CO<Subscript>2</Subscript> enrichment,nitrogen fertilization and development of freezing tolerance in Norway spruce

Plant growth and adaptation to cold and freezing temperatures in a CO₂-enriched atmosphere have received little attention despite the predicted effects of elevated CO₂ on plant distribution and productivity. Norway spruce [Picea abies (L.) Karst.] seedlings from latitudinally distinct seed sources (66°N and 60°N) were grown for one simulated growth season under controlled conditions in an atmosphere enriched in CO₂ (70 Pa) and at ambient CO₂ (40 Pa), combined factorially with low (3.6 mM) or high (15.7 mM) concentrations of nitrogen fertilization. There was a clear difference between the two provenances in height growth, in the timing of bud set, and in freezing tolerance. Nitrogen fertilization increased height growth in both provenances, while CO₂ enrichment stimulated height growth only in the southern provenance. We found no significant effects of elevated CO₂ or nitrogen fertilization on the timing of bud set. During cold acclimation, freezing tolerance increased from –10°C to –35°C, and there was a marked increase in all soluble sugars except inositol. Elevated CO₂ in combination with high nitrogen led to a slight increased freezing tolerance in both provenances during the early stages of cold acclimation. However, towards the end of cold acclimation, elevated CO₂ and high nitrogen led to reduced freezing tolerance in the southern provenance, while elevated CO₂ and low nitrogen reduced freezing tolerance in the northern provenance. These results suggest that CO₂ enrichment influences the development of freezing tolerance, and that these responses differ with available nitrogen and between provenances.     

The effects of elevated atmospheric CO<Subscript>2</Subscript> and nitrogen amendments on subsurface CO<Subscript>2</Subscript> production and concentration dynamics in a maturing pine forest

Profiles of subsurface soil CO₂ concentration, soil temperature, and soil moisture, and throughfall were measured continuously during the years 2005 and 2006 in 16 locations at the free air CO₂ enrichment facility situated within a temperate loblolly pine (Pinus taeda L.) stand. Sampling at these locations followed a 4 by 4 replicated experimental design comprised of two atmospheric CO₂ concentration levels (ambient [CO₂]^a, ambient + 200 ppmv, [CO₂]^e) and two soil nitrogen (N) deposition levels (ambient, ambient + fertilization at 11.2 g_N m⁻² year⁻¹). The combination of these measurements permitted indirect estimation of belowground CO₂ production and flux profiles in the mineral soil. Adjacent to the soil CO₂ profiles, direct (chamber-based) measurements of CO₂ fluxes from the soil–litter complex were simultaneously conducted using the automated carbon efflux system. Based on the measured soil CO₂ profiles, neither [CO₂]^e nor N fertilization had a statistically significant effect on seasonal soil CO₂, CO₂ production, and effluxes from the mineral soil over the study period. Soil moisture and temperature had different effects on CO₂ concentration depending on the depth. Variations in CO₂ were mostly explained by soil temperature at deeper soil layers, while water content was an important driver at the surface (within the first 10 cm), where CO₂ pulses were induced by rainfall events. The soil effluxes were equal to the CO₂ production for most of the time, suggesting that the site reached near steady-state conditions. The fluxes estimated from the CO₂ profiles were highly correlated to the direct measurements when the soil was neither very dry nor very wet. This suggests that a better parameterization of the soil CO₂ diffusivity is required for these soil moisture extremes.     

The effect of cold-induced increased metabolic rate on the rate of <Superscript>13</Superscript>C and <Superscript>15</Superscript>N incorporation in house sparrows (<Emphasis Type="Italic">Passer domesticus</Emphasis>)

Animals with high metabolic rates are believed to have high rates of carbon and nitrogen isotopic incorporation. We hypothesized that (1) chronic exposure to cold, and hence an increase in metabolic rate, would increase the rate of isotopic incorporation of both ¹³C and ¹⁵N into red blood cells; and (2) that the rate of isotopic incorporation into red blood cells would be allometrically related to body mass. Two groups of sparrows were chronically exposed to either 5 or 22°C and switched from a ¹³C-depleted C₃-plant diet to a more ¹³C-enriched C₄-plant one. We used respirometry to estimate the resting metabolic rate of birds exposed chronically to our two experimental temperatures. The allometric relationship between the rate of ¹³C incorporation into blood and body mass was determined from published data. The of birds at 5°C was 1.9 times higher than that of birds at 22°C. Chronic exposure to a low temperature did not have an effect on the rate of isotopic incorporation of ¹⁵N save for a very small effect on the incorporation of ¹³C. The isotopic incorporation rate of ¹³C was 1.5 times faster than that of ¹⁵N. The fractional rate of ¹³C incorporation into avian blood was allometrically related to body mass with an exponent similar to −1/4. We conclude that the relationship between metabolic rate and the rate of isotopic incorporation into an animal’s tissues is indirect. It is probably mediated by protein turnover and thus more complex than previous studies have assumed.     

Carbon allocation in shoots of alpine treeline conifers in a CO<Subscript>2</Subscript> enriched environment

With a new approach we assessed the relative contribution of stored and current carbon compounds to new shoot growth in alpine treeline conifers. Within a free air CO₂ enrichment experiment at the alpine treeline in Switzerland, ¹³C-depleted fossil CO₂ was used to trace new carbon in the two tree species Larix decidua L. and Pinus uncinata Ramond over two subsequent years. The deciduous L. decidua was found to supply new shoot growth (structural woody part) by 46% from storage. Surprisingly, the evergreen P. uncinata, assumed to use current-year photosynthates, also utilized a considerable fraction of storage (42%) for new wood growth. In contrast, the needles of P. uncinata were built up almost completely from current-year photosynthates. The isotopic composition of different wood carbon fractions revealed a similar relative allocation of current and stored assimilates to various carbon fractions. Elevated CO₂ influenced the composition of woody tissue in a species-specific way, e.g. the water soluble fraction decreased in pine in 2001 but increased in larch in 2002 compared to ambient CO₂. Heavy defoliation applied as an additional treatment factor in the second year of the experiment decreased the lipophilic fraction in current-year wood in both species compared to undefoliated trees. We conclude that storage may play an important role for new shoot growth in these treeline conifers and that altered carbon availability (elevated CO₂, defoliation) results in significant changes in the relative amount of mobile carbon fractions in woody tissue. In particular, stored carbon seems to be of greater importance in the evergreen P. uncinata than has been previously thought.     

Occurrence and altitudinal pattern of C<Subscript>4</Subscript> plants on Qinghai Plateau,Qinghai province,China

The natural occurrence and altitudinal pattern of species with C₄ photosynthesis were investigated on Qinghai Plateau, Qinghai province by using stable carbon isotopes in plant leaves and using additional data from references. A total of 58 species belonging to 10 families and 34 genera were identified using C₄ photosynthetic pathway, which is only 1.66 % of total 3 500 plant species in Qinghai province. The leading two families, i.e. Gramineae (23 species) and Chenopodiaceae (22 species) contain 77.6 % of all C₄ plants in the studied area. The number of C₄ species increased from 1 600 to 2 400 m a.s.l. and then decreases quickly till 4 400 m a.s.l. with one half of C₄ species distributing from 2 200 to 2 800 m a.s.l. (48 %). Eight plant species were found above 4 000 m a.s.l., but the distribution of these species is limited to the south of Qinghai province (low latitude area) where annual mean temperature is above 0 °C, suggesting that low temperature may generally limit the distribution of C₄ plants.     

The role of N<Subscript>2</Subscript> fixation in alleviating N limitation in wetland metaphyton: enzymatic,isotopic, and elemental evidence

The role of nitrogen (N₂) fixation in balancing N supply to wetland metaphyton was assessed by comparing primary production with enzymatic, isotopic, and elemental correlates. Primary production, N₂ fixation (acetylene reduction, AR), phosphatase activity, C:N:P ratio, and N isotopic composition of metaphyton were measured along a nutrient gradient in a freshwater marsh during May through September 2004. N₂ fixation and phosphatase activity in metaphyton were negatively correlated with inorganic N and P concentrations, respectively. Although metaphyton N₂ fixation demonstrated a clear spatial pattern along the nutrient gradient, N₂ fixation rates varied monthly and decreased sharply in September. However, the percent contribution of N₂ fixation to N uptake by metaphyton consistently decreased throughout the summer. Furthermore, the decreased contribution of N₂ fixation to N uptake corresponded with an increase in metaphyton N content during the growing season. Nitrogen isotopic data suggested the sustained importance of an atmospheric N₂ source through September at the most downstream (nutrient poor) site even though the percent contribution of N₂ fixition to N uptake was lowest in that month. This suggests that metaphyton were efficiently accumulating and recycling fixed N₂ in support of primary production. Over the course of the summer, metaphyton primary production showed a weak inverse correlation with metaphyton phosphatase activity (r ² = 0.58). The largest residuals in this regression corresponded to the largest vartiation in metaphyton N content. When metaphyton primary production was normalized to metaphyton N content, production rates for the entire growing season were more strongly inversely correlated with metaphyton phosphatase activity (r ² = 0.78). Results of the study suggest that N₂ fixation in N poor areas may adequately supplement community N requirements in metaphyton, thereby causing limitation by other elemental resources such as P.     

Keeling plots for hummingbirds: a method to estimate carbon isotope ratios of respired CO<Subscript>2</Subscript> in small vertebrates

The carbon isotope composition of an animals breath reveals the composition of the nutrients that it catabolizes for energy. Here we describe the use of Keeling plots, a method widely applied in ecosystem ecology, to measure the ¹³C of respired CO₂ of small vertebrates. We measured the ¹³C of Rufous Hummingbirds (Selasphorus rufus) in the laboratory and of Mourning (Zenaida macroura) and White-winged (Z. asiatica) Doves in the field. In the laboratory, when hummingbirds were fed a sucrose based C3 diet, the ¹³C of respired CO₂ was not significantly different from that of their diet (¹³C_{C3 diet}). The ¹³C of respired CO₂ for C3 fasted birds was slightly, albeit significantly, depleted in ¹³C relative to ¹³C_{C3 diet}. Six hours after birds were shifted to a sucrose based C4 diet, the isotopic composition of their breath revealed that birds were catabolizing a mixture of nutrients derived from both the C3 and the C4 diet. In the field, the ¹³C of respired CO₂ from Mourning and White-winged Doves reflected that of their diets: the CAM saguaro cactus (Carnegeia gigantea) and C3 seeds, respectively. Keeling plots are an easy, effective and inexpensive method to measure ¹³C of respired CO₂ in the lab and the field.     

Elevated CO<Subscript>2</Subscript> Effects on Peatland Plant Community Carbon Dynamics and DOC Production

Northern peatlands are important stores of carbon and reservoirs of biodiversity that are vulnerable to global change. However, the carbon dynamics of individual peatland plant species is poorly understood, despite the potential for rising atmospheric CO₂ to affect the vegetation’s contribution to overall ecosystem carbon function. Here, we examined the effects of 3 years exposure to elevated CO₂ (eCO₂) on (a) peatland plant community composition and biomass, and (b) plant carbon dynamics and the production of dissolved organic carbon (DOC) using a ¹³CO₂ pulse–chase approach. Results showed that under eCO₂, Sphagnum spp. cover declined by 39% (P < 0.05) and Juncus effusus L. cover increased by 40% (P < 0.001). There was a concurrent increase in above- and belowground plant biomass of 115% (P < 0.01) and 96% (P < 0.01), respectively. Vascular species assimilated and turned over more ¹³CO₂-derived carbon than Sphagnum spp. (49% greater turnover of assimilated ¹³C in J. effusus and F. ovina L. leaf tissues compared with Sphagnum, P < 0.01). Elevated CO₂ also produced a 66% rise in DOC concentrations (P < 0.001) and an order of magnitude more ‘new’ exudate ¹³DOC than control samples (24 h after ¹³CO₂ pulse-labelling 2.5 ± 0.5 and 0.2 ± 0.1% in eCO₂ and control leachate, respectively, P < 0.05). We attribute the observed increase in DOC concentrations under eCO₂ to the switch from predominantly Sphagnum spp. to vascular species (namely J. effusus), leading to enhanced exudation and decomposition (litter and peat). The potential for reduced peatland carbon accretion, increased DOC exports and positive feedback to climate change are discussed.     

Long-term effects of elevated atmospheric CO<Subscript>2</Subscript> on species composition and productivity of a southern African C<Subscript>4</Subscript> dominated grassland in the vicinity of a CO<Subscript>2</Subscript> exhalation

We describe the long-term effects of a CO₂ exhalation, created more than 70 years ago, on a natural C₄ dominated sub-tropical grassland in terms of ecosystem structure and functioning. We tested whether long-term CO₂ enrichment changes the competitive balance between plants with C₃ and C₄ photosynthetic pathways and how CO₂ enrichment has affected species composition, plant growth responses, leaf properties and soil nutrient, carbon and water dynamics. Long-term effects of elevated CO₂ on plant community composition and system processes in this sub-tropical grassland indicate very subtle changes in ecosystem functioning and no changes in species composition and dominance which could be ascribed to elevated CO₂ alone. Species compositional data and soil δ¹³C isotopic evidence suggest no detectable effect of CO₂ enrichment on C₃:C₄ plant mixtures and individual species dominance. Contrary to many general predictions C₃ grasses did not become more abundant and C₃ shrubs and trees did not invade the site. No season length stimulation of plant growth was found even after 5 years of exposure to CO₂ concentrations averaging 610 μmol mol⁻¹. Leaf properties such as total N decreased in the C₃ but not C₄ grass under elevated CO₂ while total non-structural carbohydrate accumulation was not affected. Elevated CO₂ possibly lead to increased end-of-season soil water contents and this result agrees with earlier studies despite the topographic water gradient being a confounding problem at our research site. Long-term CO₂ enrichment also had little effect on soil carbon storage with no detectable changes in soil organic matter found. There were indications that potential soil respiration and N mineralization rates could be higher in soils close to the CO₂ source. The conservative response of this grassland suggests that many of the reported effects of elevated CO₂ on similar ecosystems could be short duration experimental artefacts that disappear under long-term elevated CO₂ conditions.     

Photosynthetic CO<Subscript>2</Subscript> uptake in seedlings of two tropical tree species exposed to oscillating elevated concentrations of CO<Subscript>2</Subscript>

Do short-term fluctuations in CO₂ concentrations at elevated CO₂ levels affect net CO₂ uptake rates of plants? When exposed to 600 μl CO₂ l^?1, net CO₂ uptake rates in shoots or leaves of seedlings of two tropical C₃ tree species, teak (Tectona grandis L. f.) and barrigon [Pseudobombax septenatum (Jacq.) Dug.], increased by 28 and 52% respectively. In the presence of oscillations with half-cycles of 20 s, amplitude of ca. 170 μl CO₂ l^?1 and mean of 600 μl CO₂ l^?1, the stimulation in net CO₂ uptake by the two species was reduced to 19 and 36%, respectively, i.e. the CO₂ stimulation in photosynthesis associated with a change in exposure from 370 to 600 μl CO₂ l^?1 was reduced by a third in both species. Similar reductions in CO₂-stimulated net CO₂ uptake were observed in T. grandis exposed to 40-s oscillations. Rates of CO₂ efflux in the dark by whole shoots of T. grandis decreased by 4.8% upon exposure of plants grown at 370 μl CO₂ l^?1 to 600 μl CO₂ l^?1. The potential implications of the observations on CO₂ oscillations and dark respiration are discussed in the context of free-air CO₂ enrichment (FACE) systems in which short-term fluctuations of CO₂ concentration are a common feature.     

Biomass responses to elevated CO<Subscript>2</Subscript>, soil heterogeneity and diversity: an experimental assessment with grassland assemblages

While it is well-established that the spatial distribution of soil nutrients (soil heterogeneity) influences the competitive ability and survival of individual plants, as well as the productivity of plant communities, there is a paucity of data on how soil heterogeneity and global change drivers interact to affect plant performance and ecosystem functioning. To evaluate the effects of elevated CO₂, soil heterogeneity and diversity (species richness and composition) on productivity, patterns of biomass allocation and root foraging precision, we conducted an experiment with grassland assemblages formed by monocultures, two- and three-species mixtures of Lolium perenne, Plantago lanceolata and Holcus lanatus. The experiment lasted for 90 days, and was conducted on microcosms built out of PVC pipe (length 38 cm, internal diameter 10 cm). When nutrients were heterogeneously supplied (in discrete patches), assemblages exhibited precise root foraging patterns, and had higher total, above- and belowground biomass. Greater aboveground biomass was observed under elevated CO₂. Species composition affected the below:aboveground biomass ratio and interacted with nutrient heterogeneity to determine belowground and total biomass. Species richness had no significant effects, and did not interact with either CO₂ or nutrient heterogeneity. Under elevated CO₂ conditions, the two- and three-species mixtures showed a clear trend towards underyielding. Our results show that differences among composition levels were dependent on soil heterogeneity, highlighting its potential role in modulating diversity–productivity relationships. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible to authorized users.     

Nitrogen and carbon isotope responses of Chinese cabbage and chrysanthemum to the application of liquid pig manure

The effects of the liquid pig manure (LM) used in organic farming on the natural abundance of ¹⁵N and ¹³C signatures in plant tissues have not been studied. We hypothesized that application of LM will (1) increase δ¹⁵N of plant tissues due to the high δ¹⁵N of N in LM as compared with soil N or inorganic fertilizer N, and (2) increase δ¹³C of plant tissues as a result of high salt concentration in LM that decreases stomatal conductance of plants. To test these hypotheses, variations in the δ¹⁵N and δ¹³C of Chinese cabbage (Brassica campestris L.) and chrysanthemum (Chrysanthemum morifolium Ramatuelle) with two different LMs (with δ¹⁵N of +15.6 and +18.2‰) applied at two rates (323 and 646 kg N ha^-1 for cabbage and 150 and 300 kg N ha^-1 for chrysanthemum), or urea (δ¹⁵N = -2.7‰) applied at the lower rate above for the respective species, in addition to the control (no N input) were investigated through a 60-day pot experiment. Application of LM significantly increased plant tissue δ¹⁵N (range +9.4 to +14.9‰) over the urea (+3.2 to +3.3‰) or control (+6.8 to 7.7‰) treatments regardless of plant species, strongly reflecting the δ¹⁵N of the N source. Plant tissue δ¹³C were not affected by the treatments for cabbage (range −30.8 to −30.2‰) or chrysanthemum (−27.3 to −26.8‰). However, cabbage dry matter production decreased while its δ¹³C increased with increasing rate of LM application or increasing soil salinity (P < 0.05), suggesting that salinity stress caused by high rate of LM application likely decreased stomatal conductance and limited growth of cabbage. Our study expanded the use of the δ¹⁵N technique in N source (organic vs. synthetic fertilizer) identification and suggested that plant tissue δ¹³C maybe a sensitive indicator of plant response to salinity stress caused by high LM application rates.     

Effect of elevated CO<Subscript>2</Subscript> levels on leaf starch,nitrogen and photosynthesis of plants growing at three natural CO<Subscript>2</Subscript> springs in Japan

Plant communities around natural CO₂ springs have been exposed to elevated CO₂ levels over many generations and give us a unique opportunity to investigate the effects of long-term elevated CO₂ levels on wild plants. We searched for natural CO₂ springs in cool temperate climate regions in Japan and found three springs that were suitable for studying long-term responses of plants to elevated levels of CO₂: Ryuzin-numa, Yuno-kawa and Nyuu. At these CO₂ springs, the surrounding air was at high CO₂ concentration with no toxic gas emissions throughout the growth season, and there was natural vegetation around the springs. At each site, high-CO₂ (HC) and low-CO₂ (LC) plots were established, and three dominant species at the shrub layers were used for physiological analyses. Although the microenvironments were different among the springs, dicotyledonous species growing at the HC plots tended to have more starch and less nitrogen per unit dry mass in the leaves than those growing at the LC plots. In contrast, monocotyledonous species growing in the HC and LC plots had similar starch and nitrogen concentrations. Photosynthetic rates at the mean growth CO₂ concentration were higher in HC plants than LC plants, but photosynthetic rates at a common CO₂ concentration were lower in HC plants. Efficiency of water and nitrogen use of leaves at growth CO₂ concentration was greatly increased in HC plants. These results suggest that natural plants growing in elevated CO₂ levels under cool temperate climate conditions have down-regulated their photosynthetic capacity but that they increased photosynthetic rates and resource use efficiencies due to the direct effect of elevated CO₂ concentration.     

Effects of CO<Subscript>2</Subscript> concentration on acclimatization and physiological responses of two cultivars of carob tree

This study reports survival and physiological responses of micropropagated Ceratonia siliqua L. cvs. Galhosa and Mulata plants during ex vitro acclimatization under ambient (AC; 330 mol mol^–1) or elevated (EC; 810 mol mol^–1) CO₂ concentration and a photosynthetic photon flux density of 125 mol m^–2 s^–1. CO₂ enrichment during acclimatization did not improve survival rate that was around 80 % for both treatments. Eight weeks after ex vitro transplantation, photosynthetic capacity and apparent quantum yield in acclimatized leaves were higher in comparison with those in in vitro-grown leaves, without any significant difference between CO₂ treatments. Chlorophyll content increased after acclimatization. However, EC led to a decrease in the total amount of chlorophyll in new leaves of both cultivars, compared to those grown at AC. Soluble sugars and starch contents were not markedly affected by growth EC, although starch had significantly increased after transfer to ex vitro conditions. EC induced an increase in the stem elongation and in the effective life of leaves, and a decrease in the number of new leaves.     

Genetic dissection of cotton physiological responses to arid conditions and their inter-relationships with productivity

Testing of the extent to which different complex traits share common genetic control provides a means to distinguish associations that are truly diagnostic of genetic potential for improved adaptation to abiotic stress, from incidental phenotypic correlations. In two generations of progeny from a cross between Gossypium hirsutum and Gossypium barbadense, quantitative trait loci (QTL) mapping was used to evaluate correspondence in genetic control of selected physiological measures and productivity under water‐limited and well‐watered environments, respectively. A total of 33 QTLs were detected for five physiological variables [osmotic potential (OP), carbon isotope ratio (δ¹³C; indicator of water use efficiency), canopy temperature, chlorophyll a and b], and 46 QTLs for five measures of crop productivity [dry matter, seed cotton yield (SC), harvest index, boll weight, and boll number]. QTL likelihood intervals for high SC and low OP corresponded in three genomic regions, two of which mapped to homoeologous locations on the two subgenomes of tetraploid cotton. QTLs for δ¹³C showed only incidental association with productivity, indicating that high water use efficiency can be associated with either high or low productivity. Different cotton species have evolved different alleles related to physiological responses and productivity under water deficit, which may permit the development of genotypes that are better‐adapted to arid conditions.     

Effects of salinity on chlorophyll fluorescence and CO<Subscript>2</Subscript> fixation in C<Subscript>4</Subscript> estuarine grasses

The effects of salinity (sea water at 0 ‰ versus 30 ‰) on gross rates of O₂ evolution (J _O2) and net rates of CO₂ uptake (P _N) were measured in the halotolerant estuarine C₄ grasses Spartina patens, S. alterniflora, S. densiflora, and Distichlis spicata in controlled growth environments. Under high irradiance, salinity had no significant effect on the intercellular to ambient CO₂ concentration ratio (C _i/C _a). However, during photosynthesis under limiting irradiance, the maximum quantum efficiency of CO₂ fixation decreased under salinity across species, suggesting there is increased leakage of the CO₂ delivered to the bundle sheath cells by the C₄ pump. Growth under salinity did not affect the maximum intrinsic efficiency of photosystem 2, PS2 (F_V/F_M) in these species, suggesting salinity had no effect on photosynthesis by inactivation of PS2 reaction centers. Under saline conditions and high irradiance, P _N was reduced by 75 % in Spartina patens and S. alterniflora, whereas salinity had no effect on P _N in S. densiflora or D. spicata. This inhibition of P _N in S. patens and S. alterniflora was not due to an effect on stomatal conductance since the ratio of C _i/C _a did not decrease under saline conditions. In growth with and without salt, P _N was saturated at ∼500 μmol(quantum) m⁻² s⁻¹ while J _O2 continued to increase up to full sunlight, indicating that carbon assimilation was not tightly coupled to photochemistry in these halophytic species. This increase in alternative electron flow under high irradiance might be an inherent function in these halophytes for dissipating excess energy.     

The interacting effects of elevated atmospheric CO<Subscript>2</Subscript> concentration,drought and leaf-to-air vapour pressure deficit on ecosystem isoprene fluxes

Isoprene is the most abundant biogenic hydrocarbon released from vegetation and it plays a major role in tropospheric chemistry. Because of its link to climate change, there is interest in understanding the relationship between CO₂, water availability and isoprene emission. We explored the effect of atmospheric elevated CO₂ concentration and its interaction with vapour pressure deficit (VPD) and water stress, on gross isoprene production (GIP) and net ecosystem exchange of CO₂ (NEE) in two Populus deltoides plantations grown at ambient and elevated atmospheric CO₂ concentration in the Biosphere 2 Laboratory facility. Although GIP and NEE showed a similar response to light and temperature, their responses to CO₂ and VPD were opposite; NEE was stimulated by elevated CO₂ and depressed by high VPD, while GIP was inhibited by elevated CO₂ and stimulated by high VPD. The difference in response between isoprene production and photosynthesis was also evident during water stress. GIP was stimulated in the short term and declined only when the stress was severe, whereas NEE started to decrease from the beginning of the experiment. This contrasting response led the carbon lost as isoprene in both the ambient and the elevated CO₂ treatments to increase as water stress progressed. Our results suggest that water limitation can override the inhibitory effect of elevated CO₂ leading to increased global isoprene emissions in a climate change scenario with warmer and drier climate.     

Characterization of an urban-rural CO<Subscript>2</Subscript>/temperature gradient and associated changes in initial plant productivity during secondary succession

To examine the impact of climate change on vegetative productivity, we exposed fallow agricultural soil to an in situ temperature and CO₂ gradient between urban, suburban and rural areas in 2002. Along the gradient, average daytime CO₂ concentration increased by 21% and maximum (daytime) and minimum (nighttime) daily temperatures increased by 1.6 and 3.3°C, respectively in an urban relative to a rural location. Consistent location differences in soil temperature were also ascertained. No other consistent differences in meteorological variables (e.g. wind speed, humidity, PAR, tropospheric ozone) as a function of urbanization were documented. The urban-induced environmental changes that were observed were consistent with most short-term (~50 year) global change scenarios regarding CO₂ concentration and air temperature. Productivity, determined as final above-ground biomass, and maximum plant height were positively affected by daytime and soil temperatures as well as enhanced [CO₂], increasing 60 and 115% for the suburban and urban sites, respectively, relative to the rural site. While long-term data are needed, these initial results suggest that urban environments may act as a reasonable surrogate for investigating future climatic change in vegetative communities.     

Elevated CO<Subscript>2</Subscript> reduces sap flux in mature deciduous forest trees

We enriched in CO₂ the canopy of 14 broad-leaved trees in a species-rich, ca. 30-m-tall forest in NW Switzerland to test whether elevated CO₂ reduces water use in mature forest trees. Measurements of sap flux density (J_S) were made prior to CO₂ enrichment (summer 2000) and throughout the first whole growing season of CO₂ exposure (2001) using the constant heat-flow technique. The short-term responses of sap flux to brief (1.5–3 h) interruptions of CO₂ enrichment were also examined. There were no significant a priori differences in morphological and physiological traits between trees which were later exposed to elevated CO₂ (n=14) and trees later used as controls (n=19). Over the entire growing season, CO₂ enrichment resulted in an average 10.7% reduction in mean daily J_S across all species compared to control trees. Responses were most pronounced in Carpinus, Acer, Prunus and Tilia, smaller in Quercus and close to zero in Fagus trees. The J_S of treated trees significantly increased by 7% upon transient exposure to ambient CO₂ concentrations at noon. Hence, responses of the different species were, in the short term, similar in magnitude to those observed over the whole season (though opposite because of the reversed treatment). The reductions in mean J_S of CO₂-enriched trees were high (22%) under conditions of low evaporative demand (vapour pressure deficit, VPD <5 hPa) and small (2%) when mean daily VPD was greater than 10 hPa. During a relatively dry period, the effect of elevated CO₂ on J_S even appeared to be reversed. These results suggest that daily water savings by CO₂-enriched trees may have accumulated to a significantly improved water status by the time when control trees were short of soil moisture. Our data indicate that the magnitude of CO₂ effects on stand transpiration will depend on rainfall regimes and the relative abundance of the different species, being more pronounced under humid conditions and in stands dominated by species such as Carpinus and negligible in mono-specific Fagus forests.