Canopy profiles of photosynthetic parameters under elevated CO2 and N fertilization in a poplar plantation

A poplar plantation has been exposed to an elevated CO2 concentration for 5 years using the free air CO2 enrichment (FACE) technique. Even after such a long period of exposure, leaves of Populus x euramericana have not shown clear signs of photosynthetic acclimation. Only at the end of the growing season for shade leaves was a decrease of maximum velocity of carboxylation (Vcmax) observed. Maximum electron transport rate (Jmax) was increased by FACE treatment in July. Assimilation rates at CO2 partial pressure of 400 (A400) and 600 (A600) micromol mol(-1) were not significantly different under FACE treatment. Most notably FACE significantly decreased stomatal conductance (g(s)) both on upper and lower canopy leaves. N fertilization increased N content in the leaves on mass basis (Nm) and specific leaf area (SLA) in both CO2 treatments but did not influence the photosynthetic parameters. These data show that in poplar plantations the long-term effects of elevated CO2 on photosynthesis do not differ considerably from the short-term ones even with N deposition.     

Photosynthetic responses to elevated CO(2) and O(3) in Quercus ilex leaves at a natural CO(2) spring

Photosynthetic stimulation and stomatal conductance (Gs) depression in Quercus ilex leaves at a CO(2) spring suggested no down-regulation. The insensitivity of Gs to a CO(2) increase (from ambient 1500 to 2000 micromol mol(-1)) suggested stomatal acclimation. Both responses are likely adaptations to the special environment of CO(2) springs. At the CO(2)-enriched site, not at the control site, photosynthesis decreased 9% in leaves exposed to 2x ambient O(3) concentrations in branch enclosures, compared to controls in charcoal-filtered air. The stomatal density reduction at high CO(2) was one-third lower than the concomitant Gs reduction, so that the O(3) uptake per single stoma was lower than at ambient CO(2). No significant variation in monoterpene emission was measured. Higher trichome and mesophyll density were recorded at the CO(2)-enriched site, accounting for lower O(3) sensitivity. A long-term exposure to H(2)S, reflected by higher foliar S-content, and CO(2) might depress the antioxidant capacity of leaves close to the vent and increase their O(3) sensitivity.     

Consequences of elevated carbon dioxide and ozone for foliar chemical composition and dynamics in trembling aspen (Populus tremuloides) and paper birch (Betula papyrifera).

Atmospheric chemical composition affects foliar chemical composition, which in turn influences the dynamics of both herbivory and decomposition in ecosystems. We assessed the independent and interactive effects of CO2 and O3 fumigation on foliar chemistry of quaking aspen (Populus tremuloides) and paper birch (Betula papyrifera) at a Free-Air CO2 Enrichment (FACE) facility in northern Wisconsin. Leaf samples were collected at five time periods during a single growing season, and analyzed for nitrogen. starch and condensed tannin concentrations, nitrogen resorption efficiencies (NREs), and C:N ratios. Enriched CO2 reduced foliar nitrogen concentrations in aspen and birch; O3 only marginally reduced nitrogen concentrations. NREs were unaffected by pollution treatment in aspen, declined with 03 exposure in birch, and this decline was ameliorated by enriched CO2. C:N ratios of abscised leaves increased in response to enriched CO2 in both tree species. O3 did not significantly alter C:N ratios in aspen, although values tended to be higher in + CO2 + O3 leaves. For birch, O3 decreased C:N ratios under ambient CO2 and increased C:N ratios under elevated CO2. Thus, under the combined pollutants, the C:N ratios of both aspen and birch leaves were elevated above the averaged responses to the individual and independent trace gas treatments. Starch concentrations were largely unresponsive to CO2 and O3 treatments in aspen. but increased in response to elevated CO2 in birch. Levels of condensed tannins were negligibly affected by CO2 and O3 treatments in aspen, but increased in response to enriched CO2 in birch. Results from this work suggest that changes in foliar chemical composition elicited by enriched CO2 are likely to impact herbivory and decomposition, whereas the effects of O3 are likely to be minor, except in cases where they influence plant response to CO2.     

Effects of elevated CO2 nitrogen supply and tropospheric ozone on spring wheat. I. Growth and yield

Spring wheat (Triticum aestivum L. cv. Minaret) was exposed to three CO(2) levels, in combination with two nitrogen fertilizer levels and two levels of tropospheric ozone, from sowing to ripening in open-top chambers. Three additional nitrogen fertilizer treatments were carried out at the lowest and the highest CO(2) level, respectively. Plants were harvested at growth stages 31, 65 and 93 and separated into up to eight fractions to gain information about biomass partitioning. CO(2) enrichment (263 microl litre(-1) above ambient levels) drastically increased biomass of organs serving as long-term carbohydrate pools. Peduncle weight increased by 92%, stem weight by 73% and flag leaf sheath weight by 59% at growth stage 65. Average increase in shoot biomass due to CO(2) enrichment amounted to 51% at growth stage 65 and 36% at final harvest. Average yield increase was 34%. Elevated nitrogen application was most effective on biomass of green tissues. Yield was increased by 30% when nitrogen application was increased from 150 to 270 kg N ha(-1). Significant interactions were observed between CO(2) enrichment and nitrogen application. Yield increase due to CO(2) ranged from 23% at 120 kg N to 47% at 330 kg N. Triticum aestivum cv. Minaret was not very responsive to ozone at 1.5 times ambient levels. 1000 grain weight was slightly decreased, which was compensated by an increased number of grains.     

Regional patterns in foliar (15)N across a gradient of nitrogen deposition in the northeastern US

Recent studies have demonstrated that natural abundance (15)N can be a useful tool for assessing nitrogen saturation, because as nitrification and nitrate loss increase, delta(15)N of foliage and soil also increases. We measured foliar delta(15)N at 11 high-elevation spruce-fir stands along an N deposition gradient in 1987-1988 and at seven paired northern hardwood and spruce-fir stands in 1999. In 1999, foliar delta(15)N increased from -5.2 to -0.7 per thousand with increasing N deposition from Maine to NY. Foliar delta(15)N decreased between 1987-1988 and 1999, while foliar %N increased and foliar C:N decreased at most sites. Foliar delta(15)N was strongly correlated with N deposition, and was also positively correlated with net nitrification potential and negatively correlated with soil C:N ratio. Although the increase in foliar %N is consistent with a progression towards N saturation, other results of this study suggest that, in 1999, these stands were further from N saturation than in 1987-1988.     

Increased leaf area expansion of hybrid poplar in elevated CO2. From controlled environments to open-top chambers and to FACE.

We examined the response of hybrid poplar to elevated CO2 in contrasting growth environments: controlled environment chamber (CE). open-top chamber (OTC) and poplar free air CO2 enrichment (POPFACE) in order to compare short versus long-term effects and to determine whether generalisations in response are possible for this fast growing tree. Leaf growth, which for poplar is an important determinant of stemwood productivity was followed in all environments, as were the determinants of leaf growth-cell expansion and cell production. Elevated CO2 (550-700 micromol mol(-1), depending on environment) resulted in an increase in final leaf size for Populus trichocarpa x Populus deltoides (Populus x interamericana) and P. deltoides x Populus nigra (Populus x euramericana), irrespective of whether plants were exposed during a short-term CE glasshouse study (90 days), a long-term OTC experiment (3 years) or during the first year of a POPFACE experiment. An exception was observed in the closed canopy POPFACE experiment, where final leaf size remained unaltered by CO2. Increased leaf extension rate was observed in elevated CO2 in all experiments, at some point during leaf development, as determined by leaf length. Again the exception were the POPFACE experiment, where effects were not statistically significant. Leaf production and specific leaf area (SLA) were increased and decreased, respectively, on five out of six occasions, although both were only statistically significant on two occasions and interestingly for SLA never in the FACE experiment. Although both cell expansion and cell production were sensitive to CO2 concentration, effects appeared highly dependent on growth environment and genotype. However, increased leaf cell expansion in elevated CO2 was often associated with changes in the biophysical properties of the cell wall, usually increased cell wall plasticity. This research has shown that enhanced leaf area development was a consistent response to elevated CO2 but that the magnitude of this response is likely to decline, in long-term exposure to elevated CO2. Effects on SLA and leaf production suggest that CE and OTC experiments may not always provide good predictors of the 'qualitative' effects of elevated CO2 in long-term ecosystem experiments.     

Surface runoff and nitrogen (N) loss in a bamboo (Phyllostachys pubescens) forest under different fertilization regimes

Nitrogen (N) losses from agricultural fields have been extensively studied. In contrast, surface runoff and N losses have rarely been considered for bamboo forests that are widespread in regions such as southern China. The thriving of bamboo industries has led to increasing fertilizer use in bamboo forests. In this study, we evaluated surface runoff and N losses in runoff following different fertilization treatments under field conditions in a bamboo (Phyllostachys pubescens) forest in the catchment of Lake Taihu in Jiangsu, China. Under three different fertilization regimes, i.e., control, site-specific nutrient management (SSNM), and farmer's fertilization practice (FFP), the water runoff rate amounted to 356, 361, and 342 m³?ha^?1 and accounted for 1.91, 1.98, and 1.85 % of the water input, respectively, from June 2009 to May 2010. The total N losses via surface runoff ranged from 1.2 to 1.8 kg?ha^?1. Compared with FFP, the SSNM treatment reduced total nitrogen (TN) and dissolved nitrogen (DN) losses by 31 and 34 %, respectively. The results also showed that variations in N losses depended mainly on runoff fluxes, not N concentrations. Runoff samples collected from all treatments throughout the year showed TN concentrations greater than 0.35 mg?L^?1, with the mean TN concentration in the runoff from the FFP treatment reaching 8.97 mg?L^?1. The loss of NO₃ ^?–N was greater than the loss of NH₄ ⁺–N. The total loss of dissolved organic nitrogen (DON) reached 23–41 % of the corresponding DN. Therefore, DON is likely the main N species in runoff from bamboo forests and should be emphasized in the assessment and management of N losses in bamboo forest.     

Impact of short-term and long-term elevated CO2 on emission of carbonyls from adult Quercus petraea and Carpinus betulus trees

The study aimed to elucidate the effect of elevated CO2 in short- and long-term experiments on the emission potential of carbonyls from mature oak (Quercus petraea) and hornbeam (Carpinus betulus). The trees were investigated using a canopy crane established in a mixed forest in Switzerland. Short-term CO2 experiments were performed on single leaves with a gas exchange measuring unit by increasing CO2 in the enclosure before and during the measurements. Long-term CO2 fumigation was performed for one vegetation period by a webFACE design. Short-term as well as long-term exposure of leaves resulted in no significant changes in carbonyl exchange in neither Q. petraea nor C. betulus. Carbonyl emission was not affected by stomatal closure under elevated CO2 levels. In contrast to the emission of isoprenoids from vegetation which is thought to be reduced at elevated atmospheric CO2, the exchange of carbonyls seems to be generally unaffected under these conditions.     

Enhanced UV-B and elevated CO(2) impacts sub-arctic shrub berry abundance, quality and seed germination

This study investigated the effects of long-term-enhanced UV-B, and combined UV-B with elevated CO(2) on dwarf shrub berry characteristics in a sub-arctic heath community. Germination of Vaccinium myrtillus was enhanced in seeds produced at elevated UV-B, but seed numbers and berry size were unaffected. Elevated UV-B and CO(2) stimulated the abundance of V. myrtillus berries, whilst UV-B alone stimulated the berry abundance of V. vitis-idaea and Empetrum hermaphroditum. Enhanced UV-B reduced concentrations of several polyphenolics in V. myrtillus berries, whilst elevated CO(2) increased quercetin glycosides in V. myrtillus, and syringetin glycosides and anthocyanins in E. hermaphroditum berries. UV-B?×?CO(2) interactions were found for total anthocyanins, delphinidin-3-hexoside and peonidin-3-pentosidein in V. myrtillus berries but not E. hermaphroditum. Results suggest positive impacts of UV-B on the germination of V. myrtillus and species-specific impacts of UV-B?×?elevated CO(2) on berry abundance and quality. The findings have relevance and implications for human and animal consumers plus seed dispersal and seedling establishment.     

Changes in susceptibility of beech (Fagus sylvatica) seedlings towards Phytophthora citricola under the influence of elevated atmospheric CO2 and nitrogen fertilization

The growth-differentiation balance hypothesis (GDBH) predicts changes in susceptibility of plants against herbivores with changing resource availability. In the presented study we tested the validity of the GDBH for trees infected with a root pathogen. For this purpose Fagus sylvatica seedlings grown under different atmospheric CO₂- and soil nitrogen regimes were infected with the root pathogen Phytophthora citricola. High nitrogen supply increased total biomass of beech regardless of the CO₂-treatment, whereas elevated CO₂ enhanced biomass only in the high nitrogen treatment. The responses of beech under the different growing regimes to the Phytophthora root infection were not in line with the predictions of the GDBH. Enhanced susceptibility of beech against P. citricola was found in seedlings grown under elevated CO₂ and low nitrogen supply. Fifteen months after inoculation these plants were characterized by enhanced water use efficiency, by altered root-shoot ratios, and by enhanced specific root tip densities.     

Spatial and seasonal variations of elemental composition in Mt. Everest (Qomolangma) snow/firn

In May 2005, a total of 14 surface snow (0–10 cm) samples were collected along the climbing route from the advanced base camp to the summit (6500–8844 m a.s.l.) on the northern slope of Mt. Everest (Qomolangma). A 108 m firn/ice core was retrieved from the col of the East Rongbuk Glacier (28.03°N, 86.96°E, 6518 m a.s.l.) on the north eastern saddle of Mt. Everest in September 2002. Surface snow and the upper 3.5 m firn samples from the core were analyzed for major and trace elements by inductively coupled plasma mass spectroscopy (ICP-MS). Measurements show that crustal elements dominated both surface snow and the firn core, suggesting that Everest snow chemistry is mainly influenced by crustal aerosols from local rock or prevalent spring dust storms over southern/central Asia.There are no clear trends for element variations with elevation due to local crustal aerosol inputs or redistribution of surface snow by strong winds during the spring. Seasonal variability in snow/firn elements show that high elemental concentrations occur during the non-monsoon season and low values during the monsoon season. Ca, Cr, Cs, and Sr display the most distinct seasonal variations. Elemental concentrations (especially for heavy metals) at Mt. Everest are comparable with polar sites, generally lower than in suburban areas, and far lower than in large cities. This indicates that anthropogenic activities and heavy metal pollution have little effect on the Mt. Everest atmospheric environment. Everest firn core REE concentrations are the first reported in the region and seem to be comparable with those measured in modern and Last Glacial Maximum snow/ice samples from Greenland and Antarctica, and with precipitation samples from Japan and the East China Sea. This suggests that REE concentrations measured at Everest are representative of the background atmospheric environment.     

Responses of CH(4), CO(2) and N(2)O fluxes to increasing nitrogen deposition in alpine grassland of the Tianshan Mountains

To assess the effects of nitrogen (N) deposition on greenhouse gas (GHG) fluxes in alpine grassland of the Tianshan Mountains in central Asia, CH₄, CO₂ and N₂O fluxes were measured from June 2010 to May 2011. Nitrogen deposition tended to significantly increase CH₄ uptake, CO₂ and N₂O emissions at sites receiving N addition compared with those at site without N addition during the growing season, but no significant differences were found for all sites outside the growing season. Air temperature, soil temperature and water content were the important factors that influence CO₂ and N₂O emissions at year-round scale, indicating that increased temperature and precipitation in the future will exert greater impacts on CO₂ and N₂O emissions in the alpine grassland. In addition, plant coverage in July was also positively correlated with CO₂ and N₂O emissions under elevated N deposition rates. The present study will deepen our understanding of N deposition impacts on GHG balance in the alpine grassland ecosystem, and help us assess the global N effects, parameterize Earth System models and inform decision makers.     

Impact of coal mine dump contaminated soils on elemental uptake by Spinacia oleracea (spinach)

The elemental uptake and the growth response of Spinacia oleracea (spinach) to the soil contaminated with the South African bituminous coal mine dump soil, viz. 0%, 5%, 15%, and 25% w/w, was investigated. The contaminated soils were analyzed for pH, cation exchange capacity (CEC), soil organic matter (SOM), and concentrations of selected heavy metals. The pH, SOM, and CEC decreased with an increase in contamination indicating the acidic nature of coal mine soil and the raise in the soil binding sites. The distribution of Fe, Mn, Ni, Cd, and Pb in the in roots and leaves of the plants was determined in two stages of plant growth. Spinach showed high accumulation of Fe and increased levels of Ni and Cd with an increase in contamination. No plant growth was recorded with 25% contamination.     

Effects of decadal exposure to interacting elevated CO2 and/or O3 on paper birch (Betula papyrifera) reproduction

We studied the effects of long-term exposure (nine years) of birch (Betula papyrifera) trees to elevated CO(2) and/or O(3) on reproduction and seedling development at the Aspen FACE (Free-Air Carbon Dioxide Enrichment) site in Rhinelander, WI. We found that elevated CO(2) increased both the number of trees that flowered and the quantity of flowers (260% increase in male flower production), increased seed weight, germination rate, and seedling vigor. Elevated O(3) also increased flowering but decreased seed weight and germination rate. In the combination treatment (elevated CO(2)+O(3)) seed weight is decreased (20% reduction) while germination rate was unaffected. The evidence from this study indicates that elevated CO(2) may have a largely positive impact on forest tree reproduction and regeneration while elevated O(3) will likely have a negative impact.     

Toxic effects of anthraquinone and phenanthrenequinone upon Scenedesmus strains (green algae) at low and elevated concentration of CO2

Short-term (24h) experiments were performed to examine the effect of anthraquinone (ANTQ) and phenanthrenequinone (PHEQ) on two Scenedesmus armatus strains (B1-76 and 276-4d) grown in a batch culture system aerated with CO2 at a low (0.1%) or elevated (2%) concentration. ANTQ at concentrations within the range of 0.156-1.250 mg dm-3 inhibited the growth of B1-76 population in a concentration-dependent manner, and calculated EC50 for low-CO2 cells was 0.56 mg dm-3. The toxic effect of ANTQ on this strain was more pronounced in high-CO2 cells, where not only growth but also photosynthesis, respiration and SOD activity were significantly inhibited. In contrast, except for SOD activity, no ANTQ effects on strain 276-4d were found. PHEQ at concentrations within the range of 0.063-0.125 mg dm-3 inhibited the growth of B1-76 population in a concentration-dependent manner. The value of EC50 for low-CO2 B1-76 cells was 0.10 mg dm-3. PHEQ inhibited the growth of both strains regardless of CO2 concentration. In B1-76 cells affected by PHEQ, inhibition of photosynthesis was independent of the CO2 level, whereas the SOD activity was much higher in cultures aerated with 2% than with 0.1% CO2. Higher toxicity of PHEQ to strain 276-4d grown at 2% CO2 was accompanied by strong inhibition of photosynthesis, while in low-CO2 cells this process was slightly stimulated. The SOD activity in both low- and high-CO2 cells of strain 276-4d treated with PHEQ was 2-3 times higher compared with the controls. The pattern of SOD isoforms (PAGE analysis) obtained from cells exposed to ANTQ or PHEQ did not change compared with the controls, but the location of the SOD isoforms bands on gel was affected by the concentration of CO2. The results suggest that the strain-specific toxicity of ANTQ and PHEQ may result from oxidative stress. In addition, carbon dioxide appears to play an important role in the toxicity of quinones to algae.     

Impacts of elevated CO2 and/or O3 on leaf ultrastructure of aspen (Populus tremuloides) and birch (Betula papyrifera) in the aspen FACE experiment.

Impacts of elevated atmospheric O3 and/or CO2 on three clones of aspen (Populus tremuloides Michx.) and birch (Betula papyrifera Marsh.) were studied to determine, whether or not elevated CO2 ameliorates O3-induced damage to leaf cells. The plants were exposed for 3 years at the Aspen FACE exposure site in Wisconsin (USA) prior to sampling for ultrastructural investigations on 19 June 1999. In the aspen clones, elevated CO2 increased chloroplast cover index, leaf and spongy mesophyll layer thickness, intercellular air space volume in mesophyll, amount of starch in chloroplasts and cytoplasmic lipids but decreased the number of plastoglobuli in chloroplasts. In contrast, elevated O3 decreased chloroplast cover index, starch content, and the proportion of cytoplasm and intercellular space in mesophyll, and increased the proportion of vacuoles, the amount of condensed vacuolar tannins and the number of plastoglobuli. Ozone also caused structural thylakoid injuries (dilation, distortion) and stromal condensation in chloroplasts, which was ameliorated by elevated CO2 by 5-66% in aspen clones and by 2-10% in birch. Birch ultrastructure was less affected by elevated CO2 or O3 stress compared to aspen. In the most O3-sensitive aspen clone, thinner leaves and cell walls, lower proportion of cell wall volume, and higher volume for vacuoles was found compared to more-tolerant clones.     

Di(2-ethylhexyl)phthalate and mono(2-ethylhexyl)phthalate in media for in vitro fertilization

In vitro fertilization (IVF) is one of the most important treatments of infertility to provide a chance of conceiving. In IVF treatment, sperm are washed and motile sperm are isolated with sperm washing media (SWM) for the purpose of fertilization; fertilized ova are then incubated for a maximum of 5 or 6 d in media for IVF (IVFM). The exposure of fertilized ova to chemicals via such media has not been studied. We determined the concentrations of two contaminants; di(2-ethylhexyl)phthalate (DEHP) and its hydrolyzed product mono(2-ethylhexyl)phthalate (MEHP) in IVFM, SWM, and protein sources (PS: human serum albumin or serum substitute) for IVFM and SWM. The DEHP and MEHP in these media were extracted by a liquid-liquid extraction method and their concentrations determined by liquid chromatography coupled with tandem mass spectrometry (LC-MS/MS). Fifteen IVFM, nine SWM, and six PS obtained in Japan were examined. The concentrations of DEHP and MEHP in IVFM and SWM were <10-114 and <2.0-263 ng mL⁻¹, respectively. The concentrations of both DEHP and MEHP were higher in the media containing PS than in those without PS. Either MEHP alone or both DEHP and MEHP were detected in PS. The concentrations of DEHP and MEHP in PS were <10-982 and 47.0-1840 ng mL⁻¹, respectively. The DEHP and MEHP detected in these media were derived from PS. This is the first study on the chemical contamination of IVFM, SWM, and PS.     

Production and dispersal of Colletotrichum gloeosporioides spores on Stylosanthes scabra under elevated CO2

This paper reports the effect of twice-ambient (700 ppm) atmospheric CO(2) concentration on infection, disease development, spore production and dispersal of the anthracnose pathogen Colletotrichum gloeosporioides in susceptible (Fitzroy) and partially resistant (Seca) cultivars of the tropical pasture legume Stylosanthes scabra under controlled environment and field conditions. Reduction in plant height due to anthracnose was partially compensated for by growth enhancement at elevated CO(2) in Fitzroy but not in Seca. Anthracnose severity was reduced under elevated CO(2) although the reduction was only significant in Fitzroy. Delayed and reduced germination, germtube growth and appressoria production were partly responsible for the reduced severity. Despite an extended incubation period, C. gloeosporioides developed sporulating lesions faster and produced more spores per day within the same latent period at high CO(2) and ambient CO(2). When Fitzroy seedlings grown at 700 ppm CO(2) were exposed to pathogen inoculum under field conditions, they consistently developed more severe anthracnose with more lesions than seedlings grown at ambient CO(2). The environmental variable, which correlated most strongly with the dispersal and infection of C. gloeosporioides spores in the field, was relative humidity in plant canopy. We have shown that an enlarged Stylosanthes canopy under elevated CO(2) can trap more spores, which can lead to more severe anthracnose under favorable weather. The implications of these findings for perennial Stylosanthes pastures are discussed.     

Differential responses of dominant and rare epiphytic bacteria from a submerged macrophyte to elevated CO2

Environmental Science and Pollution Research - Epiphytic bacteria develop complex interactions with their host macrophytes and play an important role in the ecological processes in freshwater...     

Effects of elevated CO2, nitrogen supply and tropospheric ozone on spring wheat-II. Nutrients (N, P, K, S, Ca, Mg, Fe, Mn, Zn)

CO(2) enrichment is expected to alter leaf demand for nitrogen and phosphorus in plant species with C(3) carbon dioxide fixation pathway, thus possibly causing nutrient imbalances in the tissues and disturbance of distribution and redistribution patterns within the plants. To test the influence of CO(2) enrichment and elevated tropospheric ozone in combination with different nitrogen supply, spring wheat (Tritium aestivum L. cv. Minaret) was exposed to three levels of CO(2) (361, 523, and 639 microl litre(-1), 24 h mean from sowing to final harvest), two levels of ozone (28.4 and 51.3 nl litre(-1)) and two levels of nitrogen supply (150 and 270 kg ha(-1)) in a full-factorial design in open-top field chambers. Additional fertilization experiments (120, 210, and 330 kg N ha(-1)) were carried out at low and high CO(2) levels. Macronutrients (N, P, K, S, Ca, Mg) and three micronutrients (Mn, Fe, Zn) were analysed in samples obtained at three different developmental stages: beginning of shoot elongation, anthesis, and ripening. At each harvest, plant samples were separated into different organs (green and senescent leaves, stem sections, ears, grains). According to analyses of tissue concentrations at the beginning of shoot elongation, the plants were sufficiently equipped with nutrients. Elevated ozone levels neither affected tissue concentrations nor shoot uptake of the nutrients. CO(2) and nitrogen treatments affected nutrient uptake, distribution and redistribution in a complex manner. CO(2) enrichment increased nitrogen-use efficiency and caused a lower demand for nitrogen in green tissues which was reflected in a decrease of critical nitrogen concentrations, lower leaf nitrogen concentrations and lower nitrogen pools in the leaves. Since grain nitrogen uptake during grain filling depended completely on redistribution from vegetative pools in green tissues, grain nitrogen concentrations fell considerably with severe implications for grain quality. Ca, S, Mg and Zn in green tissues were influenced by CO(2) enrichment in a similar manner to nitrogen. Phosphorus concentrations in green tissues, on the other hand, were not, or only slightly, affected by elevated CO(2). In stems, 'dilution' of all nutrients except manganese was observed, caused by the huge accumulation of water soluble carbohydrates, mainly fructans, in these tissues under CO(2) enrichment. Whole shoot uptake was either remarkably increased (K, Mn, P, Mg), nearly unaffected (N, S, Fe, Zn) or decreased (Ca) under CO(2) enrichment. Thus, nutrient cycling in plant-soil systems is expected to be altered under CO(2) enrichment.