Inter‐annual changes in detritus‐based food chains can enhance plant growth response to elevated atmospheric CO2

Elevated atmospheric CO₂ generally enhances plant growth, but the magnitude of the effects depend, in part, on nutrient availability and plant photosynthetic pathway. Due to their pivotal role in nutrient cycling, changes in abundance of detritivores could influence the effects of elevated atmospheric CO₂ on essential ecosystem processes, such as decomposition and primary production. We conducted a field survey and a microcosm experiment to test the influence of changes in detritus‐based food chains on litter mass loss and plant growth response to elevated atmospheric CO₂ using two wetland plants: a C₃ sedge (Scirpus olneyi) and a C₄ grass (Spartina patens). Our field study revealed that organism's sensitivity to climate increased with trophic level resulting in strong inter‐annual variation in detritus‐based food chain length. Our microcosm experiment demonstrated that increased detritivore abundance could not only enhance decomposition rates, but also enhance plant growth of S. olneyi in elevated atmospheric CO₂ conditions. In contrast, we found no evidence that changes in the detritus‐based food chains influenced the growth of S. patens. Considered together, these results emphasize the importance of approaches that unite traditionally subdivided food web compartments and plant physiological processes to understand inter‐annual variation in plant production response to elevated atmospheric CO_2.     

Direct and indirect effects of elevated atmospheric CO2 on net ecosystem production in a Chesapeake Bay tidal wetland

The rapid increase in atmospheric CO₂ concentrations (C_a) has resulted in extensive research efforts to understand its impact on terrestrial ecosystems, especially carbon balance. Despite these efforts, there are relatively few data comparing net ecosystem exchange of CO₂ between the atmosphere and the biosphere (NEE), under both ambient and elevated C_a. Here we report data on annual sums of CO₂ (NEE_net) for 19 years on a Chesapeake Bay tidal wetland for Scirpus olneyi (C₃ photosynthetic pathway)‐ and Spartina patens (C₄ photosynthetic pathway)‐dominated high marsh communities exposed to ambient and elevated C_a (ambient + 340 ppm). Our objectives were to (i) quantify effects of elevated C_a on seasonally integrated CO₂ assimilation (NEE_net = NEE_day + NEE_night, kg C m^?2 y^?1) for the two communities; and (ii) quantify effects of altered canopy N content on ecosystem photosynthesis and respiration. Across all years, NEE_net averaged 1.9 kg m^?2 y^?1 in ambient C_a and 2.5 kg m^?2 y^?1 in elevated C_a, for the C₃‐dominated community. Similarly, elevated C_a significantly (P < 0.01) increased carbon uptake in the C₄‐dominated community, as NEE_net averaged 1.5 kg m^?2 y^?1 in ambient C_a and 1.7 kg m^?2 y^?1 in elevated C_a. This resulted in an average CO₂ stimulation of 32% and 13% of seasonally integrated NEE_net for the C₃‐ and C₄‐dominated communities, respectively. Increased NEE_day was correlated with increased efficiencies of light and nitrogen use for net carbon assimilation under elevated C_a, while decreased NEE_night was associated with lower canopy nitrogen content. These results suggest that rising C_a may increase carbon assimilation in both C₃‐ and C₄‐dominated wetland communities. The challenge remains to identify the fate of the assimilated carbon.     

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.     

Salinity and sea level mediate elevated CO2 effects on C3–C4 plant interactions and tissue nitrogen in a Chesapeake Bay tidal wetland

Elevated atmospheric carbon dioxide concentrations ([CO₂]) generally increase plant photosynthesis in C₃ species, but not in C₄ species, and reduce stomatal conductance in both C₃ and C₄ plants. In addition, tissue nitrogen concentration ([N]) often fails to keep pace with enhanced carbon gain under elevated CO₂, particularly in C₃ species. While these responses are well documented in many species, implications for plant growth and nutrient cycling in native ecosystems are not clear. Here we present data on 18 years of measurement of above and belowground biomass, tissue [N] and total standing crop of N for a Scirpus olneyi‐dominated (C₃ sedge) community, a Spartina patens‐dominated (C₄ grass) community and a C₃–C₄‐mixed species community exposed to ambient and elevated (ambient +340 ppm) atmospheric [CO₂] in natural salinity and sea level conditions of a Chesapeake Bay wetland. Increased biomass production (shoots plus roots) under elevated [CO₂] in the S. olneyi‐dominated community was sustained throughout the study, averaging approximately 35%, while no significant effect of elevated [CO₂] was found for total biomass in the C₄‐dominated community. We found a significant decline in C₄ biomass (correlated with rising sea level) and a concomitant increase in C₃ biomass in the mixed community. This shift from C₄ to C₃ was accelerated by the elevated [CO₂] treatment. The elevated [CO₂] stimulation of total biomass accumulation was greatest during rainy, low salinity years: the average increase above the ambient treatment during the three wettest years (1994, 1996, 2003) was 2.9 t ha⁻¹ but in the three driest years (1995, 1999, 2002), it was 1.2 t ha⁻¹. Elevated [CO₂] depressed tissue [N] in both species, but especially in the S. olneyi where the relative depression was positively correlated with salinity and negatively related with the relative enhancement of total biomass production. Thus, the greatest amount of carbon was added to the S. olneyi‐dominated community during years when shoot [N] was reduced the most, suggesting that the availability of N was not the most or even the main limitation to elevated [CO₂] stimulation of carbon accumulation in this ecosystem.     

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.     

Short-term decomposition of litter produced by plants grown in ambient and elevated atmospheric CO2 concentrations

The effects of elevated atmospheric CO₂ (ambient + 340 μmol mol^–1) on above-ground litter decomposition were investigated over a 6-week period using a field-based mesocosm system. Soil respiratory activity in mesocosms incubated in ambient and elevated atmospheric CO₂ concentrations were not significantly different (t-test, P > 0.05) indicating that there were no direct effects of elevated atmospheric CO₂ on litter decomposition. A study of the indirect effects of CO₂ on soil respiration showed that soil mesocosms to which naturally senescent plant litter had been added (0.5% w/w) from the C₃ sedge Scirpus olneyi grown in elevated atmospheric CO₂ was reduced by an average of 17% throughout the study when compared to soil mesocosms to which litter from Scirpus olneyi grown in ambient conditions had been added. In contrast, similar experiments using senescent material from the C₄ grass Spartina patens showed no difference in soil respiration rates between mesocosms to which litter from plants grown in elevated or ambient CO₂ conditions had been added. Analysis of the C:N ratio and lignin content of the senescent material showed that, while the C:N ratio and lignin content of the Spartina patens litter did not vary with atmospheric CO₂ conditions, the C:N ratio (but not the lignin content) of the litter from Scirpus olneyi was significantly greater (t-test;P < 0.05) when derived from plants grown under elevated CO₂ (105:1 compared to 86:1 for litter derived from Scirpus olneyi grown under ambient conditions). The results suggest that the increased C:N ratio of the litter from the C₃ plant Scirpus olneyi grown under elevated CO₂ led to the lower rates of biodegradation observed as reduced soil respiration in the mesocosms. Further long-term experiments are now required to determine the effects of elevated CO₂ on C partitioning in terrestrial ecosystems.     

Net uptake of atmospheric CO2 by coastal submerged aquatic vegetation

‘Blue Carbon’, which is carbon captured by marine living organisms, has recently been highlighted as a new option for climate change mitigation initiatives. In particular, coastal ecosystems have been recognized as significant carbon stocks because of their high burial rates and long‐term sequestration of carbon. However, the direct contribution of Blue Carbon to the uptake of atmospheric CO₂ through air‐sea gas exchange remains unclear. We performed in situ measurements of carbon flows, including air‐sea CO₂ fluxes, dissolved inorganic carbon changes, net ecosystem production, and carbon burial rates in the boreal (Furen), temperate (Kurihama), and subtropical (Fukido) seagrass meadows of Japan from 2010 to 2013. In particular, the air‐sea CO₂ flux was measured using three methods: the bulk formula method, the floating chamber method, and the eddy covariance method. Our empirical results show that submerged autotrophic vegetation in shallow coastal waters can be functionally a sink for atmospheric CO_2. This finding is contrary to the conventional perception that most near‐shore ecosystems are sources of atmospheric CO₂. The key factor determining whether or not coastal ecosystems directly decrease the concentration of atmospheric CO₂ may be net ecosystem production. This study thus identifies a new ecosystem function of coastal vegetated systems; they are direct sinks of atmospheric CO₂.     

Evapotranspiration and water use efficiency in a Chesapeake Bay wetland under carbon dioxide enrichment

Wetlands evapotranspire more water than other ecosystems, including agricultural, forest and grassland ecosystems. However, the effects of elevated atmospheric carbon dioxide (CO₂) concentration (C_a) on wetland evapotranspiration (ET) are largely unknown. Here, we present data on 12 years of measurements of ET, net ecosystem CO₂ exchange (NEE), and ecosystem water use efficiency (EWUE, i.e. NEE/ET) at 13:00–15:00 hours in July and August for a Scirpus olneyi (C3 sedge) community and a Spartina patens (C4 grass) community exposed to ambient and elevated (ambient+340 μmol mol^?1) C_a in a Chesapeake Bay wetland. Although a decrease in stomatal conductance at elevated C_a in the S. olneyi community was counteracted by an increase in leaf area index (LAI) to some extend, ET was still reduced by 19% on average over 12 years. In the S. patens community, LAI was not affected by elevated C_a and the reduction of ET was 34%, larger than in the S. olneyi community. For both communities, the relative reduction in ET by elevated C_a was directly proportional to precipitation due to a larger reduction in stomatal conductance in the control plants as precipitation decreased. NEE was stimulated about 36% at elevated C_a in the S. olneyi community but was not significantly affected by elevated C_a in S. patens community. A negative correlation between salinity and precipitation observed in the field indicated that precipitation affected ET through altered salinity and interacted with growth C_a. This proposed mechanism was supported by a greenhouse study that showed a greater C_a effect on ET in controlled low salinity conditions compared with high salinity. In spite of the differences between the two communities in their responses to elevated C_a, EWUE was increased about 83% by elevated C_a in both the S. olneyi and S. patens communities. These findings suggest that rising C_a could have significant impacts on the hydrologic cycles of coastal wetlands.     

闽江河口芦苇沼泽和短叶茳芏沼泽生态系统含碳温室气体的年收支

河口感潮沼泽是全球重要的蓝碳生态系统,具有很强的固碳能力。碳收支研究是量化生态系统碳源/汇过程及固碳规模的基础。本研究运用透明箱和不同遮光率布遮盖+红外气体分析仪/气相色谱相结合的方法,模拟不同光照条件,测定闽江河口鳝鱼滩半咸水芦苇沼泽和短叶茳芏沼泽的瞬时净生态系统二氧化碳（CO₂）交换量（net ecosystem exchange,NEE）、生态系统呼吸（ecosystem respiration,ER）以及甲烷（CH₄）排放通量,并通过对总光合吸收量（gross ecosystem exchange,GEE）与光合有效辐射的拟合以及ER与气温的拟合,外推2个沼泽生态系统CO₂气体在月、年尺度上的NEE和ER,评估其年固碳量。2个沼泽生态系统的NEE和ER均具有明显的季节变化,春夏秋季为大气中CO₂的汇,而冬季则转化为大气中CO₂的源,芦苇沼泽年尺度固碳能力显著高于短叶茳芏沼泽。芦苇沼泽与短叶茳芏沼泽CH₄排放通量差异不显著。综合考虑CH₄排放,闽江河口鳝鱼滩半咸水芦苇沼泽、短叶茳芏沼泽生态系统年固碳量分别为（5371.52±336.97） g CO₂-eq/m²和（2730.32±503.67） g CO₂-eq/m²。研究表明：闽江河口半咸水沼泽湿地在年尺度上是一个较强的碳汇,在缓解全球变暖方面发挥着重要的角色。     

Growth and senescence in plant communities exposed to elevated CO2 concentrations on an estuarine marsh

Summary Three high marsh communities on the Chesapeake Bay were exposed to a doubling in ambient CO₂ concentration for one growing season. Open-top chambers were used to raise CO₂ concentrations ca. 340 ppm above ambient over monospecific communities of Scirpus olneyi (C₃) and Spartina patens (C₄), and a mixed community of S. olneyi, S. patens, and Distichlis spicata (C₄). Plant growth and senescence were monitored by serial, nondestructive censuses. Elevated CO₂ resulted in increased shoot densities and delayed sensecence in the C₃ species. This resulted in an increase in primary productivity in S. olneyi growing in both the pure and mixed communities. There was no effect of CO₂ on growth in the C₄ species. These results demonstrate that elevated atmospheric CO₂ can cause increased aboveground production in a mature, unmanaged ecosystem.     

Insects and fungi on a C3 sedge and a C4 grass exposed to elevated atmospheric CO2 concentrations in open-top chambers in the field

The effects of elevated atmospheric CO₂ concentration on plant-fungi and plant-insect interactions were studied in an emergent marsh in the Chesapeake Bay. Stands of the C₃ sedge Scirpus olneyi Grey, and the C₄ grass Spartina patens (Ait.) Muhl. have been exposed to elevated atmospheric CO₂ concentrations during each growing season since 1987. In August 1991 the severities of fungal infections and insect infestations were quantified. Shoot nitrogen concentration ([N]) and water content (WC) were determined. In elevated concentrations of atmospheric CO₂, 32% fewer S. olneyi plants were infested by insects, and there was a 37% reduction in the severity of a pathogenic fungal infection, compared with plants grown in ambient CO₂ concentrations. S. olneyi also had reduced [N], which correlated positively with the severities of fungal infections and insect infestations. Conversely, S. patens had increased WC but unchanged [N] in elevated concentrations of atmospheric CO₂ and the severity of fungal infection increased. Elevated atmospheric CO₂ concentration increased or decreased the severity of fungal infection depending on at least two interacting factors, [N] and WC; but it did not change the number of plants that were infected with fungi. In contrast, the major results for insects were that the number of plants infected with insects decreased, and that the amount of tissue that each insect ate also decreased.     

Relationship of soil hydrogen sulfide level to net carbon assimilation ofPanicum hemitomon andSpartina patens

Panicum hemitomon Schult andSpartina patens (Ait) Muhl. plants from Louisiana Gulf Coast fresh and brackish marshes were subjected to hydrogen sulfide under controlled sediment redox conditions. Net carbon assimilation responses of both species to the combined sediment anaerobiosis and hydrogen sulfide concentrations was measured.Panicum hemitomon was more sensitive to hydrogen sulfide as compared toSpartina patens. Initiation of reduction in net carbon assimilation inP. hemitomon began when H₂S concentrations of soil solution exceeded 0.22 mgl^-1. Reductions in net carbon assimilation inS. patens were also noted at H₂S concentrations exceeding 0.34 mgl^-1. The reduction in net carbon assimilation of both species measured at elevated H₂S concentrations suggests that extreme anaerobiosis and elevated sulfide could contribute to the growth reduction of these species under certain conditions. However based on H₂S concentration in fresh and brackish marsh soil profiles, levels were too low to cause any adverse effects ofPanicum hemitomon. In brackish marsh soils containing hydrogen sulfide of 3.4 mgl^-1 in soil solution, sulfide could be a major factor limiting growth ofS. patens.     

Effects of Elevated CO2 and Nitrogen Deposition on Ecosystem Carbon Fluxes on the Sanjiang Plain Wetland in Northeast China

Background

Increasing atmospheric CO₂ and nitrogen (N) deposition across the globe may affect ecosystem CO₂ exchanges and ecosystem carbon cycles. Additionally, it remains unknown how increased N deposition and N addition will alter the effects of elevated CO₂ on wetland ecosystem carbon fluxes.

Methodology/Principal Findings

Beginning in 2010, a paired, nested manipulative experimental design was used in a temperate wetland of northeastern China. The primary factor was elevated CO₂, accomplished using Open Top Chambers, and N supplied as NH₄NO₃ was the secondary factor. Gross primary productivity (GPP) was higher than ecosystem respiration (ER), leading to net carbon uptake (measured by net ecosystem CO₂ exchange, or NEE) in all four treatments over the growing season. However, their magnitude had interannual variations, which coincided with air temperature in the early growing season, with the soil temperature and with the vegetation cover. Elevated CO₂ significantly enhanced GPP and ER but overall reduced NEE because the stimulation caused by the elevated CO₂ had a greater impact on ER than on GPP. The addition of N stimulated ecosystem C fluxes in both years and ameliorated the negative impact of elevated CO₂ on NEE.

Conclusion/Significance

In this ecosystem, future elevated CO₂ may favor carbon sequestration when coupled with increasing nitrogen deposition.     

Carbon Dioxide Dynamics and Controls in a Deep-water Wetland on the Qinghai-Tibetan Plateau

To initially characterize the dynamics and environmental controls of CO₂, ecosystem CO₂ fluxes were measured for different vegetation zones in a deep-water wetland on the Qinghai-Tibetan Plateau during the growing season of 2002. Four zones of vegetation along a gradient from shallow to deep water were dominated, respectively by the emergent species Carex allivescens V. Krez., Scirpus distigmaticus L., Hippuris vulgaris L., and the submerged species Potamogeton pectinatus L. Gross primary production (GPP), ecosystem respiration (Re), and net ecosystem production (NEP) were markedly different among the vegetation zones, with lower Re and GPP in deeper water. NEP was highest in the Scirpus-dominated zone with moderate water depth, but lowest in the Potamogeton-zone that occupied approximately 75% of the total wetland area. Diurnal variation in CO₂ flux was highly correlated with variation in light intensity and soil temperature. The relationship between CO₂ flux and these environmental variables varied among the vegetation zones. Seasonal CO₂ fluxes, including GPP, Re, and NEP, were strongly correlated with aboveground biomass, which was in turn determined by water depth. In the early growing season, temperature sensitivity (Q₁₀) for Re varied from 6.0 to 8.9 depending on vegetation zone. Q₁₀ decreased in the late growing season. Estimated NEP for the whole deep-water wetland over the growing season was 24 g C m⁻². Our results suggest that water depth is the major environmental control of seasonal variation in CO₂ flux, whereas photosynthetic photon flux density (PPFD) controls diurnal dynamics.     

Effects of inoculation of EPS-producing Pantoea agglomerans on wheat rhizosphere aggregation

We investigated plant and soil nitrogen pools and soil processes in monospecific stands of the C₃ sedge Scirpus olneyi and the C₄ grass Spartina patens grown in the field in open top chambers in a brackish marsh on the Chesapeake Bay. Stands of S. olneyi responded to eight years of elevated CO₂, by increased rates of net ecosystem gas exchange and a large stimulation of net ecosystem production. We conducted our study in the summer of 1994 and 1995 when soil cores were collected and aboveground biomass was estimated. Nitrogen concentration in elevated CO₂ treatments was reduced 15% in stems of S. olneyi and 8% in the upper 10 cm of the soil profile. While total plant nitrogen per unit of land area remained the same between treatments, total soil nitrogen showed a non-significant tendency to decrease in the upper 10 cm of the soil profile in elevated CO₂ both years of study. A significant decrease in soil bulk density largely contributed to the observed decrease in soil nitrogen. Exchangeable nitrogen and potential denitrification rates were also reduced in elevated CO₂, but net nitrogen mineralization was unchanged by elevated CO₂ treatment in S. olneyi both years. Plants and soils in a pure stand of the C₄ grass, S. patens, showed none of these effects of elevated CO₂ treatment. Our data provides evidence of changes in nitrogen dynamics of an ecosystem exposed to elevated CO₂ for eight years; however due to the variability in these data, we cannot say if or how these changes are likely to impact the effect of rising CO₂ on primary production or carbon accumulation in this ecosystem in the future.     

Reconciling Carbon-cycle Concepts, Terminology, and Methods

Recent projections of climatic change have focused a great deal of scientific and public attention on patterns of carbon (C) cycling as well as its controls, particularly the factors that determine whether an ecosystem is a net source or sink of atmospheric carbon dioxide (CO₂). Net ecosystem production (NEP), a central concept in C-cycling research, has been used by scientists to represent two different concepts. We propose that NEP be restricted to just one of its two original definitions—the imbalance between gross primary production (GPP) and ecosystem respiration (ER). We further propose that a new term—net ecosystem carbon balance (NECB)—be applied to the net rate of C accumulation in (or loss from [negative sign]) ecosystems. Net ecosystem carbon balance differs from NEP when C fluxes other than C fixation and respiration occur, or when inorganic C enters or leaves in dissolved form. These fluxes include the leaching loss or lateral transfer of C from the ecosystem; the emission of volatile organic C, methane, and carbon monoxide; and the release of soot and CO₂ from fire. Carbon fluxes in addition to NEP are particularly important determinants of NECB over long time scales. However, even over short time scales, they are important in ecosystems such as streams, estuaries, wetlands, and cities. Recent technological advances have led to a diversity of approaches to the measurement of C fluxes at different temporal and spatial scales. These approaches frequently capture different components of NEP or NECB and can therefore be compared across scales only by carefully specifying the fluxes included in the measurements. By explicitly identifying the fluxes that comprise NECB and other components of the C cycle, such as net ecosystem exchange (NEE) and net biome production (NBP), we can provide a less ambiguous framework for understanding and communicating recent changes in the global C cycle.     

Reduced N cycling in response to elevated CO2, warming,and drought in a Danish heathland: Synthesizing results of the CLIMAITE project after two years of treatments

Field‐scale experiments simulating realistic future climate scenarios are important tools for investigating the effects of current and future climate changes on ecosystem functioning and biogeochemical cycling. We exposed a seminatural Danish heathland ecosystem to elevated atmospheric carbon dioxide (CO₂), warming, and extended summer drought in all combinations. Here, we report on the short‐term responses of the nitrogen (N) cycle after 2 years of treatments. Elevated CO₂ significantly affected aboveground stoichiometry by increasing the carbon to nitrogen (C/N) ratios in the leaves of both co‐dominant species (Calluna vulgaris and Deschampsia flexuosa), as well as the C/N ratios of Calluna flowers and by reducing the N concentration of Deschampsia litter. Belowground, elevated CO₂ had only minor effects, whereas warming increased N turnover, as indicated by increased rates of microbial NH₄⁺ consumption, gross mineralization, potential nitrification, denitrification and N₂O emissions. Drought reduced belowground gross N mineralization and decreased fauna N mass and fauna N mineralization. Leaching was unaffected by treatments but was significantly higher across all treatments in the second year than in the much drier first year indicating that ecosystem N loss is highly sensitive to changes and variability in amount and timing of precipitation. Interactions between treatments were common and although some synergistic effects were observed, antagonism dominated the interactive responses in treatment combinations, i.e. responses were smaller in combinations than in single treatments. Nonetheless, increased C/N ratios of photosynthetic tissue in response to elevated CO₂, as well as drought‐induced decreases in litter N production and fauna N mineralization prevailed in the full treatment combination. Overall, the simulated future climate scenario therefore lead to reduced N turnover, which could act to reduce the potential growth response of plants to elevated atmospheric CO₂ concentration.     

Elevated atmospheric CO2 in open top chambers increases net nitrification and potential denitrification

The control of soil nitrogen (N) availability under elevated atmospheric CO₂ is central to predicting changes in ecosystem carbon (C) storage and primary productivity. The effects of elevated CO₂ on belowground processes have so far attracted limited research and they are assumed to be controlled by indirect effects through changes in plant physiology and chemistry. In this study, we investigated the effects of a 4‐year exposure to elevated CO₂ (ambient + 400 µmol mol^?1) in open top chambers under Scots pine (Pinus sylvestris L) seedlings on soil microbial processes of nitrification and denitrification. Potential denitrification (DP) and potential N₂O emissions were significantly higher in soils from the elevated CO₂ treatment, probably regulated indirectly by the changes in soil conditions (increased pH, C availability and NO₃^– production). Net N mineralization was mainly accounted for by nitrate production. Nitrate production was significantly larger for soil from the elevated CO₂ treatment in the field when incubated in the laboratory under elevated CO₂ (increase of 100%), but there was no effect when incubated under ambient CO₂. Net nitrate production of the soil originating from the ambient CO₂ treatment in the field was not influenced by laboratory incubation conditions. These results indicate that a direct effect of elevated atmospheric CO₂ on soil microbial processes might take place. We hypothesize that physiological adaptation or selection of nitrifiers could occur under elevated CO₂ through higher soil CO₂ concentrations. Alternatively, lower microbial NH₄ assimilation under elevated CO₂ might explain the higher net nitrification. We conclude that elevated atmospheric CO₂ has a major direct effect on the soil microbial processes of nitrification and denitrification despite generally higher soil CO₂ concentrations compared to atmospheric concentrations.     

Elevated atmospheric CO2 increases fine root production, respiration, rhizosphere respiration and soil CO2 efflux in Scots pine seedlings

In this study, we investigated the impact of elevated atmospheric CO₂ (ambient + 350 μmol mol^–1) on fine root production and respiration in Scots pine (Pinus sylvestris L.) seedlings. After six months exposure to elevated CO₂, root production measured by root in-growth bags, showed significant increases in mean total root length and biomass, which were more than 100% greater compared to the ambient treatment. This increased root length may have lead to a more intensive soil exploration. Chemical analysis of the roots showed that the roots in the elevated treatment accumulated more starch and had a lower C/N-ratio. Specific root respiration rates were significantly higher in the elevated treatment and this was probably attributed to increased nitrogen concentrations in the roots. Rhizospheric respiration and soil CO₂ efflux were also enhanced in the elevated treatment. These results clearly indicate that under elevated atmospheric CO₂ root production and development in Scots pine seedlings is altered and respiratory carbon losses through the root system are increased.     

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.