Dynamics of soil CO2 efflux under varying atmospheric CO2 concentrations reveal dominance of slow processes

We evaluated the effect on soil CO₂ efflux (F_CO2) of sudden changes in photosynthetic rates by altering CO₂ concentration in plots subjected to +200 ppmv for 15 years. Five‐day intervals of exposure to elevated CO₂ (eCO₂) ranging 1.0–1.8 times ambient did not affect F_CO2. F_CO2 did not decrease until 4 months after termination of the long‐term eCO₂ treatment, longer than the 10 days observed for decrease of F_CO2 after experimental blocking of C flow to belowground, but shorter than the ~13 months it took for increase of F_CO2 following the initiation of eCO₂. The reduction of F_CO2 upon termination of enrichment (~35%) cannot be explained by the reduction of leaf area (~15%) and associated carbohydrate production and allocation, suggesting a disproportionate contraction of the belowground ecosystem components; this was consistent with the reductions in base respiration and F_CO2‐temperature sensitivity. These asymmetric responses pose a tractable challenge to process‐based models attempting to isolate the effect of individual processes on F_CO2.     

Rapid turnover of DOC in temperate forests accounts for increased CO2 production at elevated temperatures

The evidence for the contribution of soil warming to changes in atmospheric CO₂ concentrations and carbon stocks of temperate forest ecosystems is equivocal. Here, we use data from a beech/oak forest on concentrations and stable isotope ratios of dissolved organic carbon (DOC), phosphate buffer-extractable organic carbon, soil organic carbon (SOC), respiration and microbial gross assimilation of N to show that respired soil carbon originated from DOC. However, the respiration was not dependent on the DOC concentration but exceeded the daily DOC pool three to four times, suggesting that DOC was turned over several times per day. A mass flow model helped to calculate that a maximum of 40% of the daily DOC production was derived from SOC and to demonstrate that degradation of SOC is limiting respiration of DOC. The carbon flow model on SOC, DOC, microbial C mobilization/immobilization and respiration is linked by temperature-dependent microbial and enzyme activity to global warming effects of CO₂ emitted to the atmosphere.     

Soil gas fluxes of N2O,CH4 and CO2 beneath Lolium perenne under elevated CO2: The Swiss free air carbon dioxide enrichment experiment

Fluxes of nitrous oxide, methane and carbon dioxide were measured from soils under ambient (350 µL L^-1) and enhanced (600 µL L^-1) carbon dioxide partial pressures (pCO₂) at the Free Air Carbon Dioxide Enrichment (FACE) experiment, Eidgenössische Technische Hochschule (ETH), Eschikon, Switzerland in July 1995, using a GC housed in a mobile laboratory. Measurements were made in plots of Lolium perenne maintained under high N input. During the data collection period N fertiliser was applied at a rate of 14 g m^-2 of N. Elevated pCO₂ appeared to result in an increased (27%) output of N₂O, thought to be the consequence of enhanced root-derived available soil C, acting as an energy source for denitrification. The climate, agricultural practices and soils at the FACE experiment combined to give rise to some of the largest N₂O emissions recorded for any terrestrial ecosystem. The amount of CO₂–C being lost from the control plot was higher (10%) than for the enhanced CO₂ plot, and is the reverse of that predicted. The control plot oxidised consistently more CH₄ than the enhanced plot, oxidising 25.5 ± 0.8 µg m^-2 hr^-1 of CH₄ for the control plot, with an average of 8.5 ± 0.4 µg m^-2 hr^-1 of CH₄ for the enhanced CO₂ plot. This suggests that elevated pCO₂ may lead to a feedback whereby less CH₄ is removed from the atmosphere. Despite the limited nature of the current study (in time and space), the observations made here on the interactions of elevated pCO₂ and soil trace gas release suggest that significant interactions are occurring. The feedbacks involved could have importance at the global scale.     

Soil microbial biomass and respiration measurements: An automated technique based on infra-red gas analysis

An automated system for continuous soil respiration and microbial biomass measurements based on Infra Red Gas Analysis was constructed. The switching device is computer controlled and allows hourly measurements of up to 24 samples when switching intervals of 2.5 min are selected. This allows the use of the substrate-induced respiration method for biomass determination. A software package to run the system was developed.     

Contrasting effects of low and high nitrogen additions on soil CO2 flux components and ectomycorrhizal fungal sporocarp production in a boreal forest

Nitrogen (N) added through atmospheric deposition or as fertilizer to boreal and temperate forests reduces both soil decomposer activity (heterotrophic respiration) and the activity of roots and mycorrhizal fungi (autotrophic respiration). However, these negative effects have been found in studies that applied relatively high levels of N, whereas the responses to ambient atmospheric N deposition rates are still not clear. Here, we compared an unfertilized control boreal forest with a fertilized forest (100 kg N ha^?1 yr^?1) and a forest subject to N‐deposition rates comparable to those in Central Europe (20 kg N ha^?1 yr^?1) to investigate the effects of N addition rate on different components of forest floor respiration and the production of ectomycorrhizal fungal sporocarps. Soil collars were used to partition heterotrophic (R_h) and autotrophic (R_a) respiration, which was further separated into respiration by tree roots (R_tr) and mycorrhizal hyphae (R_m). Total forest floor respiration was twice as high in the low N plot compared to the control, whereas there were no differences between the control and high N plot. There were no differences in R_h respiration among plots. The enhanced forest floor respiration in the low N plot was, therefore, the result of increased R_a respiration, with an increase in R_tr respiration, and a doubling of R_m respiration. The latter was corroborated by a slightly greater ectomycorrhizal (EM) fungal sporocarp production in the low N plot as compared to the control plot. In contrast, EM fungal sporocarp production was nearly eliminated, and R_m respiration severely reduced, in the high N plot, which resulted in significantly lower R_a respiration. We thus found a nonlinear response of the R_a components to N addition rate, which calls for further studies of the quantitative relations among N addition rate, plant photosynthesis and carbon allocation, and the function of EM fungi.     

The effects of elevated [CO2] on plant-soil carbon below-ground: A summary and synthesis

We undertake a synthesis of the most relevant results from the presentations at the meeting Plant-Soil Carbon Below-Ground: The Effects of Elevated CO₂ (Oxford-UK, September 1995), many of which are published in this Special Issue. Below-ground responses to elevated [CO₂] are important because the capacity of soils for long-term carbon sequestration. We draw the following conclusions: (i) several ecosystems exposed to elevated [CO₂] showed sustained increased CO₂ uptake at the plot level for many years. A few systems, however, showed complete down-regulation of net CO₂ uptake after several years of elevated [CO₂] exposure; (ii) under elevated [CO₂], a greater proportion of fixed carbon is generally allocated below-ground, potentially increasing the capacity of below-ground sinks; and (iii) some of the increased capacity of these sinks may lead to increased long-term soil carbon sequestration, although strong evidence is still lacking. We highlight the need for more soil studies to be undertaken within ongoing ecosystem-level experiments, and suggest that while some key experiments already established should be maintained to allow long term effects and feedbacks to take place, more research effort should be directed to mechanisms of soil organic matter stabilization.     

The effects of elevated atmospheric CO2 and soil P placement on cotton root deployment

Root proliferation into nutrient rich zones is an important mechanism in the exploitation of soil nutrients by plants. No studies have examined atmospheric CO₂ effects on cotton (Gossypium hirsutum L.) root distribution as affected by localized phosphorus (P). Cotton plants were grown in a Troup sand (loamy, thermic Grossarenic Kandiudults) using 17.2-l containers placed in open top field chambers (OTC) under ambient (360 mol mol^–1) or enriched (720 mol mol^–1) atmospheric CO₂ concentrations for 40 days. Equivalent amounts of P were added (150 mg P per kg of soil) to 100, 50, 25, 12.5, and 6.25% of the total soil volume; control containers with no added P were also included. Under extremely low P (controls), cotton was unresponsive to CO₂ enrichment. In treatments with both fertilized and unfertilized soil volumes, root proliferation was greater in the unfertilized soil under elevated CO₂ conditions. Stimulation of root growth occurred in the P-fertilized soil fraction; the pattern of stimulation was similar under both CO₂ levels. Under ambient CO₂, cotton plant response was positive (shoot mass, and total root mass and length) when soil P was confined to relatively small proportions of the total soil volume (6.25 and 12.5%). However, elevated CO₂ grown plants tended to respond to P regardless of its distribution.     

The interactive effects of elevated CO2 and O3 concentration on photosynthesis in spring wheat

This study investigated the interacting effects of carbon dioxide and ozone on photosynthetic physiology in the flag leaves of spring wheat (Triticum aestivum L. cv. Wembley), at three stages of development. Plants were exposed throughout their development to reciprocal combinations of two carbon dioxide and two ozone treatments: [CO₂] at 350 or 700 mol mol^–1, [O₃] at < 5 or 60 nmol mol^–1. Gas exchange analysis, coupled spectrophotometric assay for RuBisCO activity, and SDS-PAGE, were used to examine the relative importance of pollutant effects on i) stomatal conductance, ii) quantum yield, and iii) RuBisCO activity, activation, and concentration. Independently, both elevated [CO₂] and elevated [O₃] caused a loss of RuBisCO protein and V_cmax. In combination, elevated [CO₂] partially protected against the deleterious effects of ozone. It did this partly by reducing stomatal conductance, and thereby reducing the effective ozone dose. Elevated [O₃] caused stomatal closure largely via its effect on photoassimilation.     

The space‐time continuum: the effects of elevated CO2 and temperature on trees and the importance of scaling

To predict how forests will respond to rising temperatures and atmospheric CO₂ concentrations, we need to understand how trees respond to both of these environmental factors. In this review, we discuss the importance of scaling, moving from leaf‐level responses to those of the canopy, and from short‐term to long‐term responses of vegetation to climate change. While our knowledge of leaf‐level, instantaneous responses of photosynthesis, respiration, stomatal conductance, transpiration and water‐use efficiency to elevated CO₂ and temperature is quite good, our ability to scale these responses up to larger spatial and temporal scales is less developed. We highlight which physiological processes are least understood at various levels of study, and discuss how ignoring differences in the spatial or temporal scale of a physiological process impedes our ability to predict how forest carbon and water fluxes forests will be altered in the future. We also synthesize data from the literature to show that light respiration follows a generalized temperature response across studies, and that the light compensation point of photosynthesis is reduced by elevated growth CO₂. Lastly, we emphasize the need to move beyond single factorial experiments whenever possible, and to combine both CO₂ and temperature treatments in studies of tree performance.     

A meta-analytical test of elevated CO2 effects on plant respiration

Contrasting results regarding elevated CO₂ effects on leafdark respiration (Rd) have hampered efforts to incorporate this importantcomponent of the plant carbon budget into long-term predictions of ecologicalresponses to rising atmospheric CO₂. To help resolve some of theseinconsistencies in the literature, we used meta-analysis to quantitativelysummarize 45 area-based leaf Rd (Rd_a) and 44 mass-based leaf Rd(Rd_m) observations from independent studies on 33 species. Ouranalysis showed that across all studies, leaf Rd_m was significantlyreduced (–18%, P < 0.05), while leaf Rd_a was marginallyincreased (+8%, P < 0.15), under elevated CO₂. There weresignificant differences among categorical groups in CO₂ effects onleaf Rd_a and Rd_m. For example, leaf Rd_a ofherbaceous species increased 28%, but leaf Rd_a of woody speciesremained unchanged under elevated CO₂. Plants exposed to elevatedCO₂ for < 60 days had significantly higher leaf Rd_a atelevated compared to ambient CO₂, while plants exposed to elevatedCO₂ for longer period of time showed no response. The magnitude ofreduction in leaf Rd_m for plants exposed to elevated CO₂for > 100 days was significantly greater than that for plants exposed toelevated CO₂ for < 100 days. Our meta-analysis of publishedresults suggest that the amount of carbon loss through leaf Rd will likelyincrease in a higher CO₂ environment because of higher leafRd_a and a proportionally greater leaf biomass increase than leafRd_m reduction at elevated CO₂. Our results alsodemonstrated the strong dependency of Rd responses to elevated CO₂onexperimental conditions.     

The effect of plant material grown under elevated CO2 on soil respiratory activity

The biodegradability of aerial material from a C4 plant, sorghum grown under ambient (345 µmol mol^–1) and elevated (700 µmol mol^–1) atmospheric CO₂ concentrations were compared by measuring soil respiratory activity. Initial daily respiratory activity (measured over 10 h per day) increased four fold from 110 to 440 cm³ CO₂ 100g dry weight soil^–1 in soils amended with sorghum grown under either elevated or ambient CO₂. Although soil respiratory activity decreased over the following 30 days, respiration remained significantly higher (t-test;p>0.05) in soils amended with sorghum grown under elevated CO₂ concentrations. Analysis of the plant material revealed no significant differences in C:N ratios between sorghum grown under elevated or ambient CO₂. The reason for the differences in soil respiratory activity have yet to be elucidated. However if this trend is repeated in natural ecosystems, this may have important implications for C and N cycling.     

Acclimation of photosynthesis,respiration and ecosystem carbon flux of a wetland on Chesapeake Bay,Maryland to elevated atmospheric CO2 concentration

Acclimation of photosynthesis and respiration in shoots and ecosystem carbon dioxide fluxes to rising atmospheric carbon dioxide concentration (C _a ) was studied in a brackish wetland. Open top chambers were used to create test atmospheres of normal ambient and elevated C _a (=normal ambient + 34 Pa CO₂) over mono-specific stands of the C₃ sedge Scirpus olneyi, the dominant C₃ species in the wetland ecosystem, throughout each growing season since April of 1987. Acclimation of photosynthesis and respiration were evaluated by measurements of gas exchange in excised shoots. The impact of elevated C _a on the accumulation of carbon in the ecosystem was determined by ecosystem gas exchange measurements made using the open top chamber as a cuvette.Elevated C _a increased carbohydrate and reduced Rubisco and soluble protein concentrations as well as photosynthetic capacity(A) and dark respiration (R _d ; dry weight basis) in excised shoots and canopies (leaf area area basis) of Scirpus olneyi. Nevertheless, the rate of photosynthesis was stimulated 53% in shoots and 30% in canopies growing in elevated C _a compared to normal ambient concentration. Elevated C _a inhibited R _d measured in excised shoots (–19 to –40%) and in seasonally integrated ecosystem respiration (R _e ; –36 to –57%). Growth of shoots in elevated C _a was stimulated 14–21%, but this effect was not statistically significant at peak standing biomass in midseason. Although the effect of elevated C _a on growth of shoots was relatively small, the combined effect of increased number of shoots and stimulation of photosynthesis produced a 30% stimulation in seasonally integrated gross primary production (GPP). The stimulation of photosynthesis and inhibition of respiration by elevated C _a increased net ecosystem production (NEP=GPP–R _e ) 59% in 1993 and 50% in 1994. While this study consistently showed that elevated C _a produced a significant increase in NEP, we have not identified a correspondingly large pool of carbon below ground.     

Application of a two‐pool model to soil carbon dynamics under elevated CO2

Elevated atmospheric CO₂ concentrations increase plant productivity and affect soil microbial communities, with possible consequences for the turnover rate of soil carbon (C) pools and feedbacks to the atmosphere. In a previous analysis (Van Groenigen et al., 2014), we used experimental data to inform a one‐pool model and showed that elevated CO₂ increases the decomposition rate of soil organic C, negating the storage potential of soil. However, a two‐pool soil model can potentially explain patterns of soil C dynamics without invoking effects of CO₂ on decomposition rates. To address this issue, we refit our data to a two‐pool soil C model. We found that CO₂ enrichment increases decomposition rates of both fast and slow C pools. In addition, elevated CO₂ decreased the carbon use efficiency of soil microbes (CUE), thereby further reducing soil C storage. These findings are consistent with numerous empirical studies and corroborate the results from our previous analysis. To facilitate understanding of C dynamics, we suggest that empirical and theoretical studies incorporate multiple soil C pools with potentially variable decomposition rates.     

The input and fate of new C in two forest soils under elevated CO2

The aim of this study was to estimate (i) the influence of different soil types on the net input of new C into soils under CO₂ enrichment and (ii) the stability and fate of these new C inputs in soils. We exposed young beech–spruce model ecosystems on an acidic loam and calcareous sand for 4 years to elevated CO₂. The added CO₂ was depleted in ¹³C, allowing to trace new C inputs in the plant–soil system. We measured CO₂‐derived new C in soil C pools fractionated into particle sizes and monitored respiration as well as leaching of this new C during incubation for 1 year. Soil type played a crucial role in the partitioning of C. The net input of new C into soils under elevated CO₂ was about 75% greater in the acidic loam than in the calcareous sand, despite a 100% and a 45% greater above‐ and below‐ground biomass on the calcareous sand. This was most likely caused by a higher turnover of C in the calcareous sand as indicated by 30% higher losses of new C from the calcareous sand than from the acidic loam during incubation. Therefore, soil properties determining stabilization of soil C were apparently more important for the accumulation of C in soils than tree productivity. Soil fractionation revealed that about 60% of the CO₂‐derived new soil C was incorporated into sand fractions. Low natural ¹³C abundance and wide C/N ratios show that sand fractions comprise little decomposed organic matter. Consistently, incubation indicated that new soil C was preferentially respired as CO₂. During the first month, evolved CO₂ consisted to 40–55% of new C, whereas the fraction of new C in bulk soil C was 15–23% only. Leaching of DOC accounted for 8–23% of the total losses of new soil C. The overall effects of CO₂ enrichment on soil C were small in both soils, although tree growth increased significantly on the calcareous sand. Our results suggest that the potential of soils for C sequestration is limited, because only a small fraction of new C inputs into soils will become long‐term soil C.     

Experimental soil warming effects on CO2 and CH4 flux from a low elevation spruce–fir forest soil in Maine, USA

The effect of soil warming on CO₂ and CH₄ flux from a spruce–fir forest soil was evaluated at the Howland Integrated Forest Study site in Maine, USA from 1993 to 1995. Elevated soil temperatures (～5 °C) were maintained during the snow-free season (May – November) in replicated 15 × 15-m plots using electric cables buried 1–2 cm below the soil surface; replicated unheated plots served as the control. CO₂ evolution from the soil surface and soil air CO₂ concentrations both showed clear seasonal trends and significant (P < 0.0001) positive exponential relationships with soil temperature. Soil warming caused a 25–40% increase in CO₂ flux from the heated plots compared to the controls. No significant differences were observed between heated and control plot soil air CO₂ concentrations which we attribute to rapid equilibration with the atmosphere in the O horizon and minimal treatment effects in the B horizon. Methane fluxes were highly variable and showed no consistent trends with treatment.     

Environmental productivity indices and productivity for Opuntia ficus-indica under current and elevated atmospheric CO2 levels*

Abstract. The productivity of the prickly-pear cactus Opuntia ficus-indica, which is cultivated worldwide for its fruits and stem segments, was predicted based on the responses of its net CO₂ uptake to soil water status, air temperature and photosynthetic photon flux density (PPFD). Each of these environmental factors was represented by an index with a maximum value of unity when that factor was not limiting net CO₂ uptake over a 24-h period. The water index, the temperature index, and the PPFD index were determined for 87 sites in the contiguous United States using data from 189 weather stations and for 148 sites worldwide using data from 1464 weather stations. The product of these three indices, the environmental productivity index (EPI), was used to predict the productivity of O. ficus-indica under current climatic conditions and under those accompanying a possible increase in the atmospheric CO₂ level to 650μumol mol^?1. Sites with temperatures always above -10°C and hence suitable for prickly-pear cultivation numbered 37 in the United States and 110 worldwide; such sites increased by 43 and 5%, respectively, for the global warming accompanying the elevated CO₂. Productivity of O. ficus-indica was at least 15 tonnes dry weight hectare^?1 year^?1, comparable to that of many agronomic crops, for 20 sites with temperatures always above -10°C in the contiguous United States and for 12 such sites worldwide under current climatic conditions; such sites increased by 85 and 117%, respectively, under the elevated CO₂ condition, mainly because of direct effects of the atmospheric CO₂ level on net CO₂ uptake. In summary, simulations based on EPI indicate that O. ficus-indica may presently be advantageously cultivated over a substantial fraction of the earth's surface, such regions increasing markedly with a future doubling in atmospheric CO₂ levels.     

Potential effects of three antibiotics on community respiration and production measurements

The effects of three commonly available antibiotics (tetracycline hydrochloride, streptomycin and neomycin sulfate) on the micro-Winkler and D.O. probe oxygen techniques and on the pH method of CO₂ determination were investigated in the laboratory to assess potential effects on community metabolism measurements.All three antibiotics significantly altered O₂ and CO₂ concentrations through time. The results demonstrate that the use of the three antibiotics as bacterial respiratory inhibitors in community metabolism studies could cause significant overestimations of community respiration when measured as CO₂, and significant underestimation of primary productivity and overestimation of respiration when measured as O₂.     

Legacy effects override soil properties for CO2 and N2O but not CH4 emissions following digestate application to soil

The application of organic materials to soil can recycle nutrients and increase organic matter in agricultural lands. Digestate can be used as a nutrient source for crop production but it has also been shown to stimulate greenhouse gas (GHG) emissions from amended soils. While edaphic factors, such as soil texture and pH, have been shown to be strong determinants of soil GHG fluxes, the impact of the legacy of previous management practices is less well understood. Here we aim to investigate the impact of such legacy effects and to contrast them against soil properties to identify the key determinants of soil GHG fluxes following digestate application. Soil from an already established field experiment was used to set up a pot experiment, to evaluate N₂O, CH₄ and CO₂ fluxes from cattle‐slurry‐digestate amended soils. The soil had been treated with farmyard manure, green manure or synthetic N‐fertilizer, 18 months before the pot experiment was set up. Following homogenization and a preincubation stage, digestate was added at a concentration of 250 kg total N/ha eq. Soil GHG fluxes were then sampled over a 64 day period. The digestate stimulated emissions of the three GHGs compared to controls. The legacy of previous soil management was found to be a key determinant of CO₂ and N₂O flux while edaphic variables did not have a significant effect across the range of variables included in this experiment. Conversely, edaphic variables, in particular texture, were the main determinant of CH₄ flux from soil following digestate application. Results demonstrate that edaphic factors and current soil management regime alone are not effective predictors of soil GHG flux response following digestate application. Knowledge of the site management in terms of organic amendments is required to make robust predictions of the likely soil GHG flux response following digestate application to soil.     

青藏高原高寒草甸生长期CO2排放对气温升高的响应

采用高度分别为0.4 m(OTC-1)和0.8 m(OTC-2)的开顶式增温小室(open top champers,OTCs),对青藏高原高寒草甸生态系统进行模拟增温处理,研究了高寒草甸生态系统的CO2排放强度及过程对气温升高的响应.结果表明:在平均气温分别提高1.25℃(OTC-1)和3.68℃(OTC-2)的条件下,对照及2种不同幅度增温处理高寒草甸生态系统CO2排放通量表现出明显的季节变化特征,并同时在植被生长旺盛期(7-8月)达到峰值,分别为2.31、2.35和6.38μmol·m-2·s-1;CO2排放通量在不同季节都表现出随增温幅度的升高而逐渐增大的趋势,表明随着气温升高,CO2排放通量逐渐增大;不同处理CO2排放通量与气温、5 cm土壤温度和5 cm土壤含水量之间表现出显著的相关关系,CO2排放通量与气温及5 cm土壤温度之间均符合指数关系,而与5 cm土壤含水量之间则符合二次多项式关系.     

Warming impacts on boreal fen CO2 exchange under wet and dry conditions

Northern peatlands form a major soil carbon (C) stock. With climate change, peatland C mineralization is expected to increase, which in turn would accelerate climate change. A particularity of peatlands is the importance of soil aeration, which regulates peatland functioning and likely modulates the responses to warming climate. Our aim is to assess the impacts of warming on a southern boreal and a sub‐arctic sedge fen carbon dioxide (CO₂) exchange under two plausible water table regimes: wet and moderately dry. We focused this study on minerotrophic treeless sedge fens, as they are common peatland types at boreal and (sub)arctic areas, which are expected to face the highest rates of climate warming. In addition, fens are expected to respond to environmental changes faster than the nutrient poor bogs. Our study confirmed that CO₂ exchange is more strongly affected by drying than warming. Experimental water level draw‐down (WLD) significantly increased gross photosynthesis and ecosystem respiration. Warming alone had insignificant impacts on the CO₂ exchange components, but when combined with WLD it further increased ecosystem respiration. In the southern fen, CO₂ uptake decreased due to WLD, which was amplified by warming, while at northern fen it remained stable. As a conclusion, our results suggest that a very small difference in the WLD may be decisive, whether the C sink of a fen decreases, or whether the system is able to adapt within its regime and maintain its functions. Moreover, the water table has a role in determining how much the increased temperature impacts the CO₂ exchange.