Biomass production in a nitrogen-fertilized,tallgrass prairie ecosystem exposed to ambient and elevated levels of CO2

Increased biomass production in terrestrial ecosystems with elevated atmospheric CO₂ may be constrained by nutrient limitations as a result of increased requirement or reduced availability caused by reduced turnover rates of nutrients. To determine the short-term impact of nitrogen (N) fertilization on plant biomass production under elevated CO₂, we compared the response of N-fertilized tallgrass prairie at ambient and twice-ambient CO₂ levels over a 2-year period. Native tallgrass prairie plots (4.5 m diameter) were exposed continuously (24 h) to ambient and twice-ambient CO₂ from 1 April to 26 October. We compared our results to an unfertilized companion experiment on the same research site. Above- and belowground biomass production and leaf area of fertilized plots were greater with elevated than ambient CO₂ in both years. The increase in biomass at high CO₂ occurred mainly aboveground in 1991, a dry year, and belowground in 1990, a wet year. Nitrogen concentration was lower in plants exposed to elevated CO₂, but total standing crop N was greater at high CO₂. Increased root biomass under elevated CO₂ apparently increased N uptake. The biomass production response to elevated CO₂ was much greater on N-fertilized than unfertilized prairie, particularly in the dry year. We conclude that biomass production response to elevated CO₂ was suppressed by N limitation in years with below-normal precipitation. Reduced N concentration in above- and belowground biomass could slow microbial degradation of soil organic matter and surface litter, thereby exacerbating N limitation in the long term.     

Above- and belowground response of Populus grandidentata to elevated atmospheric CO2 and soil N availability

Soil N availability may play an important role in regulating the long-term responses of plants to rising atmospheric CO₂ partial pressure. To further examine the linkage between above- and belowground C and N cycles at elevated CO₂, we grew clonally propagated cuttings of Populus grandidentata in the field at ambient and twice ambient CO₂ in open bottom root boxes filled with organic matter poor native soil. Nitrogen was added to all root boxes at a rate equivalent to net N mineralization in local dry oak forests. Nitrogen added during August was enriched with ¹⁵N to trace the flux of N within the plant-soil system. Above-and belowground growth, CO₂ assimilation, and leaf N content were measured non-destructively over 142 d. After final destructive harvest, roots, stems, and leaves were analyzed for total N and ¹⁵N. There was no CO₂ treatment effect on leaf area, root length, or net assimilation prior to the completion of N addition. Following the N addition, leaf N content increased in both CO₂ treatments, but net assimilation showed a sustained increase only in elevated CO₂ grown plants. Root relative extension rate was greater at elevated CO₂, both before and after the N addition. Although final root biomass was greater at elevated CO₂, there was no CO₂ effect on plant N uptake or allocation. While low soil N availability severely inhibited CO₂ responses, high CO₂ grown plants were more responsive to N. This differential behavior must be considered in light of the temporal and spatial heterogeneity of soil resources, particularly N which often limits plant growth in temperate forests.     

Plant Nitrogen Dynamics in Shortgrass Steppe under Elevated Atmospheric Carbon Dioxide

The direct and indirect effects of increasing levels of atmospheric carbon dioxide (CO₂) on plant nitrogen (N) content were studied in a shortgrass steppe ecosystem in northeastern Colorado, USA. Beginning in 1997 nine experimental plots were established: three open-top chambers with ambient CO₂ levels (approximately 365 mol mol^–1), three open-top chambers with twice-ambient CO₂ levels (approximately 720 mol mol^–1), and three unchambered control plots. After 3 years of growing-season CO₂ treatment, the aboveground N concentration of plants grown under elevated atmospheric CO₂ decreased, and the carbon–nitrogen (C:N) ratio increased. At the same time, increased aboveground biomass production under elevated atmospheric CO₂ conditions increased the net transfer of N out of the soil of elevated-CO₂ plots. Aboveground biomass production after simulated herbivory was also greater under elevated CO₂ compared to ambient CO₂. Surprisingly, no significant changes in belowground plant tissue N content were detected in response to elevated CO₂. Measurements of individual species at peak standing phytomass showed significant effects of CO₂ treatment on aboveground plant tissue N concentration and significant differences between species in N concentration, suggesting that changes in species composition under elevated CO₂ will contribute to overall changes in nutrient cycling. Changes in plant N content, driven by changes in aboveground plant N concentration, could have important consequences for biogeochemical cycling rates and the long-term productivity of the shortgrass steppe as atmospheric CO₂ concentrations increase.     

Effects of elevated atmospheric CO2 on protozoan abundance in soil planted with wheat and on decomposition of wheat roots

The effect of elevated CO₂ on growth of wheat plants (Triticum aestivum cv. Minaret) and soil protozoan and bacterial populations was investigated in soil pots placed in open top chambers fumigated with ambient air or air enriched with CO₂ (ambient + 320 l L^–1 CO₂). We harvested plants two times during the growing season and measured the biomass and the C and N content of roots and shoots. The soil was divided into bulk and rhizosphere soil and the number of bacteria (colony-forming units, CFU) and protozoa was determined. There was no effect of atmospheric CO₂ content on the number of bacteria, but the total number of bacterivorous protozoa was higher in pots from the elevated CO₂ treatment. This increase was mainly due to an increase in the number of protozoa in the bulk soil. Density of protozoa in the rhizosphere was not affected by elevated CO₂. This suggests that the increase in protozoan numbers was a result of a general increase in rhizodeposition, presumably caused by increased root production, and not to an increased root exudation per root mass. After harvest, soil from the two treatments was incubated with and without roots and the respiration rate was estimated at intervals for 200 days. During the first 55 days, the specific root induced respiration rate was not affected by the CO₂ level at which the plants had been grown, indicating that the quality of the easily decomposable components of the roots was not affected by CO₂ level.     

Response of soil biota to elevated atmospheric CO2 in poplar model systems

We tested the hypotheses that increased belowground allocation of carbon by hybrid poplar saplings grown under elevated atmospheric CO₂ would increase mass or turnover of soil biota in bulk but not in rhizosphere soil. Hybrid poplar saplings (Populus×euramericana cv. Eugenei) were grown for 5 months in open-bottom root boxes at the University of Michigan Biological Station in northern, lower Michigan. The experimental design was a randomized-block design with factorial combinations of high or low soil N and ambient (34 Pa) or elevated (69 Pa) CO₂ in five blocks. Rhizosphere microbial biomass carbon was 1.7 times greater in high-than in low-N soil, and did not respond to elevated CO₂. The density of protozoa did not respond to soil N but increased marginally (P < 0.06) under elevated CO₂. Only in high-N soil did arbuscular mycorrhizal fungi and microarthropods respond to CO₂. In high-N soil, arbuscular mycorrhizal root mass was twice as great, and extramatrical hyphae were 11% longer in elevated than in ambient CO₂ treatments. Microarthropod density and activity were determined in situ using minirhizotrons. Microarthropod density did not change in response to elevated CO₂, but in high-N soil, microarthropods were more strongly associated with fine roots under elevated than ambient treatments. Overall, in contrast to the hypotheses, the strongest response to elevated atmospheric CO₂ was in the rhizosphere where (1) unchanged microbial biomass and greater numbers of protozoa (P < 0.06) suggested faster bacterial turnover, (2) arbuscular mycorrhizal root length increased, and (3) the number of microarthropods observed on fine roots rose. Received: 18 March 1997 / Accepted: 5 August 1997     

Carbon dynamics and microbial activity in tallgrass prairie exposed to elevated CO2 for 8 years

Alterations in microbial mineralization and nutrient cycling may control the long-term response of ecosystems to elevated CO₂. Because micro-organisms constitute a labile fraction of potentially available N and are regulators of decomposition, an understanding of microbial activity and microbial biomass is crucial. Tallgrass prairie was exposed to twice ambient CO₂ for 8 years beginning in 1989. Starting in 1991 and ending in 1996, soil samples from 0 to 5 and 5 to 15 cm depths were taken for measurement of microbial biomass C and N, total C and N, microbial activity, inorganic N and soil water content. Because of increased water-use-efficiency by plants, soil water content was consistently and significantly greater in elevated CO₂ compared to ambient treatments. Soil microbial biomass C and N tended to be greater under elevated CO₂ than ambient CO₂ in the 5–15 cm depth during most years, and in the month of October, when analyzed over the entire study period. Microbial activity was significantly greater at both depths in elevated CO₂ than ambient conditions for most years. During dry periods, the greater water content of the surface 5 cm soil in the elevated CO₂ treatments increased microbial activity relative to the ambient CO₂ conditions. The increase in microbial activity under elevated CO₂ in the 5–15 cm layer was not correlated with differences in soil water contents, but may have been related to increases in soil C inputs from enhanced root growth and possibly greater root exudation. Total soil C and N in the surface 15 cm were, after 8 years, significantly greater under elevated CO₂ than ambient CO₂. Our results suggest that decomposition is enhanced under elevated CO₂ compared with ambient CO₂, but that inputs of C are greater than the decomposition rates. Soil C sequestration in tallgrass prairie and other drought-prone grassland systems is, therefore, considered plausible as atmospheric CO₂ increases. This revised version was published online in June 2006 with corrections to the Cover Date.     

Combined effects of atmospheric CO<Subscript>2</Subscript> and N availability on the belowground carbon and nitrogen dynamics of aspen mesocosms

It is uncertain whether elevated atmospheric CO₂ will increase C storage in terrestrial ecosystems without concomitant increases in plant access to N. Elevated CO₂ may alter microbial activities that regulate soil N availability by changing the amount or composition of organic substrates produced by roots. Our objective was to determine the potential for elevated CO₂ to change N availability in an experimental plant-soil system by affecting the acquisition of root-derived C by soil microbes. We grew Populus tremuloides (trembling aspen) cuttings for 2 years under two levels of atmospheric CO₂ (36.7 and 71.5 Pa) and at two levels of soil N (210 and 970 μg N g^–1). Ambient and twice-ambient CO₂ concentrations were applied using open-top chambers, and soil N availability was manipulated by mixing soils differing in organic N content. From June to October of the second growing season, we measured midday rates of soil respiration. In August, we pulse-labeled plants with ¹⁴CO₂ and measured soil ¹⁴CO₂ respiration and the ¹⁴C contents of plants, soils, and microorganisms after a 6-day chase period. In conjunction with the August radio-labeling and again in October, we used ¹⁵N pool dilution techniques to measure in situ rates of gross N mineralization, N immobilization by microbes, and plant N uptake. At both levels of soil N availability, elevated CO₂ significantly increased whole-plant and root biomass, and marginally increased whole-plant N capital. Significant increases in soil respiration were closely linked to increases in root biomass under elevated CO₂. CO₂ enrichment had no significant effect on the allometric distribution of biomass or ¹⁴C among plant components, total ¹⁴C allocation belowground, or cumulative (6-day) ¹⁴CO₂ soil respiration. Elevated CO₂ significantly increased microbial ¹⁴C contents, indicating greater availability of microbial substrates derived from roots. The near doubling of microbial ¹⁴C contents at elevated CO₂ was a relatively small quantitative change in the belowground C cycle of our experimental system, but represents an ecologically significant effect on the dynamics of microbial growth. Rates of plant N uptake during both 6-day periods in August and October were significantly greater at elevated CO₂, and were closely related to fine-root biomass. Gross N mineralization was not affected by elevated CO₂. Despite significantly greater rates of N immobilization under elevated CO₂, standing pools of microbial N were not affected by elevated CO₂, suggesting that N was cycling through microbes more rapidly. Our results contained elements of both positive and negative feedback hypotheses, and may be most relevant to young, aggrading ecosystems, where soil resources are not yet fully exploited by plant roots. If the turnover of microbial N increases, higher rates of N immobilization may not decrease N availability to plants under elevated CO₂. Received: 12 February 1999 / Accepted: 2 March 2000     

Growth and nitrogen accretion of dinitrogen-fixing Alnus glutinosa (L.) Gaertn. under elevated carbon dioxide

Short-term studies of tree growth at elevated CO₂ suggest that forest productivity may increase as atmospheric CO₂ concentrations rise, although low soil N availability may limit the magnitude of this response. There have been few studies of growth and N₂ fixation by symbiotic N₂-fixing woody species under elevated CO₂ and the N inputs these plants could provide to forest ecosystems in the future. We investigated the effect of twice ambient CO₂ on growth, tissue N accretion, and N₂ fixation of nodulated Alnus glutinosa (L.) Gaertn. grown under low soil N conditions for 160 d. Root, nodule, stem, and leaf dry weight (DW) and N accretion increased significantly in response to elevated CO₂. Whole-plant biomass and N accretion increased 54% and 40%, respectively. Delta-¹⁵N analysis of leaf tissue indicated that plants from both treatments derived similar proportions of their total N from symbiotic fixation suggesting that elevated CO₂ grown plants fixed approximately 40% more N than did ambient CO₂ grown plants. Leaves from both CO₂ treatments showed similar relative declines in leaf N content prior to autumnal leaf abscission, but total N in leaf litter increased 24% in elevated compared to ambient CO₂ grown plants. These results suggest that with rising atmospheric CO₂ N₂-fixing woody species will accumulate greater amounts of biomass N through N₂ fixation and may enhance soil N levels by increased litter N inputs.     

Nitrogen and phosphorus dynamics of a tallgrass prairie ecosystem exposed to elevated carbon dioxide

A tallgrass prairie ecosystem was exposed to ambient and twice-ambient CO₂ concentrations in open-top chambers and compared to unchambered ambient CO₂ during the entire growing season from 1989 through 1991. Dominant species were Andropogon gerardii (C₄), A. scoparius (C₄), Sorghastrum nutans (C₄) and Poa pratensis (C₃). Nitrogen and phosphorus concentrations in A. gerardii, P. pratensis and dicotyledonous herbs above ground biomass were estimated by periodic sampling throughout the growing season in 1989 and 1990. In 1991, N and P concentrations in peak biomass were estimated by an early August harvest. N and P concentrations in root production as a function of treatment were estimated using root ingrowth bags that remained in place throughout the growing season. Total N and P in above- and belowground biomass were calculated as products of concentration and peak biomass by species groups. N concentration in A. gerardii and dicotyledonous herb aboveground biomass was lower and total N higher in elevated CO₂ plots than in ambient CO₂ plots. N concentration in P. pratensis aboveground biomass was lower in elevated CO₂ plots than in ambient, but total N did not differ among treatments in 2 out of 3 years. In 1990, N concentration in root ingrowth bag biomass was lower and total N greater in elevated CO₂ than in ambient CO₂ plots. Root ingrowth bag biomass N concentration did not differ among treatments in 1991, but total N was greater in elevated CO₂ plots than in ambient CO₂ plots. P concentration was lower under elevated CO₂ compared to ambient in 1989, but did not differ substantially among treatments in 1990 or 1991. In all years, total P in aboveground A. gerardii and root ingrowth bag biomass was greater under elevated CO₂ than ambient. P concentration and total P in P. pratensis was similar among treatments.     

Soil microbial response in tallgrass prairie to elevated CO2

Terrestrial responses to increasing atmospheric CO₂ are important to the global carbon budget. Increased plant production under elevated CO₂ is expected to increase soil C which may induce N limitations. The objectives of this study were to determine the effects of increased CO₂ on 1) the amount of carbon and nitrogen stored in soil organic matter and microbial biomass and 2) soil microbial activity. A tallgrass prairie ecosystem was exposed to ambient and twice-ambient CO₂ concentrations in open-top chambers in the field from 1989 to 1992 and compared to unchambered ambient CO₂ during the entire growing season. During 1990 and 1991, N fertilizer was included as a treatment. The soil microbial response to CO₂ was measured during 1991 and 1992. Soil organic C and N were not significantly affected by enriched atmospheric CO₂. The response of microbial biomass to CO₂ enrichment was dependent upon soil water conditions. In 1991, a dry year, CO₂ enrichment significantly increased microbial biomass C and N. In 1992, a wet year, microbial biomass C and N were unaffected by the CO₂ treatments. Added N increased microbial C and N under CO₂ enrichment. Microbial activity was consistently greater under CO₂ enrichment because of better soil water conditions. Added N stimulated microbial activity under CO₂ enrichment. Increased microbial N with CO₂ enrichment may indicate plant production could be limited by N availability. The soil system also could compensate for the limited N by increasing the labile pool to support increased plant production with elevated atmospheric CO₂. Longer-term studies are needed to determine how tallgrass prairie will respond to increased C input.     

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.     

Biomass production and species composition change in a tallgrass prairie ecosystem after long-term exposure to elevated atmospheric CO2

To determine the long-term impact of elevated CO₂ on primary production of native tallgrass prairie, we compared the responses of tallgrass prairie at ambient and twice-ambient atmospheric CO₂ levels over an 8-year period. Plots in open-top chambers (4.5 m diameter) were exposed continuously (24 h) to ambient and elevated CO₂ from early April to late October each year. Unchambered plots were monitored also. Above-ground peak biomass was determined by clipping each year in early August, and root growth was estimated by harvesting roots from root ingrowth bags. Plant community composition was censused each year in early June. In the last 2 years of the study, subplots were clipped on 1 June or 1 July, and regrowth was harvested on 1 October. Volumetric soil water content of the 0–100 cm soil layer was determined using neutron scattering, and was generally higher in elevated CO₂ plots than ambient. Peak above-ground biomass was greater on elevated CO₂ plots than ambient CO₂ plots with or without chambers during years with significant plant water stress. Above-ground regrowth biomass was greater under elevated CO₂ than under ambient CO₂ in a year with late-season water stress, but did not differ in a wetter year. Root ingrowth biomass was also greater in elevated CO₂ plots than ambient CO₂ plots when water stress occurred during the growing season. The basal cover and relative amount of warm-season perennial grasses (C4) in the stand changed little during the 8-year period, but basal cover and relative amount of cool-season perennial grasses (C3) in the stand declined in the elevated CO₂ plots and in ambient CO₂ plots with chambers. Forbs (C3) and members of the Cyperaceae (C3) increased in basal cover and relative amount in the stand at elevated compared to ambient CO₂. Greater biomass production under elevated CO₂ in C4-dominated grasslands may lead to a greater carbon sequestration by those ecosystems and reduce peak atmospheric CO₂ concentrations in the future.     

Fungal Inoculation and Elevated CO2 Mediate Growth of Lolium Mutiforum and Phytolacca Americana,Metal Uptake,and Metal Bioavailability in Metal-Contaminated Soil: Evidence from DGT Measurement

Fungal inoculation and elevated CO₂ may mediate plant growth and uptake of heavy metals, but little evidence from Diffusive Gradients in Thin-films (DGT) measurement has been obtained to characterize the process. Lolium mutiforum and Phytolacca americana were grown at ambient and elevated CO₂ on naturally Cd and Pb contaminated soils inoculated with and without Trichoderma asperellum strain C3 or Penicillium chrysogenum strain D4, to investigate plant growth, metal uptake, and metal bioavailability responses. Fungal inoculation increased plant biomass and shoot/root Cd and Pb concentrations. Elevated CO₂ significantly increased plants biomass, but decreased Cd and Pb concentrations in shoot/root to various extents, leading to a metal dilution phenomenon. Total Cd and Pb uptake by plants, and DGT-measured Cd and Pb concentrations in rhizosphere soils, were higher in all fungal inoculation and elevated CO₂ treatments than control treatments, with the combined treatments having more influence than either treatment alone. Metal dilution phenomenon occurred because the increase in DGT-measured bioavailable metal pools in plant rhizosphere due to elevated CO₂ was unable to match the increase in requirement for plant uptake of metals due to plant biomass increase.     

Plant productivity is a key driver of soil respiration response to climate change in a nutrient-limited soil.

Despite knowledge of the interaction between climate change factors significant uncertainty exists concerning the individual and interactive effects of elevated carbon dioxide (eCO₂) and elevated temperature (eT) on the soil microbiome and function. Here we examine the individual and interactive effects of eCO₂ and eT on tree growth, soil respiration (R_soil), biomass, structural and functional composition of microbial community, nitrogen (N) mineralisation and N availability in a whole tree chamber experiment. Eucalyptus globulus plants were grown from seedling to ca. 10 m tall for 15 months in a nutrient-poor sandy soil under ambient and elevated (+ 240 ppm) atmospheric CO₂ concentrations combined with ambient or elevated temperatures (+ 3 °C) in a full factorial design. Plant growth was strongly stimulated under eCO₂, but eT had little impact on any measured plant property. In contrast, R_soil was not consistently affected by eCO₂ or eT, but correlated strongly with root and leaf biomass. The response of N-mineralisation and nutrient availability to eCO₂ and eT varied across time, and available N correlated strongly with plant height. Further, the C:N ratio of the microbial biomass and leaves were both higher under eCeT treatment. However, these functional measures were not significantly linked to either structural or functional diversity of the soil microbiome. Taken together, these results suggest that in this low-nutrient soil, belowground processes are principally driven by aboveground productivity. Our work provides novel insight into mechanisms underlying above- and belowground response to climate change, and the potential to sequester C in a low-nutrient status soil under future climatic conditions may be limited .     

The effects of CO2 and nutrient enrichment on photosynthesis and growth of Poa annua in two consecutive generations

We studied short- and long-term growth responses of Poa annua L. (Gramineae) at ambient and elevated (ambient +200???mol?mol^?1) atmospheric CO₂. In experiment 1 we compared plant growth during the early, vegetative and final, reproductive growth phases. Plant growth in elevated CO₂ was significantly enhanced during the early phase, but this was reversed in the reproductive phase. Seed mass and percentage germination were significantly reduced in elevated CO₂. Experiment 2 tested for the impact of transgenerational and nutrient effects on the response of Poa annua to elevated CO₂. Plants were grown at ambient and elevated CO₂ for one or two consecutive generations at three soil nutrient levels. Leaf photosynthesis was significantly higher at elevated CO₂, but was also affected by both soil nutrient status and plant generation. Plants grown at elevated CO₂ and under conditions of low nutrient availability showed photosynthetic acclimation after 12?weeks of growth but not after 6?weeks. First-generation growth remained unaffected by elevated CO₂, while second-generation plants produced significantly more tillers and flowers when grown in elevated CO₂ compared to ambient conditions. This effect was strongest at low nutrient availability. Average above- and belowground biomass after 12?weeks of growth was enhanced in elevated CO₂ during both generations, but more so during plant generation 2. This study demonstrates the importance of temporal/maternal effects in plant responses to elevated CO₂.     

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.     

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.     

Atmospheric carbon dioxide and fertilizer nitrogen effects on radiation interception by rice 1

In order to predict the potential impacts of global change, it is important to understand the impact of increasing global atmospheric [CO₂] on the growth and yield of crop plants. The objectives of this study were to determine the interaction of N fertilization rates and atmospheric [CO₂] on radiation interception and radiation-use efficiency of rice (Oryza sativa L. cv. IR72) grown under tropical field conditions. Rice plants were grown inside open top chambers in a lowland rice field at the International Rice Research Institute in the Philippines at ambient (about 350 μmol mol^-1) or elevated (about 600 μmol mol^-1 during the 1993 wet season and 700 μmol mol^-1 during the 1994 dry season) in combination with three levels of applied N (0, 50 or 100 kg N ha^-1 in the wet season; 0, 90 or 200 kg N ha-1 in the dry season). Light interception was not directly affected by [CO₂], but elevated [CO₂] indirectly increased light interception through increasing total absorbed N. Plant N requirement for radiation interception was similar for rice grown under ambient [CO₂] or elevated [CO₂] treatments. The conversion efficiency of intercepted radiation to dry matter, radiation-use efficiency (RUE), was about 35% greater at elevated [CO₂] than at ambient [CO₂]. The relationship between leaf N and RUE was curvilinear. At ambient [CO₂], RUE was fairly stable across levels of leaf N, but leaf N less than about 2.5% resulted in lower RUE for plants grown with elevated [CO₂] than for plant grown at ambient [CO₂]. Decreased leaf N with increased [CO₂], therefore decreased RUE of rice plants grown at elevated [CO₂]. When predicting responses of rice to elevated [CO₂], RUE should be adjusted with a decrease in leaf N. This revised version was published online in June 2006 with corrections to the Cover Date.     

Populus tremuloides photosynthesis and crown architecture in response to elevated CO2 and soil N availability

We tested the hypothesis that elevated CO₂ would stimulate proportionally higher photosynthesis in the lower crown of Populus trees due to less N retranslocation, compared to tree crowns in ambient CO₂. Such a response could increase belowground C allocation, particularly in trees with an indeterminate growth pattern such as Populus tremuloides. Rooted cuttings of P. tremuloides were grown in ambient and twice ambient (elevated) CO₂ and in low and high soil N availability (89 ± 7 and 333 ± 16 ng N g⁻¹ day⁻¹ net mineralization, respectively) for 95 days using open-top chambers and open-bottom root boxes. Elevated CO₂ resulted in significantly higher maximum leaf photosynthesis (A _max) at both soil N levels. A _max was higher at high N than at low N soil in elevated, but not ambient CO₂. Photosynthetic N use efficiency was higher at elevated than ambient CO₂ in both soil types. Elevated CO₂ resulted in proportionally higher whole leaf A in the lower three-quarters to one-half of the crown for both soil types. At elevated CO₂ and high N availability, lower crown leaves had significantly lower ratios of carboxylation capacity to electron transport capacity (V _cmax/J _max) than at ambient CO₂ and/or low N availability. From the top to the bottom of the tree crowns, V _cmax/J _max increased in ambient CO₂, but it decreased in elevated CO₂ indicating a greater relative investment of N into light harvesting for the lower crown. Only the mid-crown leaves at both N levels exhibited photosynthetic down regulation to elevated CO₂. Stem biomass segments (consisting of three nodes and internodes) were compared to the total A _leaf for each segment. This analysis indicated that increased A _leaf at elevated CO₂ did not result in a proportional increase in local stem segment mass, suggesting that C allocation to sinks other than the local stem segment increased disproportionally. Since C allocated to roots in young Populus trees is primarily assimilated by leaves in the lower crown, the results of this study suggest a mechanism by which C allocation to roots in young trees may increase in elevated CO₂. Received: 12 August 1996 / Accepted: 12 November 1996     

Leaf gas exchange and nitrogen dynamics of N2-fixing, field-grown Alnus glutinosa under elevated atmospheric CO2

Few studies have investigated the effects of elevated CO₂ on the physiology of symbiotic N₂-fixing trees. Tree species grown in low N soils at elevated CO₂ generally show a decline in photosynthetic capacity over time relative to ambient CO₂ controls. This negative adjustment may be due to a reallocation of leaf N away from the photosynthetic apparatus, allowing for more efficient use of limiting N. We investigated the effect of twice ambient CO₂ on net CO₂ assimilation (A), photosynthetic capacity, leaf dark respiration, and leaf N content of N2-fixing Alnus glutinosa (black alder) grown in field open top chambers in a low N soil for 160 d. At growth CO₂, A was always greater in elevated compared to ambient CO₂ plants. Late season A vs. internal leaf p(CO₂) response curves indicated no negative adjustment of photosynthesis in elevated CO₂ plants. Rather, elevated CO₂ plants had 16% greater maximum rate of CO₂ fixation by Rubisco. Leaf dark respiration was greater at elevated CO₂ on an area basis, but unaffected by CO₂ on a mass or N basis. In elevated CO₂ plants, leaf N content (μg N cm^?2) increased 50% between Julian Date 208 and 264. Leaf N content showed little seasonal change in ambient CO₂ plants. A single point acetylene reduction assay of detached, nodulated root segments indicated a 46% increase in specific nitrogenase activity in elevated compared to ambient CO₂ plants. Our results suggest that N₂-fixing trees will be able to maintain high A with minimal negative adjustment of photosynthetic capacity following prolonged exposure to elevated CO₂ on N-poor soils.