Elevated carbon dioxide and ozone alter productivity and ecosystem carbon content in northern temperate forests

Three young northern temperate forest communities in the north‐central United States were exposed to factorial combinations of elevated carbon dioxide (CO₂) and tropospheric ozone (O₃) for 11 years. Here, we report results from an extensive sampling of plant biomass and soil conducted at the conclusion of the experiment that enabled us to estimate ecosystem carbon (C) content and cumulative net primary productivity (NPP). Elevated CO₂ enhanced ecosystem C content by 11%, whereas elevated O₃ decreased ecosystem C content by 9%. There was little variation in treatment effects on C content across communities and no meaningful interactions between CO₂ and O₃. Treatment effects on ecosystem C content resulted primarily from changes in the near‐surface mineral soil and tree C, particularly differences in woody tissues. Excluding the mineral soil, cumulative NPP was a strong predictor of ecosystem C content (r² = 0.96). Elevated CO₂ enhanced cumulative NPP by 39%, a consequence of a 28% increase in canopy nitrogen (N) content (g N m^?2) and a 28% increase in N productivity (NPP/canopy N). In contrast, elevated O₃ lowered NPP by 10% because of a 21% decrease in canopy N, but did not impact N productivity. Consequently, as the marginal impact of canopy N on NPP (?NPP/?N) decreased through time with further canopy development, the O₃ effect on NPP dissipated. Within the mineral soil, there was less C in the top 0.1 m of soil under elevated O₃ and less soil C from 0.1 to 0.2 m in depth under elevated CO₂. Overall, these results suggest that elevated CO₂ may create a sustained increase in NPP, whereas the long‐term effect of elevated O₃ on NPP will be smaller than expected. However, changes in soil C are not well‐understood and limit our ability to predict changes in ecosystem C content.     

Net primary and ecosystem production and carbon stocks of terrestrial ecosystems and their responses to climate change

Evaluating the role of terrestrial ecosystems in the global carbon cycle requires a detailed understanding of carbon exchange between vegetation, soil, and the atmosphere. Global climatic change may modify the net carbon balance of terrestrial ecosystems, causing feedbacks on atmospheric CO₂ and climate. We describe a model for investigating terrestrial carbon exchange and its response to climatic variation based on the processes of plant photosynthesis, carbon allocation, litter production, and soil organic carbon decomposition. The model is used to produce geographical patterns of net primary production (NPP), carbon stocks in vegetation and soils, and the seasonal variations in net ecosystem production (NEP) under both contemporary and future climates. For contemporary climate, the estimated global NPP is 57.0 Gt C y^–1, carbon stocks in vegetation and soils are 640 Gt C and 1358 Gt C, respectively, and NEP varies from –0.5 Gt C in October to 1.6 Gt C in July. For a doubled atmospheric CO₂ concentration and the corresponding climate, we predict that global NPP will rise to 69.6 Gt C y^–1, carbon stocks in vegetation and soils will increase by, respectively, 133 Gt C and 160 Gt C, and the seasonal amplitude of NEP will increase by 76%. A doubling of atmospheric CO₂ without climate change may enhance NPP by 25% and result in a substantial increase in carbon stocks in vegetation and soils. Climate change without CO₂ elevation will reduce the global NPP and soil carbon stocks, but leads to an increase in vegetation carbon because of a forest extension and NPP enhancement in the north. By combining the effects of CO₂ doubling, climate change, and the consequent redistribution of vegetation, we predict a strong enhancement in NPP and carbon stocks of terrestrial ecosystems. This study simulates the possible variation in the carbon exchange at equilibrium state. We anticipate to investigate the dynamic responses in the carbon exchange to atmospheric CO₂ elevation and climate change in the past and future.     

Accelerated belowground C cycling in a managed agriforest ecosystem exposed to elevated carbon dioxide concentrations

We investigated the effects of three elevated atmospheric CO₂ levels on a Populus deltoides plantation at Biosphere 2 Laboratory in Oracle Arizona. Stable isotopes of carbon have been used as tracers to separate the carbon present before the CO₂ treatments started (old C), from that fixed after CO₂ treatments began (new C). Tree growth at elevated [CO₂] increased inputs to soil organic matter (SOM) by increasing the production of fine roots and accelerating the rate of root C turnover. However, soil carbon content decreased as [CO₂] in the atmosphere increased and inputs of new C were not found in SOM. Consequently, the rates of soil respiration increased by 141% and 176% in the 800 and 1200 μL L^?1 plantations, respectively, when compared with ambient [CO₂] after 4 years of exposure. However, the increase in decomposition of old SOM (i.e. already present when CO₂ treatments began) accounted for 72% and 69% of the increase in soil respiration seen under elevated [CO₂]. This resulted in a net loss of soil C at a rate that was between 10 and 20 times faster at elevated [CO₂] than at ambient conditions. The inability to retain new and old C in the soil may stem from the lack of stabilization of SOM, allowing for its rapid decomposition by soil heterotrophs.     

An oxygen-mediated positive feedback between elevated carbon dioxide and soil organic matter decomposition in a simulated anaerobic wetland

We examined the effects of elevated atmospheric CO₂ on soil carbon decomposition in an experimental anaerobic wetland system. Pots containing either bare C₄‐derived soil or the C₃ sedge Scirpus olneyi planted in C₄‐derived soil were incubated in greenhouse chambers at either ambient or twice‐ambient atmospheric CO₂. We measured CO₂ flux from each pot, quantified soil organic matter (SOM) mineralization using δ¹³C, and determined root and shoot biomass. SOM mineralization increased in response to elevated CO₂ by 83–218% (P<0.0001). In addition, soil redox potential was significantly and positively correlated with root biomass (P= 0.003). Our results (1) show that there is a positive feedback between elevated atmospheric CO₂ concentrations and wetland SOM decomposition and (2) suggest that this process is mediated by the release of oxygen from the roots of wetland plants. Because this feedback may occur in any wetland system, including peatlands, these results suggest a limitation on the size of the carbon sink presented by anaerobic wetland soils in a future elevated‐CO₂ atmosphere.     

Carbon balance in tussock tundra under ambient and elevated atmospheric CO2

Summary Whole ecosystem CO₂ flux under ambient (340 l/l) and elevated (680 l/l) CO₂ was measured in situ in Eriophorum tussock tundra on the North Slope of Alaska. Elevated CO₂ resulted in greater carbon acquisition than control treatments and there was a net loss of CO₂ under ambient conditions at this upland tundra site. These measurements indicate a current loss of carbon from upland tundra, possibly the result of recent climatic changes. Elevated CO₂ for the duration of one growing season appeared to delay the onset of dormancy and resulted in approximately 10 additional days of positive ecosystem flux. Homeostatic adjustment of ecosystem CO₂ flux (sum of species' response) was apparent by the third week of exposure to elevated CO₂. Ecosystem dark respiration rates were not significantly higher at elevated CO₂ levels. Rapid homeostatic adjustment to elevated CO₂ may limit carbon uptake in upland tundra. Abiotic factors were evaluated as predictors of ecosystem CO₂ flux. For chambers exposed to ambient and elevated CO₂ levels for the duration of the growing season, seasonality (Julian day) was the best predictor of ecosystem CO₂ flux at both ambient and elevated CO₂ levels. Light (PAR), soil temperature, and air temperature were also predictive of seasonal ecosystem flux, but only at elevated CO₂ levels. At any combination of physical conditions, flux of the elevated CO₂ treatment was greater than that at ambient. In short-term manipulations of CO₂, tundra exposed to elevated CO₂ had threefold greater carbon gain, and had one half the ecosystem level, light compensation point when compared to ambient CO₂ treatments. Elevated CO₂-acclimated tundra had twofold greater carbon gain compared to ambient treatments, but there was no difference in ecosystem level, light compensation point between elevated and ambient CO₂ treatments. The predicted future increases in cloudiness could substantially decrease the effect of elevated atmospheric CO₂ on net ecosystem carbon budget. These analyses suggest little if any long-term stimulation of ecosystem carbon acquisition by increases in atmospheric CO₂.     

Will changes in soil organic carbon act as a positive or negative feedback on global warming?

The world's soils contain about 1500 Gt of organic carbon to a depth of 1m and a further 900 Gt from 1--2m. A change of total soil organic carbon by just 10% would thus be equivalent to all the anthropogenic CO₂ emitted over 30 years. Warming is likely to increase both the rate of decomposition and net primary production (NPP), with a fraction of NPP forming new organic carbon. Evidence from various sources can be used to assess whether NPP or the rate of decomposition has the greater temperature sensitivity, and, hence, whether warming is likely to lead to an increase or decrease in soil organic carbon.Evidence is reviewed from laboratory-based incubations, field measurements of organic carbon storage, carbon isotope ratios and soil respiration with either naturally varying temperatures or after experimentally increasing soil temperatures. Estimates of terrestrial carbon stored at the Last Glacial Maximum are also reviewed. The review concludes that the temperature dependence of organic matter decomposition can be best described as: d(T) = exp[3.36 (T – 40)/(T + 31.79)] where d(T) is the normalised decomposition rate at temperature T (in °C). In this equation, decomposition rate is normalised to 1 at 40 °C.The review concludes by simulating the likely changes in soil organic carbon with warming. In summary, it appears likely that warming will have the effect of reducing soil organic carbon by stimulating decomposition rates more than NPP. However, increasing CO₂ is likely to simultaneously have the effect of increasing soil organic carbon through increases in NPP. Any changes are also likely to be very slow. The net effect of changes in soil organic carbon on atmospheric CO₂ loading over the next decades to centuries is, therefore, likely to be small.     

Elevated CO2 evokes quantitative and qualitative changes in carbon dynamics in a plant/soil system: mechanisms and implications

It is hypothesized that carbon storage in soil will increase under an elevated atmospheric CO₂ concentration due to a combination of an increased net CO₂ uptake, a shift in carbon allocation pattern in the plant/soil system and a decreased decomposition rate of plant residues. An overview of several studies, performed in our laboratory, on the effects of elevated CO₂ on net carbon uptake, allocation to the soil and decomposition of roots is given to test this hypothesis. The studies included wheat, ryegrass and Douglas-fir and comprised both short-term and long-term studies.Total dry weight of the plants increased up to 62%, but depended on nutrient availability. These results were supported by the data on net ¹⁴CO₂ uptake. A shift in ¹⁴C-carbon distribution from shoots to roots was found in perennial species, although this depended on nutrient availability.The decomposition experiments showed that roots cultivated at 700 L L^–1 CO₂ were decomposed more slowly than those cultivated at 350 L L^–1 CO₂. Even after two growing seasons differences up to 13% were observed, although this was found to be dependent on the nitrogen level at which the roots were grown.Both an increased carbon allocation to the soil due to an increased carbon uptake, whether or not combined with a shift in distribution pattern, and a decreased decomposition of root residues will enhance the possibilities of carbon sequestration in soil, thus supporting our hypothesis. However, nutrient availability and the response of the soil microbial biomass (size and activity) play a major role in the processes involved and require attention to clarify plant/soil responses in the long term with regard to sustained stimulation of carbon input into soils and the decomposability of roots and rhizodeposition. Soil texture will also have a strong effect on decomposition rates as a result of differences in the protecting capacity for organic matter. More detailed information on these changes is needed for a proper use of models simulating soil carbon dynamics in the long term.     

Impact of grazing intensity on seasonal variations in soil organic carbon and soil CO2 efflux in two semiarid grasslands in southern Botswana

Biological soil crusts (BSCs) are an important source of organic carbon, and affect a range of ecosystem functions in arid and semiarid environments. Yet the impact of grazing disturbance on crust properties and soil CO₂ efflux remain poorly studied, particularly in African ecosystems. The effects of burial under wind-blown sand, disaggregation and removal of BSCs on seasonal variations in soil CO₂ efflux, soil organic carbon, chlorophyll a and scytonemin were investigated at two sites in the Kalahari of southern Botswana. Field experiments were employed to isolate CO₂ efflux originating from BSCs in order to estimate the C exchange within the crust. Organic carbon was not evenly distributed through the soil profile but concentrated in the BSC. Soil CO₂ efflux was higher in Kalahari Sand than in calcrete soils, but rates varied significantly with seasonal changes in moisture and temperature. BSCs at both sites were a small net sink of C to the soil. Soil CO₂ efflux was significantly higher in sand soils where the BSC was removed, and on calcrete where the BSC was buried under sand. The BSC removal and burial under sand also significantly reduced chlorophyll a, organic carbon and scytonemin. Disaggregation of the soil crust, however, led to increases in chlorophyll a and organic carbon. The data confirm the importance of BSCs for C cycling in drylands and indicate intensive grazing, which destroys BSCs through trampling and burial, will adversely affect C sequestration and storage. Managed grazing, where soil surfaces are only lightly disturbed, would help maintain a positive carbon balance in African drylands.     

Soil carbon and nitrogen cycling and storage throughout the soil profile in a sweetgum plantation after 11 years of CO2‐enrichment

Increased partitioning of carbon (C) to fine roots under elevated [CO₂], especially deep in the soil profile, could alter soil C and nitrogen (N) cycling in forests. After more than 11 years of free‐air CO₂ enrichment in a Liquidambar styraciflua L. (sweetgum) plantation in Oak Ridge, TN, USA, greater inputs of fine roots resulted in the incorporation of new C (i.e., C with a depleted δ¹³C) into root‐derived particulate organic matter (POM) pools to 90‐cm depth. Even though production in the sweetgum stand was limited by soil N availability, soil C and N contents were greater throughout the soil profile under elevated [CO₂] at the conclusion of the experiment. Greater C inputs from fine‐root detritus under elevated [CO₂] did not result in increased net N immobilization or C mineralization rates in long‐term laboratory incubations, possibly because microbial biomass was lower in the CO₂‐enriched plots. Furthermore, the δ¹³CO₂ of the C mineralized from the incubated soil closely tracked the δ¹³C of the labile POM pool in the elevated [CO₂] treatment, especially in shallower soil, and did not indicate significant priming of the decomposition of pre‐experiment soil organic matter (SOM). Although potential C mineralization rates were positively and linearly related to total SOM C content in the top 30 cm of soil, this relationship did not hold in deeper soil. Taken together with an increased mean residence time of C in deeper soil pools, these findings indicate that C inputs from relatively deep roots under elevated [CO₂] may increase the potential for long‐term soil C storage. However, C in deeper soil is likely to take many years to accrue to a significant fraction of total soil C given relatively smaller root inputs at depth. Expanded representation of biogeochemical cycling throughout the soil profile may improve model projections of future forest responses to rising atmospheric [CO₂].     

Linking microbial activity and soil organic matter transformations in forest soils under elevated CO2

Soil organic matter (SOM) dynamics ultimately govern the ability of soil to provide long‐term C sequestration and the nutrients required for ecosystem productivity. Predicting belowground responses to elevated CO₂ requires an integrated understanding of SOM transformations and the microbial activity that governs them. It remains unclear how the microorganisms upon which these transformations depend will function in an elevated CO₂ world. This study examines SOM transformations and microbial metabolism in soils from the Duke Free Air Carbon Enrichment site in North Carolina, USA. We assessed microbial respiration and net nitrogen (N) mineralization in soils with and without elevated CO₂ exposure during a 100‐day incubation. We also traced the depleted C isotopic signature of the supplemental CO₂ into SOM and the soils' phospholipid fatty acids (PLFA), which serve as biomarkers for living cells. Cumulative net N mineralization in elevated CO₂ soils was 50% that in control soils after a 100‐day incubation. Respiration was not altered with elevated CO₂. C : N ratios of bulk SOM did not change with elevated CO₂, but incubation data suggest that the C : N ratios of mineralized organic matter increased with elevated CO₂. Values of SOM δ¹³C were depleted with elevated CO₂ (?26.7±0.2 vs. ?30.2±0.3‰), reflecting the depleted signature of the supplemental CO₂. We compared δ¹³C of individual PLFA with the δ¹³C of SOM to discern incorporation of the depleted C isotopic signature into soil microbial groups in elevated CO₂ plots. PLFA i15:0, a15:0, and 10Met18:0 reflected significant incorporation of recently produced photosynthate, suggesting that the bacterial groups defined by these biomarkers are active metabolizers in elevated CO₂ soils. At least one of these groups (actinomycetes, 10Met18:0) specializes in metabolizing less labile substrates. Because control plots did not receive an equivalent ¹³C tracer, we cannot determine from these data whether this group of organisms was stimulated by elevated CO₂ compared with these organisms in control soils. Stimulation of this group, if it occurred in the elevated CO₂ plot, would be consistent with a decline in the availability of mineralizable organic matter with elevated CO₂, which incubation data suggest may be the case in these soils.     

Soil carbon sequestration in a pine forest after 9 years of atmospheric CO2 enrichment

The impact of anthropogenic CO₂ emissions on climate change may be mitigated in part by C sequestration in terrestrial ecosystems as rising atmospheric CO₂ concentrations stimulate primary productivity and ecosystem C storage. Carbon will be sequestered in forest soils if organic matter inputs to soil profiles increase without a matching increase in decomposition or leaching losses from the soil profile, or if the rate of decomposition decreases because of increased production of resistant humic substances or greater physical protection of organic matter in soil aggregates. To examine the response of a forest ecosystem to elevated atmospheric CO₂ concentrations, the Duke Forest Free‐Air CO₂ Enrichment (FACE) experiment in North Carolina, USA, has maintained atmospheric CO₂ concentrations 200 μL L^?1 above ambient in an aggrading loblolly pine (Pinus taeda) plantation over a 9‐year period (1996–2005). During the first 6 years of the experiment, forest‐floor C and N pools increased linearly under both elevated and ambient CO₂ conditions, with significantly greater accumulations under the elevated CO₂ treatment. Between the sixth and ninth year, forest‐floor organic matter accumulation stabilized and C and N pools appeared to reach their respective steady states. An additional C sink of ～30 g C m^?2 yr^?1 was sequestered in the forest floor of the elevated CO₂ treatment plots relative to the control plots maintained at ambient CO₂ owing to increased litterfall and root turnover during the first 9 years of the study. Because we did not detect any significant elevated CO₂ effects on the rate of decomposition or on the chemical composition of forest‐floor organic matter, this additional C sink was likely related to enhanced litterfall C inputs. We also failed to detect any statistically significant treatment effects on the C and N pools of surface and deep mineral soil horizons. However, a significant widening of the C : N ratio of soil organic matter (SOM) in the upper mineral soil under both elevated and ambient CO₂ suggests that N is being transferred from soil to plants in this aggrading forest. A significant treatment × time interaction indicates that N is being transferred at a higher rate under elevated CO₂ (P=0.037), suggesting that enhanced rates of SOM decomposition are increasing mineralization and uptake to provide the extra N required to support the observed increase in primary productivity under elevated CO₂.     

Increased mercury in forest soils under elevated carbon dioxide

Fossil fuel combustion is the primary anthropogenic source of both CO₂ and Hg to the atmosphere. On a global scale, most Hg that enters ecosystems is derived from atmospheric Hg that deposits onto the land surface. Increasing concentrations of atmospheric CO₂ may affect Hg deposition to terrestrial systems and storage in soils through CO₂-mediated changes in plant and soil properties. We show, using free-air CO₂ enrichment (FACE) experiments, that soil Hg concentrations are almost 30% greater under elevated atmospheric CO₂ in two temperate forests. There were no direct CO₂ effects, however, on litterfall, throughfall or stemflow Hg inputs. Soil Hg was positively correlated with percent soil organic matter (SOM), suggesting that CO₂-mediated changes in SOM have influenced soil Hg concentrations. Through its impacts on SOM, elevated atmospheric CO₂ may increase the Hg storage capacity of soils and modulate the movement of Hg through the biosphere. Such effects of rising CO₂, ones that transcend the typically studied effects on C and nutrient cycling, are an important next phase for research on global environmental change.     

Soil–Nitrogen Cycling in a Pine Forest Exposed to 5 Years of Elevated Carbon Dioxide

Empirical and modeling studies have shown that the magnitude and duration of the primary production response to elevated carbon dioxide (CO₂) can be constrained by limiting supplies of soil nitrogen (N). We have studied the response of a southern US pine forest to elevated CO₂ for 5 years (1997–2001). Net primary production has increased significantly under elevated CO₂. We hypothesized that the increase in carbon (C) fluxes to the microbial community under elevated CO₂ would increase the rate of N immobilization over mineralization. We tested this hypothesis by quantifying the pool sizes and fluxes of inorganic and organic N in the forest floor and top 30 cm of mineral soil during the first 5 years of CO₂ fumigation. We observed no statistically significant change in the gross or net rate of inorganic N mineralization and immobilization in any soil horizon under elevated CO₂. Similarly, elevated CO₂ had no statistically significant effect on the concentration or flux of organic N, including amino acids. Microbial biomass N was not significantly different between CO₂ treatments. Thus, we reject our hypothesis that elevated CO₂ increases the rate of N immobilization. The quantity and chemistry of the litter inputs to the forest floor and mineral soil horizons can explain the limited range of microbially mediated soil–N cycling responses observed in this ecosystem. Nevertheless a comparative analysis of ecosystem development at this site and other loblolly pine forests suggests that rapid stand development and C sequestration under elevated CO₂ may be possible only in the early stages of stand development, prior to the onset of acute N limitation.     

Soil carbon storage under simulated climate change is mediated by plant functional type

The stability of soil organic matter (SOM) pools exposed to elevated CO₂ and warming has not been evaluated adequately in long‐term experiments and represents a substantial source of uncertainty in predicting ecosystem feedbacks to climate change. We conducted a 6‐year experiment combining free‐air CO₂ enrichment (FACE, 550 ppm) and warming (+2 °C) to evaluate changes in SOM accumulation in native Australian grassland. In this system, competitive interactions appear to favor C₄ over C₃ species under FACE and warming. We therefore investigated the role of plant functional type (FT) on biomass and SOM responses to the long‐term treatments by carefully sampling soil under patches of C₃‐ and C₄‐dominated vegetation. We used physical fractionation to quantify particulate organic matter (POM) and long‐term incubation to assess potential decomposition rates. Aboveground production of C₄ grasses increased in response to FACE, but total root biomass declined. Across treatments, C : N ratios were higher in leaves, roots and POM of C₄ vegetation. CO₂ and temperature treatments interacted with FT to influence SOM, and especially POM, such that soil carbon was increased by warming under C₄ vegetation, but not in combination with elevated CO₂. Potential decomposition rates increased in response to FACE and decreased with warming, possibly owing to treatment effects on soil moisture and microbial community composition. Decomposition was also inversely correlated with root N concentration, suggesting increased microbial demand for older, N‐rich SOM in treatments with low root N inputs. This research suggests that C₃–C₄ vegetation responses to future climate conditions will strongly influence SOM storage in temperate grasslands.     

Biochar stimulates the decomposition of simple organic matter and suppresses the decomposition of complex organic matter in a sandy loam soil

Incorporating crop residues and biochar has received increasing attention as tools to mitigate atmospheric carbon dioxide (CO₂) emissions and promote soil carbon (C) sequestration. However, direct comparisons between biochar, torrefied biomass, and straw on both labile and recalcitrant soil organic matter (SOM) remain poorly understood. In this study, we explored the impact of biochars produced at different temperatures and torrefied biomass on the simple C substrates (glucose, amino acids), plant residues (Lolium perenne L.), and native SOM breakdown in soil using a ¹⁴C labeling approach. Torrefied biomass and biochars produced from wheat straw at four contrasting pyrolysis temperatures (250, 350, 450, and 550 °C) were incorporated into a sandy loam soil and their impact on C turnover compared to an unamended soil or one amended with unprocessed straw. Biochar, torrefied biomass, and straw application induced a shift in the soil microbial community size, activity, and structure with the greatest effects in the straw‐amended soil. In addition, they also resulted in changes in microbial carbon use efficiency (CUE) leading to more substrate C being partitioned into catabolic processes. While overall the biochar, torrefied biomass, and straw addition increased soil respiration, it reduced the turnover rate of the simple C substrates, plant residues, and native SOM and had no appreciable effect on the turnover rate of the microbial biomass. The negative SOM priming was positively correlated with biochar production temperature. We therefore ascribe the increase in soil CO₂ efflux to biochar‐derived C rather than that originating from SOM. In conclusion, the SOM priming magnitude is strongly influenced by both the soil organic C quality and the biochar properties. In comparison with straw, biochar has the greatest potential to promote soil C storage. However, straw and torrefied biomass may have other cobenefits which may make them more suitable as a CO₂ abatement strategy.     

Effects of six years atmospheric CO2 enrichment on plant,soil, and soil microbial C of a calcareous grassland

Stimulated plant production and often even larger stimulation of photosynthesis at elevated CO₂ raise the possibility of increased C storage in plants and soils. We analysed ecosystem C partitioning and soil C fluxes in calcareous grassland exposed to elevated CO₂ for 6 years. At elevated CO₂, C pools increased in plants (+23%) and surface litter (+24%), but were not altered in microbes and soil organic matter. Soils were fractionated into particle size and density separates. The amount of low-density macroorganic C, an indicator of particulate soil C inputs from root litter, was not affected by elevated CO₂. Incorporation of C fixed during the experiment (C_new) was tracked by C isotopic analysis of soil fractions which were labelled due to ¹³C depletion of the commercial CO₂ used for atmospheric enrichment. This data constrains estimates of C sequestration (absolute upper bound) and indicates where in soils potentially sequestered C is stored. C_new entered soils at an initial rate of 210±42 g C m^–2 year^–1, but only 554±39 g C_new m^–2 were recovered after 6 years due to the low mean residence time of 1.8 years. Previous process-oriented measurements did not indicate increased plant–soil C fluxes at elevated CO₂ in the same system (¹³C kinetics in soil microbes and fine roots after pulse labelling, and minirhizotron observations). Overall experimental evidence suggests that C storage under elevated CO₂ occurred only in rapidly turned-over fractions such as plants and detritus, and that potential extra soil C inputs were rapidly re-mineralised. We argue that this inference does not conflict with the observed increases in photosynthetic fixation at elevated CO₂, because these are not good predictors of plant growth and soil C fluxes for allometric reasons. C sequestration in this natural system may also be lower than suggested by plant biomass responses to elevated CO₂ because C storage may be limited by stabilisation of C_new in slowly turned-over soil fractions (a prerequisite for long-term storage) rather than by the magnitude of C inputs per se.     

Losses of soil organic carbon under wind erosion in China

Soil organic carbon (SOC) storage generally represents the long‐term net balance of photosynthesis and total respiration in terrestrial ecosystems. However, soil erosion can affect SOC content by direct removal of soil and reduction of the surface soil depth; it also affects plant growth and soil biological activity, soil air CO₂ concentration, water regimes, soil temperature, soil respiration, carbon flux to the atmosphere, and carbon deposition in soil. In arid and semi‐arid region of northern China, wind erosion caused soil degradation and desert expansion. This paper estimated the SOC loss of the surface horizon at eroded regions based on soil property and wind erosion intensity data. The SOC loss in China because of wind erosion was about 75 Tg C yr^?1 in 1990s. The spatial pattern of SOC loss indicates that SOC loss of the surface horizon increases significantly with the increase of soil wind erosion intensity. The comparison of SOC loss and annual net primary productivity (NPP) of terrestrial ecosystem was discussed in wind erosion regions of China. We found that NPP is also low in the eroded regions and heavy SOC loss often occurs in regions where NPP is very small. However, there is potential to improve our study to resolve uncertainty on the soil organic matter oxidation and soil deposition processes in eroded and deposited sites.     

Effects of drainage and temperature on carbon balance of tussock tundra micrososms

We examined the importance of temperature (7°C or 15°C) and soil moisture regime (saturated or field capacity) on the carbon (C) balance of arctic tussock tundra microcosms (intact blocks of soil and vegetation) in growth chambers over an 81-day simulated growing season. We measured gaseous CO₂ exchanges, methane (CH₄) emissions, and dissolved C losses on intact blocks of tussock (Eriophorum vaginatum) and intertussock (moss-dominated). We hypothesized that under increased temperature and/or enhanced drainage, C losses from ecosystem respiration (CO₂ respired by plants and heterotrophs) would exceed gains from gross photosynthesis causing tussock tundra to become a net source of C to the atmosphere. The field capacity moisture regime caused a decrease in net CO₂ storage (NEP) in tussock tundra micrososms. This resulted from a stimulation of ecosystem respiration (probably mostly microbial) with enhanced drainage, rather than a decrease in gross photosynthesis. Elevated temperature alone had no effect on NEP because CO₂ losses from increased ecosystem respiration at elevated temperature were compensated by increased CO₂ uptake (gross photosynthesis). Although CO₂ losses from ecosystem respiration were primarily limited by drainage, CH₄ emissions, in contrast, were dependent on temperature. Furthermore, substantial dissolved C losses, especially organic C, and important microhabitat differences must be considered in estimating C balance for the tussock tundra system. As much as 20% of total C fixed in photosynthesis was lost as dissolved organic C. Tussocks stored 2x more C and emitted 5x more methane than intertussocks. In spite of the limitations of this microcosm experiment, this study has further elucidated the critical role of soil moisture regime and dissolved C losses in regulating net C balance of arctic tussock tundra.     

Fine Root Growth and Vertical Distribution in Response to Elevated CO<Subscript>2</Subscript>, Warming and Drought in a Mixed Heathland–Grassland

Belowground plant responses have received much less attention in climate change experiments than aboveground plant responses, thus hampering a holistic understanding of climate change effects on plants and ecosystems. In addition, responses of plant roots to climate change have mostly been studied in single-factor experiments. In a Danish heathland ecosystem, we investigated both individual and combined effects of elevated CO₂, warming and drought on fine root length, net production and standing biomass by the use of minirhizotrons, ingrowth cores and soil coring. Warming increased the net root production from ingrowth cores, but decreased fine root number and length in minirhizotrons, whereas there were no significant main effects of drought. Across all treatments and soil depths, CO₂ stimulated both the total fine root length (+44%) and the number of roots observed (+39%), with highest relative increase in root length in the deeper soil layers. Our results suggest that under future climate, plants may allocate considerable resources into roots compared to aboveground biomass. Increased carbon (C) allocation to roots may have a great impact on the overall ecosystem C balance and must be considered in modelling of future ecosystem responses to climate change. To provide models with necessary validation data, more studies are needed to investigate if higher C allocation to roots will lead to long-term C storage in more recalcitrant soil C pools or if this potential increase in soil carbon storage may be offset by increased priming activity and turnover rates for soil organic matter.     

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.