Changes in microbial activity and composition in a pasture ecosystem exposed to elevated atmospheric carbon dioxide

Elevated atmospheric CO₂ increases aboveground plant growth and productivity. However, carbon dioxide-induced alterations in plant growth are also likely to affect belowground processes, including the composition of soil biota. We investigated the influence of increased atmospheric CO₂on bacterial numbers and activity, and on soil microbial community composition in a pasture ecosystem under Free-Air Carbon Dioxide Enrichment (FACE). Composition of the soil microbial communities, in rhizosphere and bulk soil, under two atmospheric CO₂ levels was evaluated by using phospholipid fatty acid analysis (PLFA), and total and respiring bacteria counts were determined by epifluorescence microscopy. While populations increased with elevated atmospheric CO₂ in bulk soil of white clover (Trifolium repens L.), a higher atmospheric CO₂ concentration did not affect total or metabolically active bacteria in bulk soil of perennial ryegrass (Lolium perenne L.). There was no effect of atmospheric CO₂ on total bacteria populations per gram of rhizosphere soil. The combined effect of elevated CO₂ on total root length of each species and the bacterial population in these rhizospheres, however, resulted in an 85% increase in total rhizosphere bacteria and a 170% increase in respiring rhizosphere bacteria for the two plant species, when assessed on a per unit land area basis. Differences in microbial community composition between rhizosphere and bulk soil were evident in samples from white clover, and these communities changed in response to CO₂ enrichment. Results of this study indicate that changes in soil microbial activity, numbers, and community composition are likely to occur under elevated atmospheric CO₂, but the extent of those changes depend on plant species and the distance that microbes are from the immediate vicinity of the plant root surface.     

The influence of plants grown under elevated CO2 and N fertilization on soil nitrogen dynamics

We investigated the effects of spring barley growth on nitrogen (N) transformations and rhizosphere microbial processes in a controlled system under elevated carbon dioxide (CO₂) at two levels of N fertilization (applied with ¹⁵N labelling). After 25 d, elevated CO₂ (twice ambient) increased plant growth (dry weight, DW) by 141% at low‐N fertilization and by 60% at high‐N fertilization, but its positive effect on the root‐to‐shoot ratio was only significant at low‐N input. As a result of this plant response, elevated CO₂ caused a greater soil CO₂ efflux, rhizosphere soil DW, and soil microbial biomass under N‐limiting conditions than under high N availability. Elevated CO₂ also caused a significant (P < 0.001) increase in the N recovered by the plant from both the labelled (N_f) and unlabelled (N_s + N_uf) N pools. The dynamics of N in the system as affected by elevated CO₂ were driven principally by mineralization–immobilization turnover, with little loss by denitrification. Under N‐limiting conditions, there is evidence to suggest enhanced nutrient release from soil organic matter (SOM) pools—a process which could be defined as priming. The results of our experiment did not indicate a direct plant‐mediated effect of elevated CO₂ on nitrous oxide (N₂O) fluxes or denitrification activity.     

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.     

Plant rhizosphere influence on microbial C metabolism: the role of elevated CO2, N availability and root stoichiometry

Microbial decomposer C metabolism is considered a factor controlling soil C stability, a key regulator of global climate. The plant rhizosphere is now recognized as a crucial driver of soil C dynamics but specific mechanisms by which it can affect C processing are unclear. Climate change could affect microbial C metabolism via impacts on the plant rhizosphere. Using continuous ¹³C labelling under controlled conditions that allowed us to quantify SOM derived-C in all pools and fluxes, we evaluated the microbial metabolism of soil C in the rhizosphere of a C4 native grass exposed to elevated CO₂ and under variation in N concentrations in soil and in plant root C:N stoichiometry. Our results demonstrated that this plant can influence soil C metabolism and further, that elevated CO₂ conditions can alter this role by increasing microbial C efficiency as indicated by a reduction in soil-derived C respiration per unit of soil C-derived microbial biomass. Moreover, under elevated CO₂ increases in soil N, and notably, root tissue N concentration increased C efficiency, suggesting elevated CO₂ shifted the stoichiometric balance so N availability was a more critical factor regulating efficiency than under ambient conditions. The root C:N stoichiometry effect indicates that plant chemical traits such as root N concentration are able to influence the metabolism of soil C and that elevated CO₂ conditions can modulate this role. Increased efficiency in soil C use was associated with negative rhizosphere priming and we hypothesize that the widely observed phenomenon of rhizosphere priming may result, at least in part, from changes in the metabolic efficiency of microbial populations. Observed changes in the microbial community support that shifting microbial populations were a contributing factor to the observed metabolic responses. Our case study points at greater efficiency of the SOM-degrading populations in a high CO₂, high N world, potentially leading to greater C storage of microbially assimilated C in soil.     

Near-term impacts of elevated CO2, nitrogen and fungal endophyte-infection on Lolium perenne L. growth, chemical composition and alkaloid production

Carbon dioxide has been rapidly accumulating in the atmosphere and is expected to continue to do so. This accumulation is presumed to have important direct effects on plant growth. The interacting affects of a small increase in CO₂ concentration (466 p.p.m., approximately 30% increase from current ambient conditions), nitrogen fertilization and fungal endophyte (Neotyphodium lolii) infection on the growth and chemical composition of perennial ryegrass (Lolium perenne) were investigated. It was found that dry mass production was approximately 50% greater under elevated CO₂ than under ambient CO₂, but only in conditions of high soil N. High molecular weight carbohydrates and total carbohydrates (LMW + HMW CHO) depended on an interaction between CO₂ and endophyte infection. Infected plants contained significantly more carbohydrate than endophyte-free plants, and the difference was greatest in ambient CO₂ conditions. Protein concentrations were also influenced by the interaction between CO₂ and endophyte-infection. Endophyte-free plants had 40% lower concentrations of soluble protein under elevated CO₂ than under ambient CO₂, but this CO₂ effect on soluble protein was largely absent in endophyte-infected plants. CO₂, endophyte-infection and nitrogen interacted to influence the total chlorophyll concentration of the grass such that chlorophyll concentration was always lower in elevated CO₂ but this decline was much greater in endophyte-free plants, particularly in conditions of high soil N. In the endophyte-infected plants, the concentrations of the pyrrolopyrazine alkaloid peramine depended on the interaction between CO₂ and N fertilization such that peramine concentrations declined with increasing N at ambient CO₂ but remained roughly constant across N levels at elevated CO₂. A similar pattern was seen for the ergot alkaloid ergovaline. The biochemical responses of perennial ryegrass to elevated CO₂ are clearly modified by the presence of endophytic fungi.     

Changes in soil carbon and nitrogen properties and microbial communities in relation to growth of Pinus radiata and Nothofagus fusca trees after 6 years at ambient and elevated atmospheric CO2

Increasing atmospheric CO₂ concentration can influence the growth and chemical composition of many plant species, and thereby affect soil organic matter pools and nutrient fluxes. Here, we examine the effects of ambient (initially 362 μL L^?1) and elevated (654 μL L^?1) CO₂ in open‐top chambers on the growth after 6 years of two temperate evergreen forest species: an exotic, Pinus radiata D. Don, and a native, Nothofagus fusca (Hook. F.) Oerst. (red beech). We also examine associated effects on selected carbon (C) and nitrogen (N) properties in litter and mineral soil, and on microbial properties in rhizosphere and hyphosphere soil. The soil was a weakly developed sand that had a low initial C concentration of about 1.0 g kg^?1 at both 0–100 and 100–300 mm depths; in the N. fusca system, it was initially overlaid with about 50 mm of forest floor litter (predominantly FH material) taken from a Nothofagus forest. A slow‐release fertilizer was added during the early stages of plant growth; subsequent foliage analyses indicated that N was not limiting. After 6 years, stem diameters, foliage N concentrations and C/N ratios of both species were indistinguishable (P>0.10) in the two CO₂ treatments. Although total C contents in mineral soil at 0–100 mm depth had increased significantly (P<0.001) after 6 years growth of P. radiata, averaging 80±0.20 g m^?2 yr^?1, they were not significantly influenced by elevated CO₂. However, CO₂‐C production in litter, and CO₂‐C production, microbial C, and microbial C/N ratios in mineral soil (0–100 mm depth) under P. radiata were significantly higher under elevated than ambient CO₂. CO₂‐C production, microbial C, and numbers of bacteria (but not fungi) were also significantly higher under elevated CO₂ in hyphosphere soil, but not in rhizosphere soil. Under N. fusca, some incorporation of the overlaid litter into the mineral soil had probably occurred; except for CO₂‐C production and microbial C in hyphosphere soil, none of the biochemical properties or microbial counts increased significantly under elevated CO₂. Net mineral‐N production, and generally the potential utilization of different substrates by microbial communities, were not significantly influenced by elevated CO₂ under either tree species. Physiological profiles of the microbial communities did, however, differ significantly between rhizosphere and hyphosphere samples and between samples under P. radiata and N. fusca. Overall, results support the concept that a major effect on soil properties after prolonged exposure of trees to elevated CO₂ is an increase in the amounts, and mineralization rate, of labile organic components.     

Influence of an Elevated Atmospheric CO2 Content on Soil and Rhizosphere Bacterial Communities Beneath Lolium perenne and Trifolium repens under Field Conditions

Abstract The increase in atmospheric CO₂ content alters C₃ plant photosynthetic rate, leading to changes in rhizodeposition and other root activities. This may influence the activity, the biomass, and the structure of soil and rhizosphere microbial communities and therefore the nutrient cycling rates and the plant growth. The present paper focuses on bacterial numbers and on community structure. The rhizospheres of two grassland plants, Lolium perenne (ryegrass) and Trifolium repens (white clover), were divided into three fractions: the bulk soil, the rhizospheric soil, and the rhizoplane–endorhizosphere. The elevated atmospheric CO₂ content increased the most probable numbers of heterotrophic bacteria in the rhizosphere of L. perenne. However, this effect lasted only at the beginning of the vegetation period for T. repens. Community structure was assessed after isolation of DNA, PCR amplification, and construction of cloned 16S rDNA libraries. Amplified ribosomal DNA restriction analysis (ARDRA) and colony hybridization with an oligonucleotide probe designed to detect Pseudomonas spp. showed under elevated atmospheric CO₂ content an increased dominance of pseudomonads in the rhizosphere of L. perenne and a decreased dominance in the rhizosphere of T. repens. This work provides evidence for a CO₂-induced alteration in the structure of the rhizosphere bacterial populations, suggesting a possible alteration of the plant-growth-promoting-rhizobacterial (PGPR) effect. Received: 14 December 1998; Accepted: 31 March 1999     

Elevated atmospheric CO2 increases microbial growth rates in soil: results of three CO2 enrichment experiments

Increasing the belowground translocation of assimilated carbon by plants grown under elevated CO₂ can cause a shift in the structure and activity of the microbial community responsible for the turnover of organic matter in soil. We investigated the long‐term effect of elevated CO₂ in the atmosphere on microbial biomass and specific growth rates in root‐free and rhizosphere soil. The experiments were conducted under two free air carbon dioxide enrichment (FACE) systems: in Hohenheim and Braunschweig, as well as in the intensively managed forest mesocosm of the Biosphere 2 Laboratory (B2L) in Oracle, AZ. Specific microbial growth rates (μ) were determined using the substrate‐induced respiration response after glucose and/or yeast extract addition to the soil. For B2L and both FACE systems, up to 58% higher μ were observed under elevated vs. ambient CO₂, depending on site, plant species and N fertilization. The μ‐values increased linearly with atmospheric CO₂ concentration at all three sites. The effect of elevated CO₂ on rhizosphere microorganisms was plant dependent and increased for: Brassica napus=Triticum aestivum<Beta vulgaris<Populus deltoides. N deficiency affected microbial growth rates directly (N limitation) and indirectly (changing the quantity of fine roots). So, 50% decrease in N fertilization caused the overall increase or decrease of microbial growth rates depending on plant species. The μ‐value increase was lower for microorganisms growing on yeast extract then for those growing on glucose, i.e. the effect of elevated CO₂ was smoothed on rich vs. simple substrate. So, the r/K strategies ratio can be better revealed by studying growth on simple (glucose) than on rich substrate mixtures (yeast extract). Our results clearly showed that the functional characteristics of the soil microbial community (i.e. specific growth rates) rather than total microbial biomass amount are sensitive to increased atmospheric CO₂. We conclude that the more abundant available organics released by roots at elevated CO₂ altered the ecological strategy of the soil microbial community specifically a shift to a higher contribution of fast‐growing r‐selected species was observed. These changes in functional structure of the soil microbial community may counterbalance higher C input into the soil under elevated atmospheric CO₂ concentration.     

Host-specific aphid population responses to elevated CO2 and increased N availability

Sap-feeding insects such as aphids are the only insect herbivores that show positive responses to elevated CO₂. Recent models predict that increased nitrogen will increase aphid population size under elevated CO₂, but few experiments have tested this idea empirically. To determine whether soil nitrogen (N) availability modifies aphid responses to elevated CO₂, we tested the performance of Macrosiphum euphorbiae feeding on two host plants; a C₃ plant (Solanum dulcamara), and a C₄ plant (Amaranthus viridis). We expected aphid population size to increase on plants in elevated CO₂, with the degree of increase depending on the N availability. We found a significant CO₂× N interaction for the response of population size for M. euphorbiae feeding on S. dulcamara: aphids feeding on plants grown in ambient CO₂, low N conditions increased in response to either high N availability or elevated CO₂. No population size responses were observed for aphids infesting A. viridis. Elevated CO₂ increased plant biomass, specific leaf weight, and C : N ratios of the C₃ plant, S. dulcamara but did not affect the C₄ plant, A. viridis. Increased N fertilization significantly increased plant biomass, leaf area, and the weight : height ratio in both experiments. Elevated CO₂ decreased leaf N in S. dulcamara and had no effect on A. viridis, while higher N availability increased leaf N in A. viridis and had no effect in S. dulcamara. Aphid infestation only affected the weight : height ratio of S. dulcamara. We only observed an increase in aphid population size in response to elevated CO₂ or increased N availability for aphids feeding on S. dulcamara grown under low N conditions. There appears to be a maximum population growth rate that M. euphorbiae aphids can attain, and we suggest that this response is because of intrinsic limits on development time and fecundity.     

Effects of elevated CO2 and Pb on the microbial community in the rhizosphere of Pinus densiflora

Rising levels of atmospheric CO₂ may stimulate forest productivity in the future, resulting in increased carbon storage in terrestrial ecosystems. However, heavy metal contamination may interfere with this, though the response is not yet known. In this study, we investigated the effect of elevated CO₂ and Pb contamination on microorganisms and decomposition in pine tree forest soil. Three-year old pine trees (Pinus densiflora) were planted in Pb contaminated soils (500 mg/kg-soil) and uncontaminated soils and cultivated for three months in a growth chamber where the CO₂ concentration was controlled at 380 or 760 mg/kg. Structures of the microbial community were comparatively analyzed in bulk and in rhizosphere soil samples using community-level physiological profiling (CLPP) and 16S rRNA gene PCR-DGGE (denaturing gradient gel electrophoresis). Additionally, microbial activity in rhizospheric soil, growth and the C/N ratio of the pine trees were measured. Elevated CO₂ significantly increased microbial activities and diversity in Pb contaminated soils due to the increase in carbon sources, and this increase was more distinctive in rhizospheric soil than in bulk soils. In addition, increased plant growth and C/N ratios of pine needles at elevated CO₂ resulted in an increase in cation exchange capacity (CEC) and dissolved organic carbon (DOC) of the rhizosphere in Pb contaminated soil. Taken together, these findings indicate that elevated CO₂ levels and heavy metals can affect the soil carbon cycle by changing the microbial community and plant metabolism.     

Carbon partitioning and rhizosphere C-flow in Lolium perenne as affected by CO2 concentration, irradiance and below-ground conditions

Plant responses to increasing atmospheric CO₂ concentrations have received considerable interest. However, major uncertainties in relation to interactive effects of CO₂ with above- and below-ground conditions remain. This microcosm study investigated the impacts of CO₂ concentration on plant growth, dry matter partitioning and rhizodeposition as affected by: (i) photon flux density (PFD), and (ii) growth matrix. Plants were grown in a sandy loam soil for 28 d under two photon flux densities: 350 (low PFD) and 1000 μmol m^–2 s^–1 (high PFD) and two CO₂ concentrations: 450 (low CO₂) and 720 μmol mol^–1 (high CO₂). Partitioning of recent assimilate amongst plant and rhizosphere C-pools was determined by use of ¹⁴CO₂ pulse-labelling. In treatments with high PFD and/or high CO₂, significant (P < 0.05) increases in dry matter production were found in comparison with the low PFD/low CO₂ treatment. In addition, significant (P < 0.05) reductions in shoot %N and SLA were found in treatments imposing high PFD and/or high CO₂. Root weight ratio (RWR) was unaffected by CO₂ concentration, however, partitioning of ¹⁴C to below ground pools was significantly (P < 0.05) increased. In a separate study, L. perenne was grown for 28 d in microcosms percolated with nutrient solution, in either a sterile sand matrix or nonsterile soil, under high or low CO₂. Dry matter production was significantly (P < 0.01) increased for both sand and soil grown seedlings. Dry matter partitioning was affected by matrix type. ¹⁴C-allocation below ground was increased for sand grown plants. Rhizodeposition was affected by CO₂ concentration for growth in each matrix, but was increased for plants grown in the soil matrix, and decreased for those in sand. The results illustrate that plant responses to CO₂ are potentially affected by (i) PFD, and (ii) by feedbacks from the growth matrix. Such feedbacks are discussed in relation to soil nutrient status and interactions with the rhizosphere microbial biomass.     

Interactions between plant growth and soil nutrient cycling under elevated CO2: a meta-analysis

free air carbon dioxide enrichment (FACE) and open top chamber (OTC) studies are valuable tools for evaluating the impact of elevated atmospheric CO₂ on nutrient cycling in terrestrial ecosystems. Using meta‐analytic techniques, we summarized the results of 117 studies on plant biomass production, soil organic matter dynamics and biological N₂ fixation in FACE and OTC experiments. The objective of the analysis was to determine whether elevated CO₂ alters nutrient cycling between plants and soil and if so, what the implications are for soil carbon (C) sequestration. Elevated CO₂ stimulated gross N immobilization by 22%, whereas gross and net N mineralization rates remained unaffected. In addition, the soil C : N ratio and microbial N contents increased under elevated CO₂ by 3.8% and 5.8%, respectively. Microbial C contents and soil respiration increased by 7.1% and 17.7%, respectively. Despite the stimulation of microbial activity, soil C input still caused soil C contents to increase by 1.2% yr^?1. Namely, elevated CO₂ stimulated overall above‐ and belowground plant biomass by 21.5% and 28.3%, respectively, thereby outweighing the increase in CO₂ respiration. In addition, when comparing experiments under both low and high N availability, soil C contents (+2.2% yr^?1) and above‐ and belowground plant growth (+20.1% and+33.7%) only increased under elevated CO₂ in experiments receiving the high N treatments. Under low N availability, above‐ and belowground plant growth increased by only 8.8% and 14.6%, and soil C contents did not increase. Nitrogen fixation was stimulated by elevated CO₂ only when additional nutrients were supplied. These results suggest that the main driver of soil C sequestration is soil C input through plant growth, which is strongly controlled by nutrient availability. In unfertilized ecosystems, microbial N immobilization enhances acclimation of plant growth to elevated CO₂ in the long‐term. Therefore, increased soil C input and soil C sequestration under elevated CO₂ can only be sustained in the long‐term when additional nutrients are supplied.     

Stimulation of microbial extracellular enzyme activities by elevated CO2 depends on soil aggregate size

Increased belowground carbon (C) transfer by plant roots at elevated CO₂ may change properties of the microbial community in the rhizosphere. Previous investigations that focused on total soil organic C or total microbial C showed contrasting results: small increase, small decrease or no changes. We evaluated the effect of 5 years of elevated CO₂ (550 ppm) on four extracellular enzymes: β‐glucosidase, chitinase, phosphatase, and sulfatase. We expected microorganisms to be differently localized in aggregates of various sizes and, therefore analyzed microbial biomass (C_mic by SIR) and enzyme activities in three aggregate‐size classes: large macro‐ (> 2 mm), small macro‐ (0.25–2 mm), and microaggregates (< 0.25 mm). To estimate the potential enzyme production, we activated microorganisms by substrate (glucose and nutrients) amendment. Although C_total and C_mic as well as the activities of β‐glucosidase, phosphatase, and sulfatase were unaffected in bulk soil and in aggregate‐size classes by elevated CO₂, significant changes were observed in potential enzyme production after substrate amendment. After adding glucose, enzyme activities under elevated CO₂ were 1.2–1.9‐fold higher than under ambient CO₂. This indicates the increased activity of microorganisms, which leads to accelerated C turnover in soil under elevated CO₂. Significantly higher chitinase activity in bulk soil and in large macroaggregates under elevated CO₂ revealed an increased contribution of fungi to turnover processes. At the same time, less chitinase activity in microaggregates underlined microaggregate stability and the difficulties for fungal hyphae penetrating them. We conclude that quantitative and qualitative changes of C input by plants into the soil at elevated CO₂ affect microbial community functioning, but not its total content. Future studies should therefore focus more on the changes of functions and activities, but less on the pools.     

Increased Quantity and Quality of Coarse Soil Organic Matter Fraction at Elevated CO2 in a Grazed Grassland are a Consequence of Enhanced Root Growth Rate and Turnover

The aims of this study were to determine whether elevated atmospheric CO₂ concentration modifies plant organic matter (OM) fluxes to the soil and whether any change in the fluxes can modify soil OM accumulation. Measurements were made in a grazed temperate grassland after almost 4 years exposure to elevated atmospheric CO₂ (475 μl l^-1) using a Free Air CO₂ Enrichment (FACE) facility located in the North Island of New Zealand. Aboveground herbage biomass and leaf litter production were not altered by elevated CO₂ but root growth rate, as measured with the ingrowth core method, and root turnover were strongly stimulated by elevated CO₂ particularly at low soil moisture contents during summer. Consequently, significantly more plant material was returned to the soil under elevated CO₂ leading to an accumulation of coarse (> 1 mm) particulate organic matter (POM) but not of finer POM fractions. The accumulating POM exhibited a lower C/N ratio, which was attributed to the higher proportion of legumes in the pasture under elevated CO₂. Only small changes were detected in the size and activity of the soil microbial biomass in response to the POM accumulation, suggesting that higher organic substrate availability did not stimulate microbial growth and activity despite the apparent lower C/N ratio of accumulating POM. As a result, elevated CO₂ may well lead to an accumulation of OM in grazed grassland soil in the long term.     

Carbon and nitrogen pools and mineralization in a grassland gley soil under elevated carbon dioxide at a natural CO2 spring

The growth and chemical composition of most plants are influenced by elevated CO₂, but accompanying effects on soil organic matter pools and mineralization are less clearly defined, partly because of the short‐term nature of most studies. Herein we describe soil properties from a naturally occurring cold CO₂ spring (Hakanoa) in Northland, New Zealand, at which the surrounding vegetation has been exposed to elevated CO₂ for at least several decades. The mean annual temperature at this site is ≈ 15.5 °C and rainfall ≈ 1550 mm. The site was unfertilized and ungrazed, with a vegetation of mainly C3 and C4 grasses, and had moderate levels of ‘available’ P. Two soils were present ? a gley soil and an organic soil – but only the gley soil is examined here. Average atmospheric CO₂ concentrations at 17 sampling locations in the gley soil area ranged from 372 to 670 ppmv. In samples at 0–5 cm depth, pH averaged 5.4; average values for organic C were 150 g, total N 11 g, microbial C 3.50 g, and microbial N 0.65 g kg^?1, respectively. Under standardized moisture conditions at 25 °C, average rates of CO₂‐C production (7–14 days) were 5.4 mg kg^?1 h^?1 and of net mineral‐N production (14 ?42 days) 0.40 mg kg^?1 h^?1. These properties were all correlated positively and significantly (P < 0.10) with atmospheric CO₂ concentrations, but not with soil moisture (except for CO₂‐C production) or with clay content; they were, however, correlated negatively and mainly significantly with soil pH. In spite of uncertainties associated with the uncontrolled environment of naturally occurring springs, we conclude that storage of C and N can increase under prolonged exposure to elevated CO₂, and may include an appreciable labile fraction in mineral soil with an adequate nutrient supply.     

Four years of experimental climate change modifies the microbial drivers of N2O fluxes in an upland grassland ecosystem

Emissions of the trace gas nitrous oxide (N₂O) play an important role for the greenhouse effect and stratospheric ozone depletion, but the impacts of climate change on N₂O fluxes and the underlying microbial drivers remain unclear. The aim of this study was to determine the effects of sustained climate change on field N₂O fluxes and associated microbial enzymatic activities, microbial population abundance and community diversity in an extensively managed, upland grassland. We recorded N₂O fluxes, nitrification and denitrification, microbial population size involved in these processes and community structure of nitrite reducers (nirK) in a grassland exposed for 4 years to elevated atmospheric CO₂ (+200 ppm), elevated temperature (+3.5 °C) and reduction of summer precipitations (?20%) as part of a long‐term, multifactor climate change experiment. Our results showed that both warming and simultaneous application of warming, summer drought and elevated CO₂ had a positive effect on N₂O fluxes, nitrification, N₂O release by denitrification and the population size of N₂O reducers and NH₄ oxidizers. In situ N₂O fluxes showed a stronger correlation with microbial population size under warmed conditions compared with the control site. Specific lineages of nirK denitrifier communities responded significantly to temperature. In addition, nirK community composition showed significant changes in response to drought. Path analysis explained more than 85% of in situ N₂O fluxes variance by soil temperature, denitrification activity and specific denitrifying lineages. Overall, our study underlines that climate‐induced changes in grassland N₂O emissions reflect climate‐induced changes in microbial community structure, which in turn modify microbial processes.     

Laboratory incubations reveal potential responses of soil nitrogen cycling to changes in soil C and N availability in Mojave Desert soils exposed to elevated atmospheric CO2

Elevated atmospheric carbon dioxide (CO₂) has the potential to alter soil carbon (C) and nitrogen (N) cycling in arid ecosystems through changes in net primary productivity. However, an associated feedback exists because any sustained increases in plant productivity will depend upon the continued availability of soil N. We took soils from under the canopies of major shrubs, grasses, and plant interspaces in a Mojave Desert ecosystem exposed to elevated atmospheric CO₂ and incubated them in the laboratory with amendments of labile C and N to determine if elevated CO₂ altered the mechanistic controls of soil C and N on microbial N cycling. Net ammonification increased under shrubs exposed to elevated CO₂, while net nitrification decreased. Elevated CO₂ treatments exhibited greater fluxes of N₂O–N under Lycium spp., but not other microsites. The proportion of microbial/extractable organic N increased under shrubs exposed to elevated CO₂. Heterotrophic N₂‐fixation and C mineralization increased with C addition, while denitrification enzyme activity and N₂O–N fluxes increased when C and N were added in combination. Laboratory results demonstrated the potential for elevated CO₂ to affect soil N cycling under shrubs and supports the hypothesis that energy limited microbes may increase net inorganic N cycling rates as the amount of soil‐available C increases under elevated CO₂. The effect of CO₂ enrichment on N‐cycling processes is mediated by its effect on the plants, particularly shrubs. The potential for elevated atmospheric CO₂ to lead to accumulation of NH₄⁺ under shrubs and the subsequent volatilization of NH₃ may result in greater losses of N from this system, leading to changes in the form and amount of plant‐available inorganic N. This introduces the potential for a negative feedback mechanism that could act to constrain the degree to which plants can increase productivity in the face of elevated atmospheric CO₂.     

Elevated atmospheric CO2 and feedback between carbon and nitrogen cycles

We tested a conceptual model describing the influence of elevated atmospheric CO₂ on plant production, soil microorganisms, and the cycling of C and N in the plant-soil system. Our model is based on the observation that in nutrient-poor soils, plants (C₃) grown in an elevated CO₂ atmosphere often increase production and allocation to belowground structures. We predicted that greater belowground C inputs at elevated CO₂ should elicit an increase in soil microbial biomass and increased rates of organic matter turnover and nitrogen availability. We measured photosynthesis, biomass production, and C allocation of Populus grandidentata Michx. grown in nutrient-poor soil for one field season at ambient and twice-ambient (i.e., elevated) atmospheric CO₂ concentrations. Plants were grown in a sandy subsurface soil i) at ambient CO₂ with no open top chamber, ii) at ambient CO₂ in an open top chamber, and iii) at twice-ambient CO₂ in an open top chamber. Plants were fertilized with 4.5 g N m⁻² over a 47 d period midway through the growing season. Following 152 d of growth, we quantified microbial biomass and the availabilities of C and N in rhizosphere and bulk soil. We tested for a significant CO₂ effect on plant growth and soil C and N dynamics by comparing the means of the chambered ambient and chambered elevated CO₂ treatments. Rates of photosynthesis in plants grown at elevated CO₂ were significantly greater than those measured under ambient conditions. The number of roots, root length, and root length increment were also substantially greater at elevated CO₂. Total and belowground biomass were significantly greater at elevated CO₂. Under N-limited conditions, plants allocated 50–70% of their biomass to roots. Labile C in the rhizosphere of elevated-grown plants was significantly greater than that measured in the ambient treatments; there were no significant differences between labile C pools in the bulk soil of ambient and elevated-grown plants. Microbial biomass C was significantly greater in the rhizosphere and bulk soil of plants grown at elevated CO₂ compared to that in the ambient treatment. Moreover, a short-term laboratory assay of N mineralization indicated that N availability was significantly greater in the bulk soil of the elevated-grown plants. Our results suggest that elevated atmospheric CO₂ concentrations can have a positive feedback effect on soil C and N dynamics producing greater N availability. Experiments conducted for longer periods of time will be necessary to test the potential for negative feedback due to altered leaf litter chemistry. ei]{gnH}{fnLambers} ei]{gnA C}{fnBorstlap}     

Terrestrial higher-plant response to increasing atmospheric [CO2] in relation to the global carbon cycle

Terrestrial higher plants exchange large amounts of CO₂ with the atmosphere each year; c. 15% of the atmospheric pool of C is assimilated in terrestrial-plant photosynthesis each year, with an about equal amount returned to the atmosphere as CO₂ in plant respiration and the decomposition of soil organic matter and plant litter. Any global change in plant C metabolism can potentially affect atmospheric CO₂ content during the course of years to decades. In particular, plant responses to the presently increasing atmospheric CO₂ concentration might influence the rate of atmospheric CO₂ increase through various biotic feedbacks. Climatic changes caused by increasing atmospheric CO₂ concentration may modulate plant and ecosystem responses to CO₂ concentration. Climatic changes and increases in pollution associated with increasing atmospheric CO₂ concentration may be as significant to plant and ecosystem C balance as CO₂ concentration itself. Moreover, human activities such as deforestation and livestock grazing can have impacts on the C balance and structure of individual terrestrial ecosystems that far outweigh effects of increasing CO₂ concentration and climatic change. In short-term experiments, which in this case means on the order of 10 years or less, elevated atmospheric CO₂ concentration affects terrestrial higher plants in several ways. Elevated CO₂ can stimulate photosynthesis, but plants may acclimate and (or) adapt to a change in atmospheric CO₂ concentration. Acclimation and adaptation of photosynthesis to increasing CO₂ concentration is unlikely to be complete, however. Plant water use efficiency is positively related to CO₂ concentration, implying the potential for more plant growth per unit of precipitation or soil moisture with increasing atmospheric CO₂ concentration. Plant respiration may be inhibited by elevated CO₂ concentration, and although a naive C balance perspective would count this as a benefit to a plant, because respiration is essential for plant growth and health, an inhibition of respiration can be detrimental. The net effect on terrestrial plants of elevated atmospheric CO₂ concentration is generally an increase in growth and C accumulation in phytomass. Published estimations, and speculations about, the magnitude of global terrestrial-plant growth responses to increasing atmospheric CO₂ concentration range from negligible to fantastic. Well-reasoned analyses point to moderate global plant responses to CO₂ concentration. Transfer of C from plants to soils is likely to increase with elevated CO₂ concentrations because of greater plant growth, but quantitative effects of those increased inputs to soils on soil C pool sizes are unknown. Whether increases in leaf-level photosynthesis and short-term plant growth stimulations caused by elevated atmospheric CO₂ concentration will have, by themselves, significant long-term (tens to hundreds of years) effects on ecosystem C storage and atmospheric CO₂ concentration is a matter for speculation, not firm conclusion. Long-term field studies of plant responses to elevated atmospheric CO₂ are needed. These will be expensive, difficult, and by definition, results will not be forthcoming for at least decades. Analyses of plants and ecosystems surrounding natural geological CO₂ degassing vents may provide the best surrogates for long-term controlled experiments, and therefore the most relevant information pertaining to long-term terrestrial-plant responses to elevated CO₂ concentration, but pollutants associated with the vents are a concern in some cases, and quantitative knowledge of the history of atmospheric CO₂ concentrations near vents is limited. On the whole, terrestrial higher-plant responses to increasing atmospheric CO₂ concentration probably act as negative feedbacks on atmospheric CO₂ concentration increases, but they cannot by themselves stop the fossil-fuel-oxidation-driven increase in atmospheric CO₂ concentration. And, in the very long-term, atmospheric CO₂ concentration is controlled by atmosphere-ocean C equilibrium rather than by terrestrial plant and ecosystem responses to atmospheric CO₂ concentration.     

Plant species, atmospheric CO2 and soil N interactively or additively control C allocation within plant-soil systems

Two plant species, Medicago truncatula (legume) and Avena sativa (non-legume), were grown in low-or high-N soils under two CO₂ concentrations to test the hypothesis whether C allocation within plant-soil system is interactively or additively controlled by soil N and atmospheric CO₂ is dependent upon plant species. The results showed the interaction between plant species and soil N had a significant impact on microbial activity and plant growth. The interaction between CO₂ and soil N had a significant impact on soil soluble C and soil microbial biomass C under Madicago but not under Avena. Although both CO₂ and soil N affected plant growth significantly, there was no interaction between CO₂ and soil N on plant growth. In other words, the effects of CO₂ and soil N on plant growth were additive. We considered that the interaction between N₂ fixation trait of legume plant and elevated CO₂ might have obscured the interaction between soil N and elevated CO₂ on the growth of legume plant. In low-N soil, the shoot-to-root ratio of Avena dropped from 2.63±0.20 in the early growth stage to 1.47±0.03 in the late growth stage, indicating that Avena plant allocated more energy to roots to optimize nutrient uptake (i.e. N) when soil N was limiting. In high-N soil, the shoot-to-root ratio of Medicago increased significantly over time (from 2.45±0.30 to 5.43±0.10), suggesting that Medicago plants allocated more energy to shoots to optimize photosynthesis when N was not limiting. The shoot-to-root ratios were not significantly different between two CO₂ levels.