Salinity and sea level mediate elevated CO2 effects on C3–C4 plant interactions and tissue nitrogen in a Chesapeake Bay tidal wetland

Elevated atmospheric carbon dioxide concentrations ([CO₂]) generally increase plant photosynthesis in C₃ species, but not in C₄ species, and reduce stomatal conductance in both C₃ and C₄ plants. In addition, tissue nitrogen concentration ([N]) often fails to keep pace with enhanced carbon gain under elevated CO₂, particularly in C₃ species. While these responses are well documented in many species, implications for plant growth and nutrient cycling in native ecosystems are not clear. Here we present data on 18 years of measurement of above and belowground biomass, tissue [N] and total standing crop of N for a Scirpus olneyi‐dominated (C₃ sedge) community, a Spartina patens‐dominated (C₄ grass) community and a C₃–C₄‐mixed species community exposed to ambient and elevated (ambient +340 ppm) atmospheric [CO₂] in natural salinity and sea level conditions of a Chesapeake Bay wetland. Increased biomass production (shoots plus roots) under elevated [CO₂] in the S. olneyi‐dominated community was sustained throughout the study, averaging approximately 35%, while no significant effect of elevated [CO₂] was found for total biomass in the C₄‐dominated community. We found a significant decline in C₄ biomass (correlated with rising sea level) and a concomitant increase in C₃ biomass in the mixed community. This shift from C₄ to C₃ was accelerated by the elevated [CO₂] treatment. The elevated [CO₂] stimulation of total biomass accumulation was greatest during rainy, low salinity years: the average increase above the ambient treatment during the three wettest years (1994, 1996, 2003) was 2.9 t ha⁻¹ but in the three driest years (1995, 1999, 2002), it was 1.2 t ha⁻¹. Elevated [CO₂] depressed tissue [N] in both species, but especially in the S. olneyi where the relative depression was positively correlated with salinity and negatively related with the relative enhancement of total biomass production. Thus, the greatest amount of carbon was added to the S. olneyi‐dominated community during years when shoot [N] was reduced the most, suggesting that the availability of N was not the most or even the main limitation to elevated [CO₂] stimulation of carbon accumulation in this ecosystem.     

Soil CO2 efflux of two silver birch clones exposed to elevated CO2 and O3 levels during three growing seasons

In the present open‐top chamber experiment, two silver birch clones (Betula pendula Roth, clone 4 and clone 80) were exposed to elevated levels of carbon dioxide (CO₂) and ozone (O₃), singly and in combination, and soil CO₂ efflux was measured 14 times during three consecutive growing seasons (1999–2001). In the beginning of the experiment, all experimental trees were 7 years old and during the experiment the trees were growing in sandy field soil and fertilized regularly. In general, elevated O₃ caused soil CO₂ efflux stimulation during most measurement days and this stimulation enhanced towards the end of the experiment. The overall soil respiration response to CO₂ was dependent on the genotype, as the soil CO₂ efflux below clone 80 trees was enhanced and below clone 4 trees was decreased under elevated CO₂ treatments. Like the O₃ impact, this clonal difference in soil respiration response to CO₂ increased as the experiment progressed. Although the O₃ impact did not differ significantly between clones, a significant time × clone × CO₂× O₃ interaction revealed that the O₃‐induced stimulation of soil respiration was counteracted by elevated CO₂ in clone 4 on most measurement days, whereas in clone 80, the effect of elevated CO₂ and O₃ in combination was almost constantly additive during the 3‐year experiment. Altogether, the root or above‐ground biomass results were only partly parallel with the observed soil CO₂ efflux responses. In conclusion, our data show that O₃ impacts may appear first in the below‐ground processes and that relatively long‐term O₃ exposure had a cumulative effect on soil CO₂ efflux. Although the soil respiration response to elevated CO₂ depended on the tree genotype as a result of which the O₃ stress response might vary considerably within a single tree species under elevated CO₂, the present experiment nonetheless indicates that O₃ stress is a significant factor affecting the carbon cycling in northern forest ecosystems.     

Elevated atmospheric CO2 stimulates aboveground biomass in a fire-regenerated scrub-oak ecosystem

The effect of elevated atmospheric CO₂ concentration (C_a) on the aboveground biomass of three oak species, Quercus myrtifolia, Q. geminata, and Q. chapmanii, was estimated nondestructively using allometric relationships between stem diameter and aboveground biomass after four years of experimental treatment in a naturally fire‐regenerated scrub‐oak ecosystem. After burning a stand of scrub‐oak vegetation, re‐growing plants were exposed to either current ambient (379 µL L^?1 CO₂) or elevated (704 µL L^?1 CO₂) C_a in 16 open‐top chambers over a four‐year period, and measurements of stem diameter were carried out annually on all oak shoots within each chamber. Elevated C_a significantly increased aboveground biomass, expressed either per unit ground area or per shoot; elevated C_a had no effect on shoot density. The relative effect of elevated C_a on aboveground biomass increased each year of the study from 44% (May 96–Jan 97), to 55% (Jan 97–Jan 98), 66% (Jan 98–Jan 99), and 75% (Jan 99–Jan 00). The effect of elevated C_a was species specific: elevated C_a significantly increased aboveground biomass of the dominant species, Q. myrtifolia, and tended to increase aboveground biomass of Q. chapmanii, but had no effect on aboveground biomass of the subdominant, Q. geminata. These results show that rising atmospheric CO₂ has the potential to stimulate aboveground biomass production in ecosystems dominated by woody species, and that species‐specific growth responses could, in the long term, alter the composition of the scrub‐oak community.     

Altered physiological and growth responses to elevated [CO2] in offspring from holm oak (Quercus ilex L.) mother trees with lifetime exposure to naturally elevated [CO2]

An important question with respect to plant performance in future climatic scenarios is whether the offspring of mature trees that have experienced lifelong exposure to elevated [CO₂] show altered physiological responses to elevated [CO₂] compared with those originating from current ambient CO₂ concentrations. To investigate this question, acorns were collected from two seed sources, denoted as ‘control’ and ‘spring’, from Quercus ilex mother trees grown at ambient (36 Pa) and at about twice ambient CO₂ concentrations, respectively, close to a natural CO₂ spring, Laiatico, central Italy. The seedlings were raised for 8 months under controlled conditions at ambient and elevated [CO₂] in a reciprocal experimental design and were used for the determination of biomass, photosynthesis and foliar carbohydrate concentrations, as well as the accumulation of structural biomass and lignin during leaf maturation. Under ambient [CO₂], biomass and foliar carbon acquisition in control progeny were not significantly different from spring progeny. However, under elevated [CO₂], spring seedlings showed less CO₂ acclimation than control seedlings but no significant differences in non‐structural carbohydrate concentrations and structural biomass per unit leaf dry mass. Developmental lignin accumulation in leaves was delayed under elevated [CO₂] compared with ambient [CO₂], but only in control progeny. Under elevated [CO₂], whole‐plant biomass, leaf area and stem diameter were significantly increased in Quercus ilex seedlings from both seed sources but with a higher stimulation of above‐ground biomass in spring than in control seedlings and a higher stimulation of below‐ground biomass in control seedlings. These results indicate that life history and/or progeny may determine the species‐specific CO₂ response and suggest that positive CO₂ acclimation is possible.     

Net ecosystem CO2 exchange in Mojave Desert shrublands during the eighth year of exposure to elevated CO2

Arid ecosystems, which occupy about 35% of the Earth's terrestrial surface area, are believed to be among the most responsive to elevated [CO₂]. Net ecosystem CO₂ exchange (NEE) was measured in the eighth year of CO₂ enrichment at the Nevada Desert Free‐Air CO₂ Enrichment (FACE) Facility between the months of December 2003–December 2004. On most dates mean daily NEE (24 h) (μmol CO₂ m^?2 s^?1) of ecosystems exposed to elevated atmospheric CO₂ were similar to those maintained at current ambient CO₂ levels. However, on sampling dates following rains, mean daily NEEs of ecosystems exposed to elevated [CO₂] averaged 23 to 56% lower than mean daily NEEs of ecosystems maintained at ambient [CO₂]. Mean daily NEE varied seasonally across both CO₂ treatments, increasing from about 0.1 μmol CO₂ m^?2 s^?1 in December to a maximum of 0.5–0.6 μmol CO₂ m^?2 s^?1 in early spring. Maximum NEE in ecosystems exposed to elevated CO₂ occurred 1 month earlier than it did in ecosystems exposed to ambient CO₂, with declines in both treatments to lowest seasonal levels by early October (0.09±0.03 μmol CO₂ m^?2 s^?1), but then increasing to near peak levels in late October (0.36±0.08 μmol CO₂ m^?2 s^?1), November (0.28±0.03 μmol CO₂ m^?2 s^?1), and December (0.54±0.06 μmol CO₂ m^?2 s^?1). Seasonal patterns of mean daily NEE primarily resulted from larger seasonal fluctuations in rates of daytime net ecosystem CO₂ uptake which were closely tied to plant community phenology and precipitation. Photosynthesis in the autotrophic crust community (lichens, mosses, and free‐living cyanobacteria) following rains were probably responsible for the high NEEs observed in January, February, and late October 2004 when vascular plant photosynthesis was low. Both CO₂ treatments were net CO₂ sinks in 2004, but exposure to elevated CO₂ reduced CO₂ sink strength by 30% (positive net ecosystem productivity=127±17 g C m^?2 yr^?1 ambient CO₂ and 90±11 g C m^?2 yr^?1 elevated CO₂, P=0.011). This level of net C uptake rivals or exceeds levels observed in some forested and grassland ecosystems. Thus, the decrease in C sequestration seen in our study under elevated CO₂– along with the extensive coverage of arid and semi‐arid ecosystems globally – points to a significant drop in global C sequestration potential in the next several decades because of responses of heretofore overlooked dryland ecosystems.     

Biosynthesis of plant phenolic compounds in elevated atmospheric CO2

Experiments were carried out to determine the effects of elevated atmospheric carbon dioxide (CO₂) on phenolic biosynthesis in four plant species growing over three generations for nine months in a model plant community. Results were compared to those obtained when the same species were grown individually in pots in the same soils and controlled environment. In the model herbaceous plant community, only two of the four species showed any increase in biomass under elevated CO₂, but this occurred only in the first generation for Spergula arvensis and in the second generation for Poa annua. Thus, the effects of CO₂ on plant biomass and carbon and nitrogen content were species‐ and generation‐specific. The activity of the principle phenolic biosynthetic enzyme, phenylalanine ammonia lyase (PAL), increased under elevated CO₂ in Senecio vulgaris only in Generation 1, but increased in three of the four plant species in Generation 2. There were no changes in the total phenolic content of the plants, except for P. annua in Generation 1. Lignin content decreased under elevated CO₂ in Cardamine hirsuta in Generation 1, but increased in Generation 2, whilst the lignin content of P. annua showed no change, decreased, then increased in response to elevated CO₂ over the three generations. When the species were grown alone in pots, elevated CO₂ increased PAL activity in plants grown in soil taken from the Ecotron community after nine months of plant growth, but not in plants grown in the soil used at the start of the experiment (‘initial' soil). In P. annua, phenolic biosynthesis decreased under elevated CO₂ in initial soil, and in both P. annua and S. vulgaris there was a significant interaction between effects of soil type and CO₂ level on PAL activity. In this study, plant chemical composition altered more in response to environmental factors such as soil type than in response to carbon supply. Results were species‐specific and changed markedly between generations.     

Responses of CAM species to increasing atmospheric CO2 concentrations

Crassulacean acid metabolism (CAM) species show an average increase in biomass productivity of 35% in response to a doubled atmospheric CO₂ concentration. Daily net CO₂ uptake is similarly enhanced, reflecting in part an increase in chlorenchyma thickness and accompanied by an even greater increase in water‐use efficiency. The responses of net CO₂ uptake in CAM species to increasing atmospheric CO₂ concentrations are similar to those for C₃ species and much greater than those for C₄ species. Increases in net daily CO₂ uptake by CAM plants under elevated atmospheric CO₂ concentrations reflect increases in both Rubisco‐mediated daytime CO₂ uptake and phosphoenolpyruvate carboxylase (PEPCase)‐mediated night‐time CO₂ uptake, the latter resulting in increased nocturnal malate accumulation. Chlorophyll contents and the activities of Rubisco and PEPCase decrease under elevated atmospheric CO₂, but the activated percentage for Rubisco increases and the K_M(HCO₃ ^? ) for PEPCase decreases, resulting in more efficient photosynthesis. Increases in root:shoot ratios and the formation of additional photosynthetic organs, together with increases in sucrose‐Pi synthase and starch synthase activity in these organs under elevated atmospheric CO₂ concentrations, decrease the potential feedback inhibition of photosynthesis. Longer‐term studies for several CAM species show no downward acclimatization of photosynthesis in response to elevated atmospheric CO₂ concentrations. With increasing temperature and drought duration, the percentage enhancement of daily net CO₂ uptake caused by elevated atmospheric CO₂ concentrations increases. Thus net CO₂ uptake, productivity, and the potential area for cultivation of CAM species will be enhanced by the increasing atmospheric CO₂ concentrations and the increasing temperatures associated with global climate change.     

Direct and indirect effects of elevated CO2 on leaf respiration in a forest ecosystem

We measured the short‐term direct and long‐term indirect effects of elevated CO₂ on leaf dark respiration of loblolly pine (Pinus taeda) and sweetgum (Liquidambar styraciflua) in an intact forest ecosystem. Trees were exposed to ambient or ambient + 200 µmol mol^?1 atmospheric CO₂ using free‐air carbon dioxide enrichment (FACE) technology. After correcting for measurement artefacts, a short‐term 200 µmol mol^?1 increase in CO₂ reduced leaf respiration by 7–14% for sweetgum and had essentially no effect on loblolly pine. This direct suppression of respiration was independent of the CO₂ concentration under which the trees were grown. Growth under elevated CO₂ did not appear to have any long‐term indirect effects on leaf maintenance respiration rates or the response of respiration to changes in temperature (Q₁₀, R₀). Also, we found no relationship between mass‐based respiration rates and leaf total nitrogen concentrations. Leaf construction costs were unaffected by growth CO₂ concentration, although leaf construction respiration decreased at elevated CO₂ in both species for leaves at the top of the canopy. We conclude that elevated CO₂ has little effect on leaf tissue respiration, and that the influence of elevated CO₂ on plant respiratory carbon flux is primarily through increased biomass.     

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

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

Biomass allocation in old-field annual species grown in elevated CO2 environments: no evidence for optimal partitioning

Increased atmospheric carbon dioxide supply is predicted to alter plant growth and biomass allocation patterns. It is not clear whether changes in biomass allocation reflect optimal partitioning or whether they are a direct effect of increased growth rates. Plasticity in growth and biomass allocation patterns was investigated at two concentrations of CO₂ ([CO₂]) and at limiting and nonlimiting nutrient levels for four fast‐ growing old‐field annual species. Abutilon theophrasti, Amaranthus retroflexus, Chenopodium album, and Polygonum pensylvanicum were grown from seed in controlled growth chamber conditions at current (350 μmol mol^?1, ambient) and future‐ predicted (700 μmol mol^?1, elevated) CO₂ levels. Frequent harvests were used to determine growth and biomass allocation responses of these plants throughout vegetative development. Under nonlimiting nutrient conditions, whole plant growth was increased greatly under elevated [CO₂] for three C3 species and moderately increased for a C4 species (Amaranthus). No significant increases in whole plant growth were observed under limiting nutrient conditions. Plants grown in elevated [CO₂] had lower or unchanged root:shoot ratios, contrary to what would be expected by optimal partitioning theory. These differences disappeared when allometric plots of the same data were analysed, indicating that CO₂‐induced differences in root:shoot allocation were a consequence of accelerated growth and development rates. Allocation to leaf area was unaffected by atmospheric [CO₂] for these species. The general lack of biomass allocation responses to [CO₂] availability is in stark contrast with known responses of these species to light and nutrient gradients. We conclude that biomass allocation responses to elevated atmospheric [CO₂] are not consistent with optimal partitioning predictions.     

Allocation responses to CO2 enrichment and defoliation by a native annual plant Heterotheca subaxillaris

Among plants grown under enriched atmospheric CO₂, root:shoot balance (RSB) theory predicts a proportionately greater allocation of assimilate to roots than among ambient‐grown plants. Conversely, defoliation, which decreases the plant's capacity to assimilate carbon, is predicted to increase allocation to shoot. We tested these RSB predictions, and whether responses to CO₂ enrichment were modified by defoliation, using Heterotheca subaxillaris, an annual plant native to south‐eastern USA. Plants were grown under near‐ambient (400 μmol mol^?1) and enriched (700 μmol mol^?1) levels of atmospheric CO₂. Defoliation consisted of the weekly removal of 25% of each new fully expanded, but not previously defoliated, leaf from either rosette or bolted plants. In addition to dry mass measurements of leaves, stems, and roots, Kjeldahl N, protein, starch and soluble sugars were analysed in these plant components to test the hypothesis that changes in C:N uptake ratio drive shifts in root:shoot ratio. Young, rapidly growing CO₂‐enriched plants conformed to the predictions of RSB, with higher root:shoot ratio than ambient‐grown plants (P < 0.02), whereas older, slower growing plants did not show a CO₂ effect on root:shoot ratio. Defoliation resulted in smaller plants, among which both root and shoot biomass were reduced, irrespective of CO₂ treatment (P < 0.03). However, H. subaxillaris plants were able to compensate for leaf area removal through flexible shoot allocation to more leaves vs. stem (P < 0.01). Increased carbon availability through CO₂ enrichment did not enhance the response to defoliation, apparently because of complete growth compensation for defoliation, even under ambient conditions. CO₂‐enriched plants had higher rates of photosynthesis (P < 0.0001), but this did not translate into increased final biomass accumulation. On the other hand, earlier and more abundant yield of flower biomass was an important consequence of growth under CO₂ enrichment.     

Plant and microbial N acquisition under elevated atmospheric CO2 in two mesocosm experiments with annual grasses

The impact of elevated CO₂ on terrestrial ecosystem C balance, both in sign or magnitude, is not clear because the resulting alterations in C input, plant nutrient demand and water use efficiency often have contrasting impacts on microbial decomposition processes. One major source of uncertainty stems from the impact of elevated CO₂ on N availability to plants and microbes. We examined the effects of atmospheric CO₂ enrichment (ambient+370 μmol mol^?1) on plant and microbial N acquisition in two different mesocosm experiments, using model plant species of annual grasses of Avena barbata and A. fatua, respectively. The A. barbata experiment was conducted in a N‐poor sandy loam and the A. fatua experiment was on a N‐rich clayey loam. Plant–microbial N partitioning was examined through determining the distribution of a ¹⁵N tracer. In the A. barbata experiment, ¹⁵N tracer was introduced to a field labeling experiment in the previous year so that ¹⁵N predominantly existed in nonextractable soil pools. In the A. fatua experiment, ¹⁵N was introduced in a mineral solution [(¹⁵NH₄)₂SO₄ solution] during the growing season of A. fatua. Results of both N budget and ¹⁵N tracer analyses indicated that elevated CO₂ increased plant N acquisition from the soil. In the A. barbata experiment, elevated CO₂ increased plant biomass N by ca. 10% but there was no corresponding decrease in soil extractable N, suggesting that plants might have obtained N from the nonextractable organic N pool because of enhanced microbial activity. In the A. fatua experiment, however, the CO₂‐led increase in plant biomass N was statistically equal to the reduction in soil extractable N. Although atmospheric CO₂ enrichment enhanced microbial biomass C under A. barbata or microbial activity (respiration) under A. fatua, it had no significant effect on microbial biomass N in either experiment. Elevated CO₂ increased the colonization of A. fatua roots by arbuscular mycorrhizal fungi, which coincided with the enhancement of plant competitiveness for soluble soil N. Together, these results suggest that elevated CO₂ may tighten N cycling through facilitating plant N acquisition. However, it is unknown to what degree results from these short‐term microcosm experiments can be extrapolated to field conditions. Long‐term studies in less‐disturbed soils are needed to determine whether CO₂‐enhancement of plant N acquisition can significantly relieve N limitation over plant growth in an elevated CO₂ environment.     

Above- and below-ground responses of C3–C4 species mixtures to elevated CO2 and soil water availability

We evaluated the influences of CO₂[Control, ～ 370 µ mol mol ^{? 1}; 200 µ mol mol ^{? 1} above ambient applied by free‐air CO₂ enrichment (FACE)] and soil water (Wet, Dry) on above‐ and below‐ground responses of C₃ (cotton, Gossypium hirsutum) and C₄ (sorghum, Sorghum bicolor) plants in monocultures and two density mixtures. In monocultures, CO₂ enrichment increased height, leaf area, above‐ground biomass and reproductive output of cotton, but not sorghum, and was independent of soil water treatment. In mixtures, cotton, but not sorghum, above‐ground biomass and height were generally reduced compared to monocultures, across both CO₂ and soil water treatments. Density did not affect individual plant responses of either cotton or sorghum across the other treatments. Total (cotton + sorghum) leaf area and above‐ground biomass in low‐density mixtures were similar between CO₂ treatments, but increased by 17–21% with FACE in high‐density mixtures, due to a 121% enhancement of cotton leaf area and a 276% increase in biomass under the FACE treatment. Total root biomass in the upper 1.2 m of the soil was not influenced by CO₂ or by soil water in monoculture or mixtures; however, under dry conditions we observed significantly more roots at lower soil depths ( > 45 cm). Sorghum roots comprised 81–85% of the total roots in the low‐density mixture and 58–73% in the high‐density mixture. CO₂‐enrichment partly offset negative effects of interspecific competition on cotton in both low‐ and high‐density mixtures by increasing above‐ground biomass, with a greater relative increase in the high‐density mixture. As a consequence, CO₂‐enrichment increased total above‐ground yield of the mixture at high density. Individual plant responses to CO₂ enrichment in global change models that evaluate mixed plant communities should be adjusted to incorporate feedbacks for interspecific competition. Future field studies in natural ecosystems should address the role that a CO₂‐mediated increase in C₃ growth may have on subsequent vegetation change.     

Ozone and density affect the response of biomass and seed yield to elevated CO2 in rice

Tropospheric O₃ reduces growth and yield of many crop species, whereas CO₂ ameliorates the negative effects of O₃. Thus, in a combined elevated CO₂ and O₃ atmosphere, seed yield is at least restored to that of charcoal‐filtered (CF) air at ambient CO₂. The CO₂‐induced yield increase in CF air is highly variable, suggesting other potential resource limitations. To understand such variability in response, we tested that (1) competition for resources precludes some of the CO₂ enhancement on biomass and yield; and (2) O₃ reduces competition in elevated CO₂. We grew rice (Oryza sativa L.) at five densities in CF and O₃‐fumigated (+O3) air at ambient (A) and elevated [CO₂] (+CO2) in 1997 and 1998. O₃ reduced biomass by 25% and seed yield by 13–20% in A, but had little effect in +CO2. A competition model of biomass and yield response to density based on resource availability without competition showed that fewer resources were used for biomass in +O3 than in CF (average 53% vs. 70%) in A, while in +CO2 85% of resources were used for biomass regardless of O₃ suggesting greater depletion of resources. The enhanced biomass response to CO₂ with O₃ is consistent with a 22% greater CO₂ enhancement ratio [mass in +CO2 air/mass in A air; enhancement ratio (ER)] in +O3 than in CF air. For seed yield, few resources were used (average 17% and 25% for CF in 1997 and 1998, respectively), and ER was 13% greater in +O3. With competition the rate of change of individual plant biomass to density was not affected by +CO2 in CF air in 1997 but was increased 19% with more nutrients in 1998, indicating resource limitations with +CO2. The rate of change of individual plant yield to density was reduced with CO₂ in 1997 and unchanged in 1998 showing a different response to resource limitation for reproductive biomass. The resource use in +O3‐A suggested that increased density and soil fertility might compensate for pollutant damage. Although ambient [O₃] can modulate the response to elevated CO₂, resource limitation precludes the CO₂ fertilization impact and both factors need consideration for better management and forecasts of future productivity.     

Effects of elevated atmospheric CO2 on net ecosystem CO2 exchange of a scrub–oak ecosystem

We report the results of a 2‐year study of effects of the elevated (current ambient plus 350 μmol CO₂ mol^?1) atmospheric CO₂ concentration (C_a) on net ecosystem CO₂ exchange (NEE) of a scrub–oak ecosystem. The measurements were made in open‐top chambers (OTCs) modified to function as open gas‐exchange systems. The OTCs enclosed samples of the ecosystem (ca. 10 m² surface area) that had regenerated after a fire, 5 years before, in either current ambient or elevated C_a. Throughout the study, elevated C_a increased maximum NEE (NEE_max) and the apparent quantum yield of the NEE (φ_NEE) during the photoperiod. The magnitude of the stimulation of NEE_max, expressed per unit ground area, was seasonal, rising from 50% in the winter to 180% in the summer. The key to this stimulation was effects of elevated C_a, and their interaction with the seasonal changes in the environment, on ecosystem leaf area index, photosynthesis and respiration. The separation of these factors was difficult. When expressed per unit leaf area the stimulation of the NEE_max ranged from 7% to 60%, with the increase being dependent on increasing soil water content (W_soil). At night, the CO₂ effluxes from the ecosystem (NEE_night) were on an average 39% higher in elevated C_a. However, the increase varied between 6% and 64%, and had no clear seasonality. The partitioning of NEE_night into its belowground (R_below) and aboveground (R_above) components was carried out in the winter only. A 35% and 27% stimulation of NEE_night in December 1999 and 2000, respectively, was largely due to a 26% and 28% stimulation of R_below in the respective periods, because R_below constituted ca. 87% of NEE_night. The 37% and 42% stimulation of R_above in December 1999 and 2000, respectively, was less than the 65% and 80% stimulation of the aboveground biomass by elevated C_a at these times. An increase in the relative amount of the aboveground biomass in woody tissue, combined with a decrease in the specific rate of stem respiration of the dominant species Quercus myrtifolia in elevated C_a, was responsible for this effect. Throughout this study, elevated C_a had a greater effect on carbon uptake than on carbon loss, in terms of both the absolute flux and relative stimulation. Consequently, for this scrub–oak ecosystem carbon sequestration was greater in the elevated C_a during this 2‐year study period.     

Growth and phenology of mature temperate forest trees in elevated CO2

Are mature forest trees carbon limited at current CO₂ concentrations? Will ‘mid‐life’, 35 m tall deciduous trees grow faster in a CO₂‐enriched atmosphere? To answer these questions we exposed ca. 100‐year‐old temperate forest trees at the Swiss Canopy Crane site near Basel, Switzerland to a ca. 540 ppm CO₂ atmosphere using web‐FACE technology. Here, we report growth responses to elevated CO₂ for 11 tall trees (compared with 32 controls) of five species during the initial four treatment years. Tested across all trees, there was no CO₂ effect on stem basal area (BA) increment (neither when tested per year nor cumulatively for 4 years). In fact, the 4th year means were almost identical for the two groups. Stem growth data were standardized by pretreatment growth (5 years) in order to account for a priori individual differences in vigor. Although this experiment was not designed to test species specific effects, one species, the common European beech, Fagus sylvatica, showed a significant growth enhancement in the first year, which reoccurred during a centennial drought in the third year. None of the other dominant species (Quercus petraea, Carpinus betulus) showed a growth response to CO₂ in any of the 4 years or for all years together. The inclusion or exclusion of single individuals of Prunus avium and Tilia platyphyllos did not change the picture. In elevated CO₂, lateral branching in terminal shoots was higher in Fagus in 2002, when shoots developed from buds that were formed during the first season of CO₂ enrichment (2001), but there was no effect in later years and no change in lateral branching in any of the other species. In Quercus, there was a steady stimulation of leading shoot length in high‐CO₂ trees. Phenological variables (bud break, leaf fall, leaf duration) were highly species specific and were not affected by elevated CO₂ in any consistent way. Our 4‐year data set reflects a very dynamic and species‐specific response of tree growth to a step change in CO₂ supply. Stem growth after 4 years of exposure does not support the notion that mature forest trees will accrete wood biomass at faster rates in a future CO₂‐enriched atmosphere.     

Seed production and seed quality in a calcareous grassland in elevated CO2

In diverse plant communities the relative contribution of species to community biomass may change considerably in response to elevated CO₂. Along with species‐specific biomass responses, reproduction is likely to change as well with increasing CO₂ and might further accelerate shifts in species composition. Here, we ask if, after 5 years of CO₂ exposure, seed production and seed quality in natural nutrient‐poor calcareous grassland are affected by elevated CO₂ (650 μ L L^?1 vs 360 μ L L^?1) and how this might affect long‐term community dynamics. The effect of elevated CO₂ on the number of flowering shoots (+ 24%, P < 0.01) and seeds (+ 29%, P = 0.06) at the community level was similar to above ground biomass responses in this year, suggesting that the overall allocation to sexual reproduction remained unchanged. Compared among functional groups of species we found a 42% increase in seed number (P < 0.01) of graminoids, a 33% increase (P = 0.07) in forbs, and no significant change in legumes (? 38%, n.s.) under elevated CO₂. Large responses particularly of two graminoid species and smaller responses of many forb species summed up to the significant or marginally significant increase in seed number of graminoids and forbs, respectively. In several species the increase in seed number resulted both from an increase in flowering shoots and an increase in inflorescence size. In most species, seeds tended to be heavier (+ 12%, P < 0.01), and N‐concentration of seeds was significantly reduced in eight out of 13 species. The fraction of germinating seeds did not differ between seeds produced in ambient and elevated CO₂, but time to germination was significantly shortened in two species and prolonged in one species when seeds had been produced in elevated CO₂. Results suggest that species specific increases in seed number and changes in seed quality will exert substantial cumulative effects on community composition in the long run.     

Growth in elevated CO2 protects photosynthesis against high-temperature damage

We present evidence that plant growth at elevated atmospheric CO₂ increases the high‐temperature tolerance of photosynthesis in a wide variety of plant species under both greenhouse and field conditions. We grew plants at ambient CO₂ (~ 360 μ mol mol ^{? 1}) and elevated CO₂ (550–1000 μ mol mol ^{? 1}) in three separate growth facilities, including the Nevada Desert Free‐Air Carbon Dioxide Enrichment (FACE) facility. Excised leaves from both the ambient and elevated CO₂ treatments were exposed to temperatures ranging from 28 to 48 °C. In more than half the species examined (4 of 7, 3 of 5, and 3 of 5 species in the three facilities), leaves from elevated CO₂‐grown plants maintained PSII efficiency (F_v/F_m) to significantly higher temperatures than ambient‐grown leaves. This enhanced PSII thermotolerance was found in both woody and herbaceous species and in both monocots and dicots. Detailed experiments conducted with Cucumis sativus showed that the greater F_v/F_m in elevated versus ambient CO₂‐grown leaves following heat stress was due to both a higher F_m and a lower F_o, and that F_v/F_m differences between elevated and ambient CO₂‐grown leaves persisted for at least 20 h following heat shock. Cucumis sativus leaves from elevated CO₂‐grown plants had a critical temperature for the rapid rise in F_o that averaged 2·9 °C higher than leaves from ambient CO₂‐grown plants, and maintained a higher maximal rate of net CO₂ assimilation following heat shock. Given that photosynthesis is considered to be the physiological process most sensitive to high‐temperature damage and that rising atmospheric CO₂ content will drive temperature increases in many already stressful environments, this CO₂‐induced increase in plant high‐temperature tolerance may have a substantial impact on both the productivity and distribution of many plant species in the 21st century.     

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

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

Partitioning net ecosystem carbon exchange with isotopic fluxes of CO2

Because biological and physical processes alter the stable isotopic composition of atmospheric CO₂, variations in isotopic content can be used to investigate those processes. Isotopic flux measurements of ¹³CO₂ above terrestrial ecosystems can potentially be used to separate net ecosystem CO₂ exchange (NEE) into its component fluxes, net photosynthetic assimilation (F_A) and ecosystem respiration (F_R). In this paper theory is developed to partition measured NEE into F_A and F_R, using measurements of fluxes of CO₂ and ¹³CO₂, and isotopic composition of respired CO₂ and forest air. The theory is then applied to fluxes measured (or estimated, for ¹³CO₂) in a temperate deciduous forest in eastern Tennessee (Walker Branch Watershed). It appears that there is indeed enough additional information in ¹³CO₂ fluxes to partition NEE into its photosynthetic and respiratory components. Diurnal patterns in F_A and F_R were obtained, which are consistent in magnitude and shape with patterns obtained from NEE measurements and an exponential regression between night‐time NEE and temperature (a standard technique which provides alternate estimates of F_R and F_A). The light response curve for photosynthesis (F_A vs. PAR) was weakly nonlinear, indicating potential for saturation at high light intensities. Assimilation‐weighted discrimination against ¹³CO₂ for this forest during July 1999 was 16.8–17.1‰, depending on canopy conductance. The greatest uncertainties in this approach lie in the evaluation of canopy conductance and its effect on whole‐canopy photosynthetic discrimination, and thus the indirect methods used to estimate isotopic fluxes. Direct eddy covariance measurements of ¹³CO₂ flux are needed to assess the validity of the assumptions used and provide defensible isotope‐based estimates of the component fluxes of net ecosystem exchange.