Elevated [CO2] alleviates the impacts of water deficit on xylem anatomy and hydraulic properties of maize stems

Plants can modify xylem anatomy and hydraulic properties to adjust to water status. Elevated [CO₂] can increase plant water potential via reduced stomatal conductance and water loss. This raises the question of whether elevated [CO₂], which thus improves plant water status, will reduce the impacts of soil water deficit on xylem anatomy and hydraulic properties of plants. To analyse the impacts of water and [CO₂] on maize stem xylem anatomy and hydraulic properties, we exposed potted maize plants to varying [CO₂] levels (400, 700, 900, and 1,200 ppm) and water levels (full irrigation and deficit irrigation). Results showed that at current [CO₂], vessel diameter, vessel roundness, stem cross-section area, specific hydraulic conductivity, and vulnerability to embolism decreased under deficit irrigation; yet, these impacts of deficit irrigation were reduced at elevated [CO₂]. Across all treatments, midday stem water potential was tightly correlated with xylem traits and displayed similar responses. A distinct trade-off between efficiency and safety in stem xylem water transportation in response to water deficit was observed at current [CO₂] but not observed at elevated [CO₂]. The results of this study enhance our knowledge of plant hydraulic acclimation under future climate environments and provide insights into trade-offs in xylem structure and function.     

Functional ecology of shrub seedlings after a natural recruitment event at the Nevada Desert FACE Facility

Seedling recruitment is an important determinant of community structure in desert ecosystems. Positive photosynthetic growth and water balance responses to increasing atmospheric carbon dioxide (CO₂) concentrations ([CO₂]) are predicted to be substantial in desert plants, suggesting that recruitment could be stimulated. However, to date no studies have addressed the response of perennial plant recruitment in natural populations of desert shrubs exposed to elevated [CO₂]. In April 1997, we employed Free‐Air Carbon Dioxide Enrichment (FACE) in order to increase atmospheric [CO₂] in an undisturbed Mojave Desert ecosystem from ambient (～～ 370 µmol mol^?1) to elevated CO₂ (～～ 550 µmol mol^?1). From 1997 to 2001 we seasonally examined survival, growth, gas exchange and water potential responses of Larrea tridentata and Ambrosia dumosa seedlings that germinated in Fall, 1997. Recruitment densities were not influenced by [CO₂] in either species, although a two‐fold higher adult Ambrosia density under elevated [CO₂] resulted in two‐fold higher seedling density (0.87 vs 0.40 seedlings m^?2). Mortality was greatest for both species during the first summer (1998), despite above‐average rainfall during the previous Winter–Spring. A significant [CO₂] × time interaction revealed that early survival was greater under elevated CO₂, whereas a significant species time interaction revealed that overall survival was greater for Ambrosia (28%) than for Larrea (15%), regardless of [CO₂]. Microsite (understorey or interspace) alone had no significant influence on survival. Significant species, microsite and species × microsite effects on growth (seedling height, stem diameter and canopy size) were found, but elevated CO₂ had minimal impact on these parameters. Photosynthetic rates (A_sat) for both species were higher at elevated [CO₂] during certain seasons, but not consistently so. These results suggest that increased atmospheric [CO₂] may enhance carbon (C) assimilation and survival of aridland perennial shrubs during favourable growing conditions, but that it may not counteract the effects of prolonged drought on mortality.     

Seasonal patterns of tissue water relations in three Mediterranean shrubs co‐occurring at a natural CO2 spring

Seasonal changes in tissue water relations of Erica arborea L., Myrtus communis L. and Juniperus communis L., grown in a Mediterranean environment, were analysed under field conditions over a 12 month period by comparing plants grown in the proximity of a natural CO₂ spring (about 700 μ mol mol ^{? 1} atmospheric CO₂ concentration, [CO₂]) with plants in ambient conditions. Tissue water relations varied in response to changes in water availability, but the seasonal course of tissue water relations parameters was also related to ontogeny. Tissue water relations of these co‐occurring shrubs were not alike. Osmotic potentials and saturated mass/dry mass ratio were lowest during peak drought stress periods. Diurnal changes in osmotic potential at the point of turgor loss were least early in the season, maximal in mid‐season, and decreased again in autumn. Turgor potentials decreased as drought progressed and were highest in late fall and mid‐winter. Symplastic water fraction was highest in mid‐spring for E. arborea and M. communis and decreased during the summer, while the opposite was observed for J. communis. Common to all species, under elevated [CO₂], was an increase of turgor pressure, particularly during the summer months. Other parameters showed species‐specific responses to long‐term elevated [CO₂]. In particular, exposure to elevated [CO₂] increased osmotic potentials in E. arborea under drought, while the opposite was the case for J. communis. Site differences in predawn to midday shifts were not strong in any of the species. Differences in tissue water relations suggest that the coexistence of these shrubs in the same environment with similar water availability are partially based on differential water relations strategies and water use patterns. Regardless of the mechanisms, growth of these shrubs in elevated [CO₂] may be either less, similarly or more affected by drought stress than plants in ambient [CO₂] depending on the species and season.     

Interactive direct and plant‐mediated effects of elevated atmospheric [CO2] and temperature on a eucalypt‐feeding insect herbivore

Understanding the direct and indirect effects of elevated [CO₂] and temperature on insect herbivores and how these factors interact are essential to predict ecosystem‐level responses to climate change scenarios. In three concurrent glasshouse experiments, we measured both the individual and interactive effects of elevated [CO₂] and temperature on foliar quality. We also assessed the interactions between their direct and plant‐mediated effects on the development of an insect herbivore of eucalypts. Eucalyptus tereticornis saplings were grown at ambient or elevated [CO₂] (400 and 650 μmol mol^?1 respectively) and ambient or elevated ( + 4 °C) temperature for 10 months. Doratifera quadriguttata (Lepidoptera: Limacodidae) larvae were feeding directly on these trees, on their excised leaves in a separate glasshouse, or on excised field‐grown leaves within the temperature and [CO₂] controlled glasshouse. To allow insect gender to be determined and to ensure that any sex‐specific developmental differences could be distinguished from treatment effects, insect development time and consumption were measured from egg hatch to pupation. No direct [CO₂] effects on insects were observed. Elevated temperature accelerated larval development, but did not affect leaf consumption. Elevated [CO₂] and temperature independently reduced foliar quality, slowing larval development and increasing consumption. Simultaneously increasing both [CO₂] and temperature reduced these shifts in foliar quality, and negative effects on larval performance were subsequently ameliorated. Negative nutritional effects of elevated [CO₂] and temperature were also independently outweighed by the direct positive effect of elevated temperature on larvae. Rising [CO₂] and temperature are thus predicted to have interactive effects on foliar quality that affect eucalypt‐feeding insects. However, the ecological consequences of these interactions will depend on the magnitude of concurrent temperature rise and its direct effects on insect physiology and feeding behaviour.     

Elevated CO2 and limited nitrogen nutrition can restrict excitation energy dissipation in photosystem II of Japanese white birch (Betula platyphylla var. japonica) leaves

Elevated atmospheric CO₂ concentration [CO₂] and different levels of nitrogen (N) nutrition can influence the amount of excess excitation energy in photosystem (PS) II and related photosynthetic properties. The interactive effect of two [CO₂] levels (ambient: 360 µM M⁻¹ and elevated: 720 µM M⁻¹) and two N levels (high: 700 mg N plant⁻¹ and low: 100 mg N plant⁻¹) on these properties was examined in seedlings of Japanese white birch (Betula platyphylla var. japonica) using simultaneous measurements of gas exchange and chlorophyll fluorescence. Photosynthetic acclimation to elevated [CO₂], as indicated by a decline in carboxylation efficiency (CE), was observed in plants grown at elevated [CO₂] especially under low N. Elevated [CO₂] resulted in a decrease in area-based leaf N content (N_area) irrespective of N treatment. The adverse effect of elevated [CO₂] and low N on CE may have been exacerbated by a greater accumulation of leaf sugar and starch contents in these plants leading to a lower electron transport rate (ETR). While these plants also showed higher non-photochemical quenching (Nq_P) that could offset the reduction in energy dissipation through ETR to some extent, they still have a higher risk of photoinhibition from excessive excitation energy in PSII as indicated by a decrease in photochemical quenching (q_P). However, chronic photoinhibition was not observed in plant grown at elevated [CO₂] and low N because they showed no difference in F_v/F_m (the maximum photochemical efficiency of PSII) from those grown at ambient [CO₂] and low N after an overnight dark adaptation. High levels of Nq_P in plants grown at elevated [CO₂] and low N reflect a near saturation of thermal energy dissipation. This impaired capacity of photoprotection would render these plants more vulnerable to photoinhibition in the event of additional environmental stresses such as drought, low or high temperature.     

Elevated CO2 increases water use efficiency by sustaining photosynthesis of water-limited maize and sorghum

Maize and grain sorghum seeds were sown in pots and grown for 39 days in sunlit controlled-environment chambers at 360 (ambient) and 720 (double-ambient, elevated) μmol mol⁻¹ carbon dioxide concentrations [CO₂]. Canopy net photosynthesis (PS) and evapotranspiration (TR) was measured throughout and summarized daily from 08:00 to 17:00 h Eastern Standard Time. Irrigation was withheld from matched pairs of treatments starting on 26 days after sowing (DAS). By 35 DAS, cumulative PS of drought-stress maize, compared to well-watered plants, was 41% lower under ambient [CO₂] but only 13% lower under elevated [CO₂]. In contrast, by 35 DAS, cumulative PS of drought-stress grain sorghum, compared to well-watered plants, was only 9% lower under ambient [CO₂] and 7% lower under elevated [CO₂]. During the 27-35 DAS drought period, water use efficiency (WUE, mol CO₂ Kmol⁻¹ H₂O), was 3.99, 3.88, 5.50, and 8.65 for maize and 3.75, 4.43, 5.26, and 9.94 for grain sorghum, for ambient-[CO₂] well-watered, ambient-[CO₂] stressed, elevated-[CO₂] well-watered and elevated-[CO₂] stressed plants, respectively. Young plants of maize and sorghum used water more efficiently at elevated [CO₂] than at ambient [CO₂], especially under drought. Reductions in biomass by drought for young maize and grain sorghum plants were 42 and 36% at ambient [CO₂], compared to 18 and 14% at elevated [CO₂], respectively. Results of our water stress experiment demonstrated that maintenance of relatively high canopy photosynthetic rates in the face of decreased transpiration rates enhanced WUE in plants grown at elevated [CO₂]. This confirms experimental evidence and conceptual models that suggest that an increase of intercellular [CO₂] (or a sustained intercellular [CO₂]) in the face of decreased stomatal conductance results in relative increases of growth of C₄ plants. In short, drought stress in C₄ crop plants can be ameliorated at elevated [CO₂] as a result of lower stomatal conductance and sustaining intercellular [CO₂]. Furthermore, less water might be required for C₄ crops in future higher CO₂ atmospheres, assuming weather and climate similar to present conditions.     

The effects of elevated atmospheric CO2 on the amount and depth distribution of plant water uptake in a California annual grassland

Soil moisture profiles can affect species composition and ecosystem processes, but the effects of increased concentrations of atmospheric carbon dioxide ([CO₂]) on the vertical distribution of plant water uptake have not been studied. Because plant species composition affects soil moisture profiles, and is likely to shift under elevated [CO₂], it is also important to test whether the indirect effects of [CO₂] on soil water content may depend on species composition. We examined the effects of elevated [CO₂] and species composition on soil moisture profiles in an annual grassland of California. We grew monocultures and a mixture of Avena barbata and Hemizonia congesta– the dominant species of two phenological groups – in microcosms exposed to ambient (～370 μmol mol^?1) and elevated (～700 μmol mol^?1) [CO₂]. Both species increased intrinsic and yield‐based water use efficiency under elevated [CO₂], but soil moisture increased only in communities with A. barbata, the dominant early‐season annual grass. In A. barbata monocultures, the [CO₂] treatment did not affect the depth distribution of soil water loss. In contrast to communities with A. barbata, monocultures of H. congesta, a late‐season annual forb, did not conserve water under elevated [CO₂], reflecting the increased growth of these plants. In late spring, elevated [CO₂] also increased the efficiency of deep roots in H. congesta monocultures. Under ambient [CO₂], roots below 60 cm accounted for 22% of total root biomass and were associated with 9% of total water loss, whereas in elevated [CO₂], 16% of total belowground biomass was associated with 34% of total water loss. Both soil moisture and isotope data showed that H. congesta monocultures grown under elevated [CO₂] began extracting water from deep soils 2 weeks earlier than plants in ambient [CO₂].     

Effect of elevated [CO2] on stem wood properties of mature Norway spruce grown at different soil nutrient availability

The objective of the present study was to investigate the interactive effects of elevated [CO₂] and soil nutrient availability on secondary xylem structure and chemical composition of 41‐year‐old Norway spruce (Picea abies (L.) Karst.) trees. The nonfertilized and irrigated‐fertilized trees were, for 3 years, continuously exposed to elevated [CO₂] in whole‐tree chambers. Elevated [CO₂] decreased concentrations of soluble sugars, acid‐soluble lignin and nitrogen in stem wood, but the effects were not consistent between sampling height and/or fertilization. The effect of 2*ambient [CO₂] on wood structure depended on the exposure year and/or fertilization. Radial lumen diameter decreased and annual ring width increased in the second year of exposure (1999) in elevated [CO₂]. In the latter, the CO₂ effect was significant only in the nonfertilized trees. Stem wood chemistry and structure were significantly affected by fertilization. Fertilization increased the concentrations of nitrogen and gravimetric lignin, annual ring width, and radial lumen diameter. Fertilization decreased C/N ratio, mean ring density, earlywood density, latewood density, cell wall thickness, cell wall index, and latewood percentage. We conclude that elevated [CO₂] had only minor effects on wood properties while fertilization had more marked effects and thus may affect ecosystem processes and suitability of wood for different end‐use purposes.     

Increased night temperature reduces the stimulatory effect of elevated carbon dioxide concentration on methane emission from rice paddy soil

To determine how elevated night temperature interacts with carbon dioxide concentration ([CO₂]) to affect methane (CH₄) emission from rice paddy soil, we conducted a pot experiment using four controlled‐environment chambers and imposed a combination of two [CO₂] levels (ambient: 380 ppm; elevated: 680 ppm) and two night temperatures (22 and 32 °C). The day temperature was maintained at 32 °C. Rice (cv. IR72) plants were grown outside until the early‐reproductive growth stage and then transferred to the chambers. After onset of the treatment, day and night CH₄ fluxes were measured every week. The CH₄ fluxes changed significantly with the growth stage, with the largest fluxes occurring around the heading stage in all treatments. The total CH₄ emission during the treatment period was significantly increased by both elevated [CO₂] (P=0.03) and elevated night temperature (P<0.01). Elevated [CO₂] increased CH₄ emission by 3.5% and 32.2% under high and low night temperature conditions, respectively. Elevated [CO₂] increased the net dry weight of rice plants by 12.7% and 38.4% under high and low night temperature conditions, respectively. These results imply that increasing night temperature reduces the stimulatory effect of elevated [CO₂] on both CH₄ emission and rice growth. The CH₄ emission during the day was larger than at night even under the high‐night‐temperature treatment (i.e. a constant temperature all day). This difference became larger after the heading stage. We observed significant correlations between the night respiration and daily CH₄ flux (P<0.01). These results suggest that net plant photosynthesis contributes greatly to CH₄ emission and that increasing night temperature reduces the stimulatory effect of elevated [CO₂] on CH₄ emission from rice paddy soil.     

Seasonal patterns of photosynthetic response and acclimation to elevated carbon dioxide in field-grown strawberry

Strawberry (Fragaria × ananassa) plants were grown in field plots at the current ambient [CO₂], and at ambient + 300 and ambient + 600 μmol mol⁻¹ [CO₂]. Approximately weekly measurements were made of single leaf gas exchange of upper canopy leaves from early spring through fall of two years, in order to determine the temperature dependence of the stimulation of photosynthesis by elevated [CO₂], whether growth at elevated [CO₂] resulted in acclimation of photosynthesis, and whether any photosynthetic acclimation was reduced when fruiting created additional demand for the products of photosynthesis. Stimulation of photosynthetic CO₂ assimilation by short-term increases in [CO₂] increased strongly with measurement temperature. The stimulation exceeded that predicted from the kinetic characteristics of ribulose-1,5-bisphosphate carboxylase at all temperatures. Acclimation of photosynthesis to growth at elevated [CO₂] was evident from early spring through summer, including the fruiting period in early summer, with lower rates under standard measurement conditions in plants grown at elevated [CO₂]. The degree of acclimation increased with growth [CO₂]. However, there were no significant differences between [CO₂] treatments in total nitrogen per leaf area, and photosynthetic acclimation was reversed one day after switching the [CO₂] treatments. Tests showed that acclimation did not result from a limitation of photosynthesis by triose phosphate utilization rate at elevated [CO₂]. Photosynthetic acclimation was not evident during dry periods in midsummer, when the elevated [CO₂] treatments conserved soil water and photosynthesis declined more at ambient than at elevated [CO₂]. Acclimation was also not evident during the fall, when plants were vegetative, despite wet conditions and continued higher leaf starch content at elevated [CO₂]. Stomatal conductance responded little to short-term changes in [CO₂] except during drought, and changed in parallel with photosynthetic acclimation through the seasons in response to the long-term [CO₂] treatments. The data do not support the hypothesis that source-sink balance controls the seasonal occurrence of photosynthetic acclimation to elevated [CO₂] in this species. This revised version was published online in June 2006 with corrections to the Cover Date.     

Nitrogen assimilation and transpiration: key processes conditioning responsiveness of wheat to elevated [CO2] and temperature

Although climate scenarios have predicted an increase in [CO₂] and temperature conditions, to date few experiments have focused on the interaction of [CO₂] and temperature effects in wheat development. Recent evidence suggests that photosynthetic acclimation is linked to the photorespiration and N assimilation inhibition of plants exposed to elevated CO₂. The main goal of this study was to analyze the effect of interacting [CO₂] and temperature on leaf photorespiration, C/N metabolism and N transport in wheat plants exposed to elevated [CO₂] and temperature conditions. For this purpose, wheat plants were exposed to elevated [CO₂] (400 vs 700 µmol mol^?1) and temperature (ambient vs ambient + 4°C) in CO₂ gradient greenhouses during the entire life cycle. Although at the agronomic level, elevated temperature had no effect on plant biomass, physiological analyses revealed that combined elevated [CO₂] and temperature negatively affected photosynthetic performance. The limited energy levels resulting from the reduced respiratory and photorespiration rates of such plants were apparently inadequate to sustain nitrate reductase activity. Inhibited N assimilation was associated with a strong reduction in amino acid content, conditioned leaf soluble protein content and constrained leaf N status. Therefore, the plant response to elevated [CO₂] and elevated temperature resulted in photosynthetic acclimation. The reduction in transpiration rates induced limitations in nutrient transport in leaves of plants exposed to elevated [CO₂] and temperature, led to mineral depletion and therefore contributed to the inhibition of photosynthetic activity.     

Soybean nodulation and N2 fixation response to drought under carbon dioxide enrichment

The combined effects of carbon dioxide (CO₂) enrichment and water deficits on nodulation and N₂ fixation were analysed in soybean [Glycine max (L.) Merr.]. Two short-term experiments were conducted in greenhouses with plants subjected to soil drying, while exposed to CO₂ atmospheres of either 360 or 700 μmol CO₂ mol^–1. Under drought-stressed conditions, elevated [CO₂] resulted in a delay in the decrease in N₂ fixation rates associated with drying of the soil used in these experiments. The elevated [CO₂] also allowed the plants under drought to sustain significant increases in nodule number and mass relative to those under ambient [CO₂]. The total non-structural carbohydrate (TNC) concentration was lower in the shoots of the plants exposed to drought; however, plants under elevated CO₂ had much higher TNC levels than those under ambient CO₂. For both [CO₂] treatments, drought stress induced a substantial accumulation of TNC in the nodules that paralleled N₂ fixation decline, which indicates that nodule activity under drought may not be carbon limited. Under drought stress, ureide concentration increased in all plant tissues. However, exposure to elevated [CO₂] resulted in substantially less drought-induced ureide accumulation in leaf and petiole tissues. A strong negative correlation was found between ureide accumulation and TNC levels in the leaves. This relationship, together with the large effect of elevated [CO₂] on the decrease of ureide accumulation in the leaves, indicated the importance of ureide breakdown in the response of N₂ fixation to drought and of feedback inhibition by ureides on nodule activity. It is concluded that an important effect of CO₂ enrichment on soybean under drought conditions is an enhancement of photoassimilation, an increased partitioning of carbon to nodules and a decrease of leaf ureide levels, which is associated with sustained nodule growth and N₂ rates under soil water deficits. We suggest that future [CO₂] increases are likely to benefit soybean production by increasing the drought tolerance of N₂ fixation.     

Seasonal acclimation of leaf respiration in Eucalyptus saligna trees: impacts of elevated atmospheric CO2 and summer drought

Understanding the impacts of atmospheric [CO₂] and drought on leaf respiration (R) and its response to changes in temperature is critical to improve predictions of plant carbon‐exchange with the atmosphere, especially at higher temperatures. We quantified the effects of [CO₂]‐enrichment (+240 ppm) on seasonal shifts in the diel temperature response of R during a moderate summer drought in Eucalyptus saligna growing in whole‐tree chambers in SE Australia. Seasonal temperature acclimation of R was marked, as illustrated by: (1) a downward shift in daily temperature response curves of R in summer (relative to spring); (2)≈60% lower R measured at 20^oC (R₂₀) in summer compared with spring; and (3) homeostasis over 12 months of R measured at prevailing nighttime temperatures. R₂₀, measured during the day, was on average 30–40% higher under elevated [CO₂] compared with ambient [CO₂] across both watered and droughted trees. Drought reduced R₂₀ by≈30% in both [CO₂] treatments resulting in additive treatment effects. Although [CO₂] had no effect on seasonal acclimation, summer drought exacerbated the seasonal downward shift in temperature response curves of R. Overall, these results highlight the importance of seasonal acclimation of leaf R in trees grown under ambient‐ and elevated [CO₂] as well as under moderate drought. Hence, respiration rates may be overestimated if seasonal changes in temperature and drought are not considered when predicting future rates of forest net CO₂ exchange.     

Stomatal responses of Douglas-fir seedlings to elevated carbon dioxide and temperature during the third and fourth years of exposure

Two major components of climate change, increasing atmospheric [CO₂] and increasing temperature, may substantially alter the effects of water availability to plants through effects on the rate of water loss from leaves. We examined the interactive effects of elevated [CO₂] and temperature on seasonal patterns of stomatal conductance (g_s), transpiration (E) and instantaneous transpiration efficiency (ITE) in Douglas‐fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings. Seedlings were grown in sunlit chambers at either ambient CO₂ (AC) or ambient + 180 µmol mol^?1 CO₂ (EC), and at ambient temperature (AT) or ambient + 3·5 °C (ET) in a full‐factorial design. Needle gas exchange at the target growth conditions was measured approximately monthly over 21 months. Across the study period and across temperature treatments, growth in elevated [CO₂] decreased E by an average of 12% and increased ITE by an average of 46%. The absolute reduction of E associated with elevated [CO₂] significantly increased with seasonal increases in the needle‐to‐air vapour pressure deficit (D). Across CO₂ treatments, growth in elevated temperature increased E an average of 37%, and did not affect ITE. Combined, growth in elevated [CO₂] and elevated temperature increased E an average of 19% compared with the ACAT treatment. The CO₂ supply and growth temperature did not significantly affect stomatal sensitivity to D or the relationship between g_s and net photosynthetic rates. This study suggests that elevated [CO₂] may not completely ameliorate the effect of elevated temperature on E, and that climate change may substantially alter needle‐level water loss and water use efficiency of Douglas‐fir seedlings.     

Elevated CO2 alleviates the impact of drought on barley improving water status by lowering stomatal conductance and delaying its effects on photosynthesis

We analysed the impact of elevated CO₂ on water relations, water use efficiency and photosynthetic gas exchange in barley (Hordeum vulgare L.) under wet and drying soil conditions. Soil moisture was less depleted under elevated compared to ambient [CO₂]. Elevated CO₂ had no significant effect on the water relations of irrigated plants, except on whole plant hydraulic conductance, which was markedly decreased at elevated compared to ambient CO₂ concentrations. The values of relative water content, water potential and osmotic potential were higher under elevated CO₂ during the entire drought period. The better water status of water-limited plants grown at elevated CO₂ was the result of stomatal control rather than of osmotic adjustment. Despite the low stomatal conductance produced by elevated CO₂, net photosynthesis was higher under elevated than ambient CO₂ concentrations. With water shortage, photosynthesis was maintained for longer at higher rates under elevated CO₂. The reduction of stomatal conductance and therefore transpiration, and the enhancement of carbon assimilation by elevated CO₂, increased instantaneous and whole plant water use efficiency in both irrigated and droughted plants. Thus, the metabolism of barley plants grown under elevated CO₂ and moderate or mild water deficit conditions is benefited by increased photosynthesis and lower transpiration. The reduction in plant water use results in a marked increase in soil water content which delays the onset and severity of water deficit.     

The influence of elevated temperature,elevated atmospheric CO2 concentration and water stress on net photosynthesis of loblolly pine (Pinus taeda L.) at northern,central and southern sites in its native range

We investigated the effect of elevated [CO₂] (700 μmol mol^?1), elevated temperature (+2 °C above ambient) and decreased soil water availability on net photosynthesis (A_net) and water relations of one‐year old potted loblolly pine (Pinus taeda L.) seedlings grown in treatment chambers with high fertility at three sites along a north‐south transect covering a large portion of the species native range. At each location (Blairsville, Athens and Tifton, GA) we constructed four treatment chambers and randomly assigned each chamber one of four treatments: ambient [CO₂] and ambient temperature, elevated [CO₂] and ambient temperature, ambient [CO₂] and elevated temperature, or elevated [CO₂] and elevated temperature. Within each chamber half of the seedlings were well watered and half received much less water (1/4 that of the well watered). Measurements of net photosynthesis (A_net), stomatal conductance (g_s), leaf water potential and leaf fluorescence were made in June and September, 2008. We observed a significant increase in A_net in response to elevated [CO₂] regardless of site or temperature treatment in June and September. An increase in air temperature of over 2 °C had no significant effect on A_net at any of the sites in June or September despite over a 6 °C difference in mean annual temperature between the sites. Decreased water availability significantly reduced A_net in all treatments at each site in June. The effects of elevated [CO₂] and temperature on g_s followed a similar trend. The temperature, [CO₂] and water treatments did not significantly affect leaf water potential or chlorophyll fluorescence. Our findings suggest that predicted increases in [CO₂] will significantly increase A_net, while predicted increases in air temperature will have little effect on A_net across the native range of loblolly pine. Potential decreases in precipitation will likely cause a significant reduction in A_net, though this may be mitigated by increased [CO₂].     

Low soil temperature inhibits the stimulatory effect of elevated [CO2] on height and biomass accumulation of white birch seedlings grown under three non‐limiting phosphorus conditions

White birch (Betula papyrifera Marsh.) seedlings were exposed to ambient or doubled ambient carbon dioxide concentration ([CO₂]), three soil temperatures (T_soil) (low, intermediate, high), and three phosphorus (P) regimes (low, medium, high) in environment‐controlled greenhouses. Height (H), root‐collar diameter (RCD), biomass, and leaf phosphorus concentration (leaf P) were determined four months after initiation of treatments. The low T_soil reduced H, RCD, shoot biomass, root biomass and total seedling biomass whereas the high‐P level and the [CO₂] elevation increased all the growth and biomass parameters. Elevated [CO₂] significantly reduced leaf P. There were significant two‐factor interactions suggesting that the effect of elevated [CO₂] on (1) H, total biomass, biomass of plant components, and leaf P was dependent on T_soil, (2) total biomass was contingent on P regime. For instance, the positive response of H and total biomass to elevated [CO₂] was limited to seedlings raised under the intermediate and high T_soil, respectively. In addition, [CO₂] elevation increased total biomass only at the high‐P regime but not at the low‐ or medium‐P level where the effect of [CO₂] was statistically insignificant. No significant main effect of treatment or interaction was observed for root to shoot biomass ratio.     

Long-term exposure to elevated [CO2] in a natural Quercus ilex L. community: net photosynthesis and photochemical efficiency of PSII at different levels of water stress

Naturally grown trees of Mediterranean evergreen oak (Quercus ilex L.), representing the climax species of the region, were enclosed in six large open-top chambers and exposed to ambient and elevated CO₂ concentrations during a 3 year period. Maximum daily net photosynthetic rates measured at the two different CO₂ concentrations were from 30 to 100% higher in elevated than in ambient [CO₂] throughout the experimental period. The increase in maximum daily photosynthesis was also accompanied by a 93% rise in the apparent quantum yield of CO₂ assimilation, measured during periods of optimum soil moisture conditions. Hence, no clear evidence of down-regulation of net photosynthetic activity was found. Interactions between atmospheric CO₂ concentration and plant water stress were studied by following the natural evolution of drought in different seasons and years. At each level of water stress, the maximum rate of carbon assimilation was higher in elevated than in ambient [CO₂] by up to 100%. Analysis of in vivo chlorophyll fluorescence parameters in normal (21%) and low (2%) oxygen concentrations provided useful insights into the functioning and stability of the photosynthetic processes. The photochemical efficiency of PSII (F_v/F_m) progressively decreased as drought conditions became more evident; this trend was accentuated under elevated [CO₂]. Thermal de-excitation processes were possibly more significant under elevated than under ambient [CO₂], in a combination of environmental stresses. This research suggests two possible conclusions: (i) a ‘positive’ interaction between elevated [CO₂] and carbon metabolism can be obtained through relief of water stress limitation in the summer months, and (ii) elevated [CO₂], under drought conditions, may also enhance the significance of slow-relaxing quenching.     

Embolism recovery strategies and nocturnal water loss across species influenced by biogeographic origin

Drought‐induced tree mortality is expected to increase in future climates with the potential for significant consequences to global carbon, water, and energy cycles. Xylem embolism can accumulate to lethal levels during drought, but species that can refill embolized xylem and recover hydraulic function may be able to avoid mortality. Yet the potential controls of embolism recovery, including cross‐biome patterns and plant traits such as nonstructural carbohydrates (NSCs), hydraulic traits, and nocturnal stomatal conductance, are unknown. We exposed eight plant species, originating from mesic (tropical and temperate) and semi‐arid environments, to drought under ambient and elevated CO₂ levels, and assessed recovery from embolism following rewatering. We found a positive association between xylem recovery and NSCs, and, surprisingly, a positive relationship between xylem recovery and nocturnal stomatal conductance. Arid‐zone species exhibited greater embolism recovery than mesic zone species. Our results indicate that nighttime stomatal conductance often assumed to be a wasteful use of water, may in fact be a key part of plant drought responses, and contribute to drought survival. Findings suggested distinct biome‐specific responses that partially depended on species climate‐of‐origin precipitation or aridity index, which allowed some species to recover from xylem embolism. These findings provide improved understanding required to predict the response of diverse plant communities to drought. Our results provide a framework for predicting future vegetation shifts in response to climate change.     

Root-zone CO2 and root-zone temperature effects on photosynthesis and nitrogen metabolism of aeroponically grown lettuce (Lactuca sativa L.) in the tropics

Effects of elevated root-zone (RZ) CO₂ concentration (RZ [CO₂]) and RZ temperature (RZT) on photosynthesis, productivity, nitrate (NO₃ ^?), total reduced nitrogen (TRN), total leaf soluble and Rubisco proteins were studied in aeroponically grown lettuce plants in a tropical greenhouse. Three weeks after transplanting, four different RZ [CO₂] concentrations (ambient, 360 ppm, and elevated concentrations of 2,000; 10,000; and 50,000 ppm) were imposed on plants at 20°C-RZT or ambient(A)-RZT (24–38°C). Elevated RZ [CO₂] resulted in significantly higher light-saturated net photosynthetic rate, but lower light-saturated stomatal conductance. Higher elevated RZ [CO₂] also protected plants from both chronic and dynamic photoinhibition (measured by chlorophyll fluorescence Fv/Fm ratio) and reduced leaf water loss. Under each RZ [CO₂], all these variables were significantly higher in 20°C-RZT plants than in A-RZT plants. All plants accumulated more biomass at elevated RZ [CO₂] than at ambient RZ [CO₂]. Greater increases of biomass in roots than in shoots were manifested by lower shoot/root ratios at elevated RZ [CO₂]. Although the total biomass was higher at 20°C-RZT, the increase in biomass under elevated RZ [CO₂] was greater at A-RZT. Shoot NO₃ ^? and TRN concentrations, total leaf soluble and Rubisco protein concentrations were higher in all elevated RZ [CO₂] plants than in plants under ambient RZ [CO₂] at both RZTs. Under each RZ [CO₂], total leaf soluble and Rubisco protein concentrations were significantly higher at 20°C-RZT than at A-RZT. Our results demonstrated that increased P _Nmax and productivity under elevated [CO₂] was partially due to the alleviation of midday water loss, both dynamic and chronic photoinhibition as well as higher turnover of Calvin cycle with higher Rubisco proteins.