Responses to elevated temperature and CO2 in the perennial grass Agrostis curtisii in relation to population origin

1. Evolutionary responses to climate change will depend on the presence of heritable variation within species populations for traits that increase fitness under the changing conditions. Patterns of ecotypic differentiation in relation to latitude in some species suggest that such variation exists in relation to temperature responses. Response to elevated CO₂, whether heritable or not, is not expected to be related to latitudinal or climatic differences within temperate regions.
2. To test these ideas, seeds were collected from 10 populations of the outbreeding perennial grass Agrostis curtisii across its range in Europe from south Wales to Portugal. Plants were grown under ambient and elevated temperature and CO₂ conditions, in a factorial design, in solardomes; two half sibs from each population were planted in separate pots in each of the two replicate domes with each combination of treatments. One half sib was harvested at the end of the first summer, the second at the end of the second summer.
3. Survival was uniformly high and flowering uniformly low across treatments and populations.
4. Responses to temperature and CO₂ treatments varied over time for almost all populations. Treatment effects were not significant on plants harvested in year 1, although there was a trend towards higher shoot biomass under the elevated temperature and CO₂ treatment. In year 2 shoot biomass was significantly higher under the elevated temperature treatment across all populations and there was a strong trend towards decreased biomass under elevated CO₂.
5. There were no significant correlations of plant response to either CO₂ or temperature with climate at origin.
6. These results warn of the dangers of extrapolating evolutionary plant responses to CO₂ from short-term experiments.     

Nitrogen‐15 budget in model ecosystems of white clover and perennial ryegrass exposed for four years at elevated atmospheric pCO2

Although there are many indications that N cycling in grassland ecosystems changes under elevated atmospheric CO₂ partial pressure (pCO₂), most information has been obtained in short‐term studies. Thus, N budgets were established for four years under ambient and 60 Pa pCO₂ at two levels of N fertilization in two contrasting model ecosystems: Trifolium repens L. (white clover) and Lolium perenne L. (perennial ryegrass) were planted in soil in boxes in the Swiss FACE experiment. While T. repens showed an 80% increase in harvested biomass with no change in biomass allocation under elevated atmospheric pCO₂ compared to ambient conditions, L. perenne showed an increase only in the biomass of the roots. During the four years of the experiment, the systems gained N both from N retained in the soil and from stubble/stolon and roots left after the final harvest; in total between 11 and 86 gN m⁻². Nitrogen retention in the soil was between 4 and 64 g m². The L. perenne system gained the most N and retained the most N in the soil at high N fertilization and elevated atmospheric pCO₂. The input of new C and N into the soil correlated well in the L. perenne systems but not in the T. repens systems. Elevated atmospheric pCO₂ led neither to an increase in N retention in the soil nor did it reduce the loss of N from the soil. In the L. perenne systems, N fertilization played the main role in both the retention of N and the sequestration of C, while in the T. repens systems symbiotic N₂ fixation may have controlled N retention in the soil.     

Carbon and water fluxes in a calcareous grassland under elevated CO2

1. As part of a long-term study of the effects of elevated CO₂ on biodiversity and ecosystem function in a calcareous grassland, we measured ecosystem carbon dioxide and water-vapour fluxes over 24-h periods during the 1994 and 1995 growing seasons. Data were used to derive CO₂ and H₂O gas-exchange response functions to quantum flux density (QFD).
2. The relative increase in net ecosystem CO₂ flux (NEC) owing to CO₂ enrichment increased as QFD rose. Daytime NEC at high QFD under elevated CO₂ increased by 25% to 60%, with the greatest increases in the spring and after mowing in June when above-ground biomass was lowest. There was much less stimulation of NEC in early June and again in October when the canopy was fully developed. Night-time NEC was not significantly altered under elevated CO₂.
3. Short-term reversal of CO₂ concentrations between treatments after two seasons of CO₂ exposure provided evidence for a 50% downward adjustment of NEC expressed per unit above-ground plant dry weight. However, when expressed on a land area basis, this difference disappeared because of a c. 20% increase in above-ground biomass under elevated CO₂.
4. Ecosystem evapotranspiration (ET) was not significantly altered by elevated CO₂ when averaged over all measurement dates and positions. However, ET was reduced 3–18% at high QFD in plots at the top of the slope at our study site. In summary, CO₂ enrichment resulted in a large stimulation of ecosystem CO₂ capture, especially during periods of a large demand of carbon in relationship to its supply, and resulted in a relatively small and variable effect on ecosystem water consumption.     

Root competition and elevated CO2: effects on seedling growth in Linum usitatissimum populations and Linum–Silene cretica mixtures

1. Root competition can be an important determinant of the performance of neighbours within plant populations and communities. Because plants often maintain larger root systems and allocate more of their carbon to root systems under elevated atmospheric CO₂ than they do at lower CO₂ concentrations, root–root interactions could play an increasingly important role in determining competitive outcomes among individuals and plant species as global CO₂ concentration continues to rise.
2. We established 12 pure stands of Linum usitatissimum (flax) and 12 mixed stands of Linum and its naturally co-occurring weed species Silene cretica in opaque plastic trays each filled with the same amount of nutrient-rich soil mix. In half of the trays from each of these stand types, vertical waterproof partitions separated the root systems of individual plants from each other to prevent root competition, while in the other half no partitions were present. Half of the trays from all treatments were allowed to grow under low atmospheric CO₂ concentration (320μll^–1) and the other half under elevated CO₂ (600μll^–1), in daylight growth chambers for 30 days from seedling emergence until harvest in mid-June. All trays received equal amounts of water so that soils in the low CO₂ treatment were maintained at field capacity.
3. Our results indicate that under high soil fertilities: (1) intra-specific root–root interactions alone play a relatively insignificant role in determining plant biomass production within pure Linum populations and (2) the impact of an aggressive species ( Silene ) on co-occurring less aggressive species ( Linum ) becomes more severe under elevated CO₂ as a result of amplified interspecific root competition.     

Does photosynthetic acclimation to elevated CO2 increase photosynthetic nitrogen-use efficiency? A study of three native UK grassland species in open-top chambers

1. The photosynthetic response to elevated CO₂ and nutrient stress was investigated in Agrostis capillaris, Lolium perenne and Trifolium repens grown in an open-top chamber facility for 2 years under two nutrient regimes. Acclimation was evaluated by measuring the response of light-saturated photosynthesis to changes in the substomatal CO₂ concentration.
2. Growth at elevated CO₂ resulted in reductions in apparent Rubisco activity in vivo in all three species, which were associated with reductions of total leaf nitrogen content on a unit area basis for A. capillaris and L. perenne . Despite this acclimation, photosynthesis was significantly higher at elevated CO₂ for T. repens and A. capillaris , the latter exhibiting the greatest increase of carbon uptake at the lowest nutrient supply.
3. The photosynthetic nitrogen-use efficiency (the rate of carbon assimilation per unit leaf nitrogen) increased at elevated CO₂, not purely owing to higher values of photosynthesis at elevated CO₂, but also as a result of lower leaf nitrogen contents.
4. Contrary to most previous studies, this investigation indicates that elevated CO₂ can stimulate photosynthesis under a severely limited nutrient supply. Changes in photosynthetic nitrogen-use efficiency may be a critical determinant of competition within low nutrient ecosystems and low input agricultural systems.     

Comparison of gas use efficiency and treatment uniformity in a forest ecosystem exposed to elevated [CO2] using pure and prediluted free-air CO2 enrichment technology

A direct comparison of treatment uniformity and CO₂ use of pure and prediluted free-air CO₂ enrichment (FACE) systems was conducted in a forest ecosystem. A vertical release pure CO₂ fumigation system was superimposed on an existing prediluted CO₂ fumigation system and operated on alternate days. The FACE system using prediluted CO₂ fumigation technology exhibited less temporal and spatial variability than the pure CO₂ fumigation system. The pure CO₂ fumigation system tended to over-fumigate the upwind portions of the plot and used 25% more CO₂ than the prediluted CO₂ fumigation system. The increased CO₂ use by the pure CO₂ system was exacerbated at low wind speeds. It is not clear if this phenomenon will also be observed in plots with smaller diameters and low-stature vegetation.     

In situ responses to elevated CO2 in tropical forest understorey plants

1. Plants growing in deep shade and high temperature, such as in the understorey of humid tropical forests, have been predicted to be particularly sensitive to rising atmospheric CO₂. We tested this hypothesis in five species whose microhabitat quantum flux density (QFD) was documented as a covariable. After 7 (tree seedlings of Tachigalia versicolor and Beilschmiedia pendula ) and 18 months (shrubs Piper cordulatum and Psychotria limonensis, and grass Pharus latifolius ) of elevated CO₂ treatment ( c. 700 μl litre^–1) under mean QFD of less than 11 μmol m^–2 s^–1, all species produced more biomass (25–76%) under elevated CO₂.
2. Total plant biomass tended to increase with microhabitat QFD (daytime means varying from 5 to 11μmol m^–2 s^–1) but the relative stimulation by elevated CO₂ was higher at low QFD except in Pharus .
3. Non-structural carbohydrate concentrations in leaves increased significantly in Pharus (+ 27%) and Tachigalia (+ 40%).
4. The data support the hypothesis that tropical plants growing near the photosynthetic light compensation point are responsive to elevated CO₂. An improved plant carbon balance in deep shade is likely to influence understorey plant recruitment and competition as atmospheric CO₂ continues to rise.     

An open-top chamber for measuring soil respiration and the influence of pressure difference on CO2 efflux measurement

1. A new open-top chamber for measuring CO₂ efflux from the soil is reported here. The new design enables measurement of the equilibrium CO₂ efflux, when there is no detectable pressure difference between the chamber and outside nor leakage of CO₂ into or out of the chamber.
2. In previous dynamic-chamber techniques, the measured CO₂ efflux is dependent on the pressure difference between the inside and outside of the chamber, and a negative pressure difference of –1Pa may cause an order of magnitude increase in measured CO₂ efflux. Although the measured CO₂ efflux is less sensitive to a positive pressure difference than to a negative one, a positive pressure difference of even a few tenths of a Pa will lead to a considerable underestimation in soil CO₂ evolution.
3. The influence of pressure difference on measured CO₂ efflux is negligible in the new design and the estimated CO₂ efflux is close to the undisturbed soil respiration rate. Flow rates up to 8lmin^–1, or air movement over the soil surface up to 55cmmin^–1, will not affect CO₂ evolution from the soil. The influence of pressure difference is related to the type of soil being measured and this has also been reported here for the new design.     

The density dependence of plant responses to elevated CO2

1 Stands of the annual Brassica kaber were grown at a range of six densities in both ambient and elevated CO₂ environments, and measurements of shoot growth were made from seedling emergence through to reproduction.
2 Early in stand development (21 days following emergence), CO₂ enhancement (β) for above-ground biomass was highly density-dependent, ranging from 1.41 at the lowest density (20 plants m−2) to 0.59 at the highest density (652 plants m−2).
3 As stands matured and total biomass exceeded a relatively low threshold level (<10.0 g m^{−2; c.}  20% of final yield), the density-dependence of β disappeared. Above this shoot biomass threshold, β-values remained remarkably stable (β = 0.34) across a broad range of stand biomass, independent of a stand's initial density or age.
4 Average stand-level reproductive β-values at a final harvest were very similar to biomass values (β = 0.38) and, as with biomass values at later stages, showed no apparent density-dependence.
5 These results highlight the importance of considering density and the time-course of stand development simultaneously when assessing the potential for CO₂-induced growth enhancements in plants.     

Beech trees exposed to high CO2 and to simulated summer ozone levels: Effects on photosynthesis, chloroplast components and leaf enzyme activity

Young trees of European beech ( Fagus sylvatica L.) were exposed in a phytotron to different levels of ozone and CO₂ under the climatic simulation of one vegetation period. High ozone levels were simulated similar to high ozone concentration in the field (up to 110 ppb), while CO₂ was added as 300 ppm to the present level of ca 380 ppm. Our study describes different aspects of photosynthesis from the leaf level to the reactions of selected thylakoid components at different harvest times during growth of the beech trees under the different fumigation regimes. Ozone effects appeared in the first weeks of the treatment as a stimulation of chlorophyll fluorescence (F_v/F_m), in oxygen production and in ribulose-1,5-bisphosphate carboxylase/oxygenase activity, while the summer and early autumn harvests showed strong reductions in these parameters. Only phosphoenolpyruvate carboxylase (PEPcase) activity remained higher under high ozone. The effects of high CO₂ appeared in general as a small stimulation in enzyme activity like PEPcase in spring. However, with increasing time of fumigation, reductions of all parameters were observed. Especially chlorophylls showed strong reductions under high CO₂. The combined treatment with high ozone plus high CO₂ resulted mostly in an amelioration of the negative ozone effects, although control levels were not reached.     

Productivity and community structure of ectomycorrhizal fungal sporocarps under increased atmospheric CO2 and O3

Sporocarp production is essential for ectomycorrhizal fungal recombination and dispersal, which influences fungal community dynamics. Increasing atmospheric carbon dioxide (CO₂) and ozone (O₃) affect host plant carbon gain and allocation, which may in turn influence ectomycorrhizal sporocarp production if the carbon available to the ectomycorrhizal fungus is dependant upon the quantity of carbon assimilated by the host. We measured sporocarp production of ectomycorrhizal fungi over 4 years at the Aspen FACE (free air CO₂ enrichment) site, which corresponded to stand ages seven to 10 years. Total mean sporocarp biomass was greatest under elevated CO₂, regardless of O₃ concentration, while it was generally lowest under elevated O₃ with ambient CO₂. Community composition differed significantly among the treatments, with less difference in the final year of the study. Whether this convergence was due to succession or environmental factors is uncertain. CO₂ and O₃ affect ectomycorrhizal sporocarp productivity and community composition, with likely effects on dispersal, colonization and sporocarp-dependent food webs.     

Influences of soil volume and an elevated CO2 level on growth and CO2 exchange for the Crassulacean acid metabolism plant Opuntia ficus-indica

Effects of the current (38 Pa) and an elevated (74 Pa) CO₂ partial pressure on root and shoot areas, biomass accumulation and daily net CO₂ exchange were determined for Opuntia ficus-indica (L.) Miller, a highly productive Crassulacean acid metabolism species cultivated worldwide. Plants were grown in environmentally controlled rooms for 18 weeks in pots of three soil volumes (2 600, 6 500 and 26 000 cm³), the smallest of which was intended to restrict root growth. For plants in the medium-sized soil volume, basal cladodes tended to be thicker and areas of main and lateral roots tended to be greater as the CO₂ level was doubled. Daughter cladodes tended to be initiated sooner at the current compared with the elevated CO₂ level but total areas were similar by 10 weeks. At 10 weeks, daily net CO₂ uptake for the three soil volumes averaged 24% higher for plants growing under elevated compared with current CO₂ levels, but at 18 weeks only 3% enhancement in uptake occurred. Dry weight gain was enhanced 24% by elevated CO₂ during the first 10 weeks but only 8% over 18 weeks. Increasing the soil volume 10-fold led to a greater stimulation of daily net CO₂ uptake and biomass production than did doubling the CO₂ level. At 18 weeks, root biomass doubled and shoot biomass nearly doubled as the soil volume was increased 10-fold; the effects of soil volume tended to be greater for elevated CO₂. The amount of cladode nitrogen per unit dry weight decreased as the CO₂ level was raised and increased as soil volume increased, the latter suggesting that the effects of soil volume could be due to nitrogen limitations.     

Influence of leaf chemistry of Lotus corniculatus (Fabaceae) on larval development of Polyommatus icarus (Lepidoptera, Lycaenidae): effects of elevated CO2 and plant genotype

1. Four Lotus corniculatus genotypes differing in cyanoglycoside and condensed tannin concentrations were grown in either low (350 ppm) or high (700 ppm) atmospheric CO₂ environments. Larval performance, consumption and conversion efficiency of Polyommatus icarus feeding on this plant material were measured.
2. Plants grown under elevated CO₂ contained less cyanoglycosides, more condensed tannins and more starch than control plants. However, water concentration, nitrogen and protein as well as nitrogen concentration in relation to carbon concentration did not differ between CO₂ treatments.
3. The four genotypes differed significantly in condensed tannins, cyanoglucoside, leaf water and leaf nitrogen but no genotype–CO₂ interaction was detected, except for total phenolics and condensed tannins in which two plant genotypes showed stronger increases under elevated CO₂ than the other two.
4. Larvae of P . icarus consumed more plant material and used and converted it more efficiently from plants grown at high atmospheric CO₂.
5. Larvae developed significantly faster and were significantly heavier when fed plant material grown under elevated CO₂. The observed difference in mass disappeared in the pupal and adult stages. However, lipid concentration of adults from the elevated CO₂ treatment was marginally significantly higher than of controls.
6. It is concluded that the higher carbohydrate concentration of L . corniculatus plants grown at elevated CO₂ renders leaves more suitable and better digestible to P . icarus . Furthermore, differences in allelochemicals might influence the palatability of L . corniculatus leaves for this specialist on Fabaceae.     

Stimulation of r- vs. K-selected microorganisms by elevated atmospheric CO2 depends on soil aggregate size

Increased root exudation under elevated atmospheric CO₂ and the contrasting environments in soil macro- and microaggregates could affect microbial growth strategies. We investigated the effect of elevated CO₂ on the contribution of fast- ( r -strategists) and slow-growing ( K -strategists) microorganisms in soil macro- and microaggregates. We fractionated the bulk soil from the ambient and elevated (for 5 years) CO₂ treatments of FACE-Hohenheim (Stuttgart) into large macro- (>2 mm), small macro- (0.25–2.00 mm), and microaggregates (<0.25 mm) using 'optimal moist' sieving. Microbial biomass (C_mic), the maximum specific growth rate (μ), growing microbial biomass (GMB) and lag-period ( t _lag) were estimated by the kinetics of CO₂ emission from bulk soil and aggregates amended with glucose and nutrients. Although C_org and C_mic were unaffected by elevated CO₂, μ values were significantly higher under elevated than ambient CO₂ for bulk soil, small macroaggregates, and microaggregates. Substrate-induced respiratory response increased with decreasing aggregate size under both CO₂ treatments. Based on changes in μ, GMB and lag period, we conclude that elevated atmospheric CO₂ stimulated the r- selected microorganisms, especially in soil microaggregates. Such an increase in r -selected microorganisms indicates acceleration of available C mineralization in soil, which may counterbalance the additional C input by roots in soils in a future elevated atmospheric CO₂ environment.     

Adjustments of net photosynthesis in Solanum tuberosum in response to reciprocal changes in ambient and elevated growth CO2 partial pressures

Single leaf photosynthetic rates and various leaf components of potato ( Solanum tuberosum L.) were studied 1–3 days after reciprocally transferring plants between the ambient and elevated growth CO₂ treatments. Plants were raised from individual tuber sections in controlled environment chambers at either ambient (36 Pa) or elevated (72 Pa) CO₂. One half of the plants in each growth CO₂ treatment were transferred to the opposite CO₂ treatment 34 days after sowing (DAS). Net photosynthesis (P_n) rates and various leaf components were then measured 34, 35 and 37 DAS at both 36 and 72 Pa CO₂. Three-day means of single leaf P_n rates, leaf starch, glucose, initial and total Rubisco activity, Rubisco protein, chlorophyll ( a + b ), chlorophyll ( a/b ), α -amino N, and nitrate levels differed significantly in the continuous ambient and elevated CO₂ treatments. Acclimation of single leaf P_n rates was partially to completely reversed 3 days after elevated CO₂-grown plants were shifted to ambient CO₂, whereas there was little evidence of photosynthetic acclimation 3 days after ambient CO₂-grown plants were shifted to elevated CO₂. In a four-way comparison of the 36, 72, 36 to 72 (shifted up) and 72 to 36 (shifted down) Pa CO₂ treatments 37 DAS, leaf starch, soluble carbohydrates, Rubisco protein and nitrate were the only photosynthetic factors that differed significantly. Simple and multiple regression analyses suggested that negative changes of P_n in response to growth CO₂ treatment were most closely correlated with increased leaf starch levels.     

Using stable isotopes to determine soil carbon input differences under ambient and elevated atmospheric CO2 conditions

Quantitative estimates of soil C input under ambient (35 Pa) and elevated (60 Pa) CO₂-partial pressure (pCO₂) were determined in a Free-Air Carbon dioxide Enrichment (FACE) experiment. To facilitate ¹³C-tracing, Trifolium repens L. was grown in a soil with an initial δ¹³C distinct by at least 5‰ from the δ¹³C of T. repens grown under ambient or elevated pCO₂. A shift in δ¹³C of the soil organic C was detected after one growing season. Calculated new soil C inputs in soil under ambient and elevated pCO₂ were 2 and 3 t ha^–1, respectively. Our findings suggest that under elevated CO₂ conditions, soil C sequestration may be altered by changes in plant biomass production and quality.     

Elevated atmospheric CO2 alters wood production, wood quality and wood strength of Scots pine (Pinus sylvestris L) after three years of enrichment

Scots pine ( Pinus sylvestris L.) trees were grown in open top chambers for three years under ambient and elevated CO₂ concentrations. The trees were aged 3 y at the beginning of the CO₂ exposure, and the effects of the treatment on total stem volume, stem wood biomass, wood quality and wood anatomy were examined at the end of the exposure. The elevated CO₂ treatment lead to a 49% and 38% increase in stem biomass and stem wood volume, respectively. However, no significant effects of the elevated CO₂ treatment on wood density were observed, neither when green wood density was estimated from stem biomass and stem volume, nor when oven-dry wood density was measured on small wood samples. Under elevated CO₂ significantly wider growth rings were observed. The effect of elevated CO₂ on growth ring width was primarily the result of an increase in earlywood width. Wood compression strength decreased under elevated CO₂ conditions, which could be explained by significantly larger tracheids and the increased earlywood band, that has thinner walls and larger cavities. A significant decrease of the number of resin canals in the third growth ring was observed under the elevated treatment; this might indicate that trees produced and contained less resin, which has implications for disease and pest resistance. So, although wood volume yield in Scots pine increased significantly with elevated CO₂ after three years of treatment, wood density remained unchanged, while wood strength decreased. Whilst wood volume and stem biomass production may increase in this major boreal forest tree species, wood quality and resin production might decrease under future elevated CO₂ conditions.     

Species-specific responses to atmospheric carbon dioxide and tropospheric ozone mediate changes in soil carbon

We repeatedly sampled the surface mineral soil (0–20 cm depth) in three northern temperate forest communities over an 11-year experimental fumigation to understand the effects of elevated carbon dioxide (CO₂) and/or elevated phyto-toxic ozone (O₃) on soil carbon (C). After 11 years, there was no significant main effect of CO₂ or O₃ on soil C. However, within the community containing only aspen ( Populus tremuloides Michx.), elevated CO₂ caused a significant decrease in soil C content. Together with the observations of increased litter inputs, this result strongly suggests accelerated decomposition under elevated CO_2. In addition, an initial reduction in the formation of new (fumigation-derived) soil C by O₃ under elevated CO₂ proved to be only a temporary effect, mirroring trends in fine root biomass. Our results contradict predictions of increased soil C under elevated CO₂ and decreased soil C under elevated O₃ and should be considered in models simulating the effects of Earth's altered atmosphere.     

Effect of enhanced atmospheric CO2 on mycorrhizal colonization and phosphorus inflow in 10 herbaceous species of contrasting growth strategies

1. Ten herbaceous species were grown over a 4-month period under ambient (360 μmol mol^–1) and elevated (610 μmol mol^–1) atmospheric CO₂ conditions. Plants were inoculated with the arbuscular mycorrhizal (AM) fungus Glomus mosseae and given a phosphorus (P) supply which was not immediately available to the plants.
2. Multiple harvests were taken in order to determine whether the effect of elevated CO₂ on mycorrhizal colonization and phosphorus inflow was independent of its effect on plant growth.
3. All species grew faster under elevated CO₂ and carbon partitioning was altered, generally in favour of the shoots. All species responded similarly to elevated CO₂.
4. Elevated CO₂ did not affect the percentage of root length colonized by AM fungi, but the total amount of colonized root length was increased, because the plants were bigger.
5. Elevated CO₂ increased total P content, but had little or no effect on P concentration. At a given age, P inflow was stimulated by elevated CO₂, but when root length was taken into account the CO₂ effect disappeared.
6. In these host species there is no evidence for a direct effect of elevated CO₂ on mycorrhizal functioning, because both internal mycorrhizal colonization and P inflow are unaffected.
7. Future research should concentrate on the potential for carbon flow to the soil via the external mycelial network.     

The Effect of Elevated [CO2] on Uptake and Allocation of 13C and 15N in Beech (Fagus sylvatica L.) during Leafing

Abstract: A continuous dual ¹³CO₂ and ¹⁵NH₄¹⁵NO₃ labelling experiment was undertaken to determine the effects of ambient (350μmol mol^-1) or elevated (700μmol mol^-1) atmospheric CO₂ concentrations on C and N uptake and allocation within 3-year-old beech ( Fagus sylvatica L.) during leafing. After six weeks of growth, total carbon uptake was increased by 63 % (calculated on total C content) under elevated CO₂ but the carbon partitioning was not altered. 56 % of the new carbon was found in the leaves. On a dry weight basis was the content of structural biomass in leaves 10 % lower and the lignin content remained unaffected under elevated as compared to ambient [CO₂]. Under ambient [CO₂] 37 %, and under elevated [CO₂] 51 %, of the lignin C of the leaves derived from new assimilates. For both treatments, internal N pools provided more than 90 % of the nitrogen used for leaf-growth and the partitioning of nitrogen was not altered under elevated [CO₂]. The C/N ratio was unaffected by elevated [CO₂] at the whole plant level, but the C/N ratio of the new C and N uptake was increased by 32 % under elevated [CO₂].