have similar effects on the mineralization of surface (0–25 cm) and subsurface (below 25 cm) C stores. Samples were collected from a 2 m deep unsaturated mollisol profile located near Santa Barbara, CA, USA. In a series of experiments, we measured the influence of nutrient additions (N and P), soil temperature (10–35°C), and soil water potential (?0.5 to ?10 MPa) on the microbial mineralization of native soil organic C. Surface and subsurface soils were slightly different with respect to the effects of water potential on microbial CO₂ production; C mineralization rates in surface soils were more affected by conditions of moderate drought than rates in subsurface soils. With respect to the effects of soil temperature and nutrient levels on C mineralization rates, subsurface horizons were significantly more sensitive to increases in temperature or nutrient availability than surface horizons. The mean Q₁₀ value for C mineralization rates was 3.0 in surface horizons and 3.9 in subsurface horizons. The addition of either N or P had negligible effects on microbial CO₂ production in surface soil layers; in the subsurface horizons, the addition of either N or P increased CO₂ production by up to 450% relative to the control. The results of these experiments suggest that alterations of the soil environment may have different effects on CO₂ production through the profile and that the mineralization of subsurface C stores may be particularly susceptible to increases in temperature or nutrient inputs to soil.     

The turnover of carbon pools contributing to soil CO2 and soil respiration in a temperate forest exposed to elevated CO2 concentration

Soil carbon is returned to the atmosphere through the process of soil respiration, which represents one of the largest fluxes in the terrestrial C cycle. The effects of climate change on the components of soil respiration can affect the sink or source capacity of ecosystems for atmospheric carbon, but no current techniques can unambiguously separate soil respiration into its components. Long‐term free air CO₂ enrichment (FACE) experiments provide a unique opportunity to study soil C dynamics because the CO₂ used for fumigation has a distinct isotopic signature and serves as a continuous label at the ecosystem level. We used the ¹³C tracer at the Duke Forest FACE site to follow the disappearance of C fixed before fumigation began in 1996 (pretreatment C) from soil CO₂ and soil‐respired CO₂, as an index of belowground C dynamics during the first 8 years of the experiment. The decay of pretreatment C as detected in the isotopic composition of soil‐respired CO₂ and soil CO₂ at 15, 30, 70, and 200 cm soil depth was best described by a model having one to three exponential pools within the soil system. The majority of soil‐respired CO₂ (71%) originated in soil C pools with a turnover time of about 35 days. About 55%, 50%, and 68% of soil CO₂ at 15, 30, and 70 cm, respectively, originated in soil pools with turnover times of less than 1 year. The rest of soil CO₂ and soil‐respired CO₂ originated in soil pools that turn over at decadal time scales. Our results suggest that a large fraction of the C returned to the atmosphere through soil respiration results from dynamic soil C pools that cannot be easily detected in traditionally defined soil organic matter standing stocks. Fast oxidation of labile C substrates may prevent increases in soil C accumulation in forests exposed to elevated [CO₂] and may consequently result in shorter ecosystem C residence times.     

Soil CO2 efflux in contrasting boreal deciduous and coniferous stands and its contribution to the ecosystem carbon balance

Similar nonsteady‐state automated chamber systems were used to measure and partition soil CO₂ efflux in contrasting deciduous (trembling aspen) and coniferous (black spruce and jack pine) stands located within 100 km of each other near the southern edge of the Boreal forest in Canada. The stands were exposed to similar climate forcing in 2003, including marked seasonal variations in soil water availability, which provided a unique opportunity to investigate the influence of climate and stand characteristics on soil CO₂ efflux and to quantify its contribution to the net ecosystem CO₂ exchange (NEE) as measured with the eddy‐covariance technique. Partitioning of soil CO₂ efflux between soil respiration (including forest‐floor vegetation) and forest‐floor photosynthesis showed that short‐ and long‐term temporal variations of soil CO₂ efflux were related to the influence of (1) soil temperature and water content on soil respiration and (2) below‐canopy light availability, plant water status and forest‐floor plant species composition on forest‐floor photosynthesis. Overall, the three stands were weak to moderate sinks for CO₂ in 2003 (NEE of ?103, ?80 and ?28 g C m^?2 yr^?1 for aspen, black spruce and jack pine, respectively). Forest‐floor respiration accounted for 86%, 73% and 75% of annual ecosystem respiration, in the three respective stands, while forest‐floor photosynthesis contributed to 11% and 14% of annual gross ecosystem photosynthesis in the black spruce and jack pine stands, respectively. The results emphasize the need to perform concomitant measurements of NEE and soil CO₂ efflux at longer time scales in different ecosystems in order to better understand the impacts of future interannual climate variability and vegetation dynamics associated with climate change on each component of the carbon balance.     

Potential effects of interaction between CO2 and temperature on forest landscape response to global warming

Projected temperature increases under global warming could benefit southern tree species by providing them the optimal growing temperature and could be detrimental to northern species by exposing them to the supra optimal growing temperatures. This benefit-detriment trade-off could increase the competitive advantage of southern species in the northern species range and cause the increase or even dominance of southern species in the northern domain. However, the optimum temperature for photosynthesis of C3 plants may increase due to CO₂ enrichment. An increase in the optimum temperature could greatly reduce the benefit-detriment effect. In this study, we coupled a forest ecosystem process model (PnET-II) and a forest GAP model (LINKAGES) with a spatially dynamic forest landscape model (LANDIS-II) to study how an optimum temperature increase could affect forest landscape response due to global warming. We simulated 360 years of forest landscape change in the Boundary Water Canoe Area (BWCA) in northern Minnesota, which is transitional between boreal and temperate forest. Our results showed that, under the control scenario of continuing the historic 1984–1993 mean climate (mainly temperature, precipitation and CO₂), the BWCA will become a spruce-fir dominated boreal forest. However, under the scenario of predicted climatic change [the 2000–2099 climates are predicted by Canadian Climate Center (CCC), followed by 200 years of continuing the predicted 2090–2099 mean climate], the BWCA will become a pine-dominated mixed forest. If the optimum temperature increases gradually with [CO₂] (the increase in optimum temperature is assumed to change gradually from 0 °C in year 2000 to 5 °C in year 2099 when [CO₂] reaches 711 ppm and stabilizes at 5 °C after year 2099), the BWCA would remain a fir-dominated boreal forest in areas with relatively high water-holding capacity, but not in areas with relatively low water-holding capacity. Our results suggest that the [CO₂] induced increases in optimum temperature could substantially reduce forest landscape change caused by global warming. However, not all tree species would be able to successfully adapt to future warming as predicted by CCC, regardless of optimum temperature acclimations.     

Interactions between increasing CO2 concentration and temperature on plant growth

The global environment is changing with increasing temperature and atmospheric carbon dioxide concentration, [CO₂]. Because these two factors are concomitant, and the global [CO₂] rise will affect all biomes across the full global range of temperatures, it is essential to review the theory and observations on effects of temperature and [CO₂] interactions on plant carbon balance, growth, development, biomass accumulation and yield. Although there are sound theoretical reasons for expecting a larger stimulation of net CO₂ assimilation rates by increased [CO₂] at higher temperatures, this does not necessarily mean that the pattern of biomass and yield responses to increasing [CO₂] and temperature is determined by this response. This paper reviews the interactions between the effects of [CO₂] and temperature on plants. There is little unequivocal evidence for large differences in response to [CO₂] at different temperatures, as studies are confounded by the different responses of species adapted and acclimated to different temperatures, and the interspecific differences in growth form and development pattern. We conclude by stressing the importance of initiation and expansion of meristems and organs and the balance between assimilate supply and sink activity in determining the growth response to increasing [CO₂] and temperature.     

Influence of atmospheric CO2 enrichment on methane consumption in a temperate forest soil

Rates of atmospheric CH₄ consumption of soils in temperate forest were compared in plots continuously enriched with CO₂ at 200 µL L^?1 above ambient and in control plots exposed to the ambient atmosphere of 360 µL CO₂ L^?1. The purpose was to determine if ecosystem atmospheric CO₂ enrichment would alter soil microbial CH₄ consumption at the forest floor and if the effect of CO₂ would change with time or with environmental conditions. Reduced CH₄ consumption was observed in CO₂‐enriched plots relative to control plots on 46 out of 48 sampling dates, such that CO₂‐enriched plots showed annual reductions in CH₄ consumption of 16% in 1998 and 30% in 1999. No significant differences were observed in soil moisture, temperature, pH, inorganic‐N or rates of N‐mineralization between CO₂‐enriched and control plots, indicating that differences in CH₄ consumption between treatments were likely the result of changes in the composition or size of the CH₄‐oxidizing microbial community. A repeated measures analysis of variance that included soil moisture, soil temperature (from 0 to 30 cm), and time as covariates indicated that the reduction of CH₄ consumption under elevated CO₂ was enhanced at higher soil temperatures. Additionally, the effect of elevated CO₂ on CH₄ consumption increased with time during the two‐year study. Overall, these data suggest that rising atmospheric CO₂ will reduce atmospheric CH₄ consumption in temperate forests and that the effect will be greater in warmer climates. A 30% reduction in atmospheric CH₄ consumption by temperate forest soils in response to rising atmospheric CO₂ will result in a 10% reduction in the sink strength of temperate forest soils in the atmospheric CH₄ budget and a positive feedback to the greenhouse effect.     

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.     

Net grassland carbon flux over a subambient to superambient CO2 gradient

Increasing atmospheric CO₂ concentrations may have a profound effect on the structure and function of plant communities. A previously grazed, central Texas grassland was exposed to a 200‐µmol mol^?1 to 550 µmol mol^?1 CO₂ gradient from March to mid‐December in 1998 and 1999 using two, 60‐m long, polyethylene‐ covered chambers built directly onto the site. One chamber was operated at subambient CO₂ concentrations (200–360 µmol mol^?1 daytime) and the other was regulated at superambient concentrations (360–550 µmol mol^?1). Continuous CO₂ gradients were maintained in each chamber by photosynthesis during the day and respiration at night. Net ecosystem CO₂ flux and end‐of‐year biomass were measured in each of 10, 5‐m long sections in each chamber. Net CO₂ fluxes were maximal in late May (c. day 150) in 1998 and in late August in 1999 (c. day 240). In both years, fluxes were near zero and similar in both chambers at the beginning and end of the growing season. Average daily CO₂ flux in 1998 was 13 g CO₂ m^?2 day^?1 in the subambient chamber and 20 g CO₂ m^?2 day^?1 in the superambient chamber; comparable averages were 15 and 26 g CO₂ m^?2 day^?1 in 1999. Flux was positively and linearly correlated with end‐of‐year above‐ground biomass but flux was not linearly correlated with CO₂ concentration; a finding likely to be explained by inherent differences in vegetation. Because C₃ plants were the dominant functional group, we adjusted average daily flux in each section by dividing the flux by the average percentage C₃ cover. Adjusted fluxes were better correlated with CO₂ concentration, although scatter remained. Our results indicate that after accounting for vegetation differences, CO₂ flux increased linearly with CO₂ concentration. This trend was more evident at subambient than superambient CO₂ concentrations.     

A multiyear synthesis of soil respiration responses to elevated atmospheric CO2 from four forest FACE experiments

The rapidly rising concentration of atmospheric CO₂ has the potential to alter forest and global carbon cycles by altering important processes that occur in soil. Forest soils contain the largest and longest lived carbon pools in terrestrial ecosystems and are therefore extremely important to the land–atmosphere exchange of carbon and future climate. Soil respiration is a sensitive integrator of many soil processes that control carbon storage in soil, and is therefore a good metric of changes to soil carbon cycling. Here, we summarize soil respiration data from four forest free‐air carbon dioxide enrichment (FACE) experiments in developing and established forests that have been exposed to elevated atmospheric [CO₂] (168 μL L^?1 average enrichment) for 2–6 years. The sites have similar experimental design and use similar methodology (closed‐path infrared gas analyzers) to measure soil respiration, but differ in species composition of the respective forest communities. We found that elevated atmospheric [CO₂] stimulated soil respiration at all sites, and this response persisted for up to 6 years. Young developing stands experienced greater stimulation than did more established stands, increasing 39% and 16%, respectively, averaged over all years and communities. Further, at sites that had more than one community, we found that species composition of the dominant trees was a major controller of the absolute soil CO₂ efflux and the degree of stimulation from CO₂ enrichment. Interestingly, we found that the temperature sensitivity of bulk soil respiration appeared to be unaffected by elevated atmospheric CO₂. These findings suggest that stage of stand development and species composition should be explicitly accounted for when extrapolating results from elevated CO₂ experiments or modeling forest and global carbon cycles.     

The response of two Glomus mycorrhizal fungi and a fine endophyte to elevated atmospheric CO2, soil warming and drought

Plantago lanceolata plants were grown under various environmental conditions in association with the mycorrhizal fungi Glomus mosseae, G. caledonium and a fine endophyte either individually or all together. Using a time‐course approach, we investigated the effects of elevated atmospheric CO₂ (eCO₂), soil warming and drought and their interactions on root length colonized (RLC) by mycorrhizal fungi and extraradical mycorrhizal hyphal (EMH) production. Plant growth responded as would be expected to the environmental manipulations. There was no plant growth‐independent effect of eCO₂ on mycorrhizal colonization; however, EMH production was stimulated by eCO₂, i.e. there was increased partitioning of below‐ground carbon to the EMH. Soil warming directly stimulated both percent RLC by the Glomus species and EMH density; soil warming did not affect RLC by the fine endophyte. Drought decreased percent RLC for the fine endophyte, but not for the Glomus species. The presence of one mycorrhizal fungus did not affect the response of another to the environmental variables. There was no evidence of any interactive effects of the environmental variables on RLC, but there were significant environmental interactions on EMH production. In particular, the stimulatory effects of eCO₂ and soil warming on EMH density were not additive. The results are discussed in terms of the soil carbon cycle, highlighting some crucial gaps in our knowledge. If future environmental changes affect mycorrhizal fungal turnover and respiration, then this could have important implications for the terrestrial carbon cycle.     

The sensitivity of C3 photosynthesis to increasing CO2 concentration: a theoretical analysis of its dependence on temperature and background CO2 concentration

The atmospheric CO₂ concentration has increased from the pre-industrial concentration of about 280 μmol mol⁻¹ to its present concentration of over 350 μmol mol⁻¹, and continues to increase. As the rate of photosynthesis in C₃ plants is strongly dependent on CO₂ concentration, this should have a marked effect on photosynthesis, and hence on plant growth and productivity. The magnitude of photo-synthetic responses can be calculated based on the well-developed theory of photosynthetic response to intercellular CO₂ concentration. A simple biochemically based model of photosynthesis was coupled to a model of stomatal conductance to calculate photosynthetic responses to ambient CO₂ concentration. In the combined model, photosynthesis was much more responsive to CO₂ at high than at low temperatures. At 350 μmol mol⁻¹, photosynthesis at 35°C reached 51% of the rate that would have been possible with non-limiting CO₂, whereas at 5°C, 77% of the CO₂ non-limited rate was attained. Relative CO₂ sensitivity also became smaller at elevated CO₂, as CO₂ concentration increased towards saturation. As photosynthesis was far from being saturated at the current ambient CO₂ concentration, considerable further gains in photosynthesis were predicted through continuing increases in CO₂ concentration. The strong interaction with temperature also leads to photosynthesis in different global regions experiencing very different sensitivities to increasing CO₂ concentrations.     

Water availability controls microbial temperature responses in frozen soil CO2 production

Soil processes in high-latitude regions during winter are important contributors to global carbon circulation, but our understanding of the mechanisms controlling these processes is poor and observed temperature response coefficients of CO₂ production in frozen soils deviate markedly from thermodynamically predicted responses (sometimes by several orders of magnitude). We investigated the temperature response of CO₂ production in 23 unfrozen and frozen surface soil samples from various types of boreal forests and peatland ecosystems and also measured changes in water content in them after freezing. We demonstrate that deviations in temperature responses at subzero temperatures primarily emanates from water deficiency caused by freezing of the soil water, and that the amount of unfrozen water is mainly determined by the quality of the soil organic matter, which is linked to the vegetation cover. Factoring out the contribution of water limitation to the CO₂ temperature responses yields response coefficients that agree well with expectations based on thermodynamic theory concerning biochemical temperature responses. This partitioning between a pure temperature response and the effect of water availability on the response of soil CO₂ production at low temperatures is crucial for a thorough understanding of low-temperature soil processes and for accurate predictions of C-balances in northern terrestrial ecosystems.