Stomatal regulation in a changing climate: a field study using Free Air Temperature Increase (FATI) and Free Air CO2 Enrichment (FACE)

This study investigates effects of climate warming (+ 2.5°C ubove ambient) and elevated CO₂ concentration (600 μmol mol^?1) on the stomatal functioning and the water relations of Lolium perenne, using Free Air Temperature Increase (FATI) and Free Air CO₂ Enrichment (FACE). Compared to growth at ambient temperature, whole-season temperature increase reduced leaf stomatal conductance, but only at the top of the canopy (-14.6 and -8.8% at ambient and elevated CO₂, respectively). However, because higher canopy temperature raised the leaf-to-air vapour pressure difference, leaf transpiration rate increased (+28% at ambient and +48% at elevated CO₂) and instantaneous leaf water use efficiency, derived from short-term measurements of assimilation and transpiration rate, declined (-11% at ambient and -13% at elevated CO₂). Nevertheless, at the stand level, growth at + 2.5°C reduced transpiration due to fewer tillers per plant and a smaller leaf area per tiller. This sparser vegetation was also more closely coupled to the atmosphere and maintained a drier internal microclimate. To assess whether the stomatal behaviour observed in this experiment could be explained by prevailing concepts of stomatal functioning, three models were applied (Cowan 1977; Ball, Woodrow & Berry 1987; Leuning 1995). The latter model accounted for the highest proportion of variability in the data (58%) and was insensitive to CO₂ and temperature regime, which suggests that the principles of stomatal regulation are not affected by changes in CO₂ or climate.     

CO2 enrichment in a maturing pine forest: are CO2 exchange and water status in the canopy affected?

A _net, leaf net CO₂ assimilation
c_a, CO₂ concentration of air surrounding a leaf
c_i, leaf intercellular CO₂ concentration
Δ, ¹³C isotope discrimination
δ¹³C, relative stable carbon isotope content
?, ratio of A_net at c_a = 560μmol mol^–1 to A_net at c_a = 360 μmol mol^–1
FACE, free-air CO₂ enrichment
g_w, stomatal conductance to water vapour
Π_i, initial leaf osmotic potential
R_t, relative water content at incipient turgor loss
Ψ_l, xylem water potential of leaves
Ψ_m, soil matric potential

Elevated CO₂ is expected to reduce forest water use as a result of CO₂-induced stomatal closure, which has implications for ecosystem-scale phenomena controlled by water availability. Leaf-level CO₂ and H₂O exchange responses and plant and soil water relations were examined in a maturing loblolly pine (Pinus taeda L.) stand in a free-air CO₂ enrichment (FACE) experiment in North Carolina, USA to test if these parameters were affected by elevated CO₂. Current-year foliage in the canopy was continuously exposed to elevated CO₂ (ambient CO₂+200μmol mol^–1) in free-air during needle growth and development for up to 400 d. Photosynthesis in upper canopy foliage was stimulated by 50–60% by elevated CO₂ compared with ambient controls. This enhancement was similar in current-year, ambient-grown foliage temporarily measured at elevated CO₂ compared with long-term elevated CO₂ grown foliage. Significant photosynthetic enhancement by CO₂ was maintained over a range of conditions except during peak drought. There was no evidence of water savings in elevated CO₂ plots in FACE compared to ambient plots under drought and non-drought conditions. This was supported by evidence from three independent measures. First, stomatal conductance was not significantly different in elevated CO₂ versus ambient trees of P. taeda. Calculations of time-integrated c_i/c_a ratios from analysis of foliar δ¹³C showed that these ratios were maintained in foliage under elevated CO₂. Second, soil moisture was not significantly different between ambient and elevated CO₂ plots during drought. Third, pre-dawn and mid-day leaf water potentials were also unaffected by the seasonal CO₂ exposure, as were tissue osmotic potentials and turgor loss points. Together the results strongly support the hypothesis that maturing P. taeda trees have low stomatal responsiveness to elevated CO₂. Elevated CO₂ effects on water relations in loblolly pine-dominated forest ecosystems may be absent or small apart from those mediated by leaf area. Large photosynthetic enhancements in the upper canopy of P. taeda by elevated CO₂ indicate that this maturing forest may have a large carbon sequestration capacity with limiting water supply.     

CO2 enrichment predisposes foliage of a eucalypt to freezing injury and reduces spring growth

Seedlings of Eucalyptus pauciflora, were grown in open-top chambers fumigated with ambient and elevated [CO₂], and were divided into two populations using 10% light transmittance screens. The aim was to separate the effects of timing of light interception, temperature and [CO₂] on plant growth. The orientation of the screens exposed plants to a similar total irradiance, but incident during either cold mornings (east-facing) or warm afternoons (west-facing). Following the first autumn freezing event elevated CO₂-grown plants had 10 times more necrotic leaf area than ambient CO₂ plants. West-facing plants had significantly greater (25% more) leaf damage and lower photochemical efficiency (F_v/F_m) in comparison with east-facing plants. Following a late spring freezing event east-facing elevated CO₂ plants suffered a greater sustained loss in F_v/F_m than west-facing elevated CO₂- and ambient CO₂-grown plants. Stomatal conductance was lower under elevated CO₂ than ambient CO₂ except during late spring, with the highest leaf temperatures occurring in west-facing plants under elevated CO₂. These higher leaf temperatures apparently interfered with cold acclimation thereby enhancing frost damage and reducing the ability to take advantage of optimal growing conditions under elevated CO₂.     

Elevated CO2 affects the interactions between aphid pests and host plant flowering

1 Broad beans (Vicia faba L.) were grown at either ambient (350 μL/L) or elevated (700 μL/L) CO₂. Elevated CO₂ increased shoot weight by 14% and root weight by 24% compared to ambient, but did not affect flowering. 2 A single pea aphid (Acyrthosiphon pisum (Harris)) and its progeny decreased shoot and root weights by 20 and 24%, respectively, at ambient CO₂ after 20 days, but did not affect flower number. At elevated CO₂A. pisum decreased shoot and root weights by 27 and 34% and flower number decreased by 73%. 3 A single glasshouse and potato aphid (Aulacorthum solani (Kaltenbach)) and its progeny had no effect on the growth of bean plants after 20 days at ambient CO₂. At elevated CO₂, A. solani decreased shoot and root weights by 20 and 18%, and flower number by 60%. 4 The large reduction in flowering caused by aphids at elevated CO₂ suggests a change in resource allocation within the plants to compensate for aphid infestation. 5 Aphid density was unaffected by elevated CO₂, although there were significant effects of CO₂ on the resulting population structure of both A. pisum and A solani. We suggest that at elevated CO₂, aphids appear not to achieve their maximum reproductive potential and their populations are limited by the lower carrying capacity of their host plants.     

Will photosynthesis of maize (Zea mays) in the US Corn Belt increase in future [CO2] rich atmospheres? An analysis of diurnal courses of CO2 uptake under free‐air concentration enrichment (FACE)

The C₄ grass Zea mays (maize or corn) is the third most important food crop globally in terms of production and demand is predicted to increase 45% from 1997 to 2020. However, the effects of rising [CO₂] upon C₄ plants, and Z. mays specifically, are not sufficiently understood to allow accurate predictions of future crop production. A rainfed, field experiment utilizing free‐air concentration enrichment (FACE) technology in the primary area of global corn production (US Corn Belt) was undertaken to determine the effects of elevated [CO₂] on corn. FACE technology allows experimental treatments to be imposed upon a complete soil–plant–atmosphere continuum with none of the effects of experimental enclosures on plant microclimate. Crop performance was compared at ambient [CO₂] (354 μ mol mol^?1) and the elevated [CO₂] (549 μmol mol^?1) predicted for 2050. Previous laboratory studies suggest that under favorable growing conditions C₄ photosynthesis is not typically enhanced by elevated [CO₂]. However, stomatal conductance and transpiration are decreased, which can indirectly increase photosynthesis in dry climates. Given the deep soils and relatively high rainfall of the US Corn Belt, it was predicted that photosynthesis would not be enhanced by elevated [CO₂]. The diurnal course of gas exchange of upper canopy leaves was measured in situ across the growing season of 2002. Contrary to the prediction, growth at elevated [CO₂] significantly increased leaf photosynthetic CO₂ uptake rate (A) by up to 41%, and 10% on average. Greater A was associated with greater intercellular [CO₂], lower stomatal conductance and lower transpiration. Summer rainfall during 2002 was very close to the 50‐year average for this site, indicating that the year was not atypical or a drought year. The results call for a reassessment of the established view that C₄ photosynthesis is insensitive to elevated [CO₂] under favorable growing conditions and that the production potential of corn in the US Corn Belt will not be affected by the global rise in [CO₂].     

Elevated atmospheric CO2 alters stomatal responses to variable sunlight in a C4 grass

Native tallgrass prairie in NE Kansas was exposed to elevated (twice ambient) or ambient atmospheric CO₂ levels in open-top chambers. Within chambers or in adjacent unchambered plots, the dominant C₄ grass, Andropogon gerardii, was subjected to fluctuations in sunlight similar to that produced by clouds or within canopy shading (full sun > 1500 μmol m⁻² s⁻¹ versus 350 μmol m⁻² s⁻¹ shade) and responses in gas exchange were measured. These field experiments demonstrated that stomatal conductance in A. gerardii achieved new steady state levels more rapidly after abrupt changes in sunlight at elevated CO₂ when compared to plants at ambient CO₂. This was due primarily to the 50% reduction in stomatal conductance at elevated CO₂, but was also a result of more rapid stomatal responses. Time constants describing stomatal responses were significantly reduced (29–33%) at elevated CO₂. As a result, water loss was decreased by as much as 57% (6.5% due to more rapid stomatal responses). Concurrent increases in leaf xylem pressure potential during periods of sunlight variability provided additional evidence that more rapid stomatal responses at elevated CO₂ enhanced plant water status. CO₂-induced alterations in the kinetics of stomatal responses to variable sunlight will likely enhance direct effects of elevated CO₂ on plant water relations in all ecosystems.     

The effects on Arbutus unedo L. of long-term exposure to elevated CO2

Arbutus unedo is a sclerophyllous evergreen, characteristic of Mediterranean coastal scrub vegetation. In Italy, trees of A. unedo have been found close to natural CO₂ vents where the mean atmospheric carbon dioxide concentration is about 2200 μmol mol^?1. Comparisons were made between trees growing in elevated and ambient CO₂ concentrations to test for evidence of adaptation to long-term exposure to elevated CO₂. Leaves formed at elevated CO₂ have a lower stomatal density and stomatal index and higher specific leaf area than those formed at ambient CO₂, but there was no change in carbon to nitrogen ratios of the leaf tissue. Stomatal conductance was lower at elevated CO₂ during rapid growth in the spring. In mid-summer, under drought stress, stomatal closure of all leaves occurred and in the autumn, when stress was relieved, the conductance of leaves at both elevated and ambient CO₂ increased. In the spring, the stomatal conductance of the new flush of leaves at ambient CO₂ was higher than the leaves at elevated CO₂, increasing instantaneous water use efficiency at elevated CO₂. Chlorophyll fluorescence measurements suggested that elevated CO₂ provided some protection against photoinhibition in mid-summer. Analysis of A/C_i curves showed that there was no evidence of either upward or downward regulation of photosynthesis at elevated CO₂. It is therefore anticipated that A. unedo will have higher growth rates as the ambient CO₂ concentrations increase.     

Effects of elevated CO2, drought and temperature on the water relations and gas exchange of groundnut (Arachis hypogaea) stands grown in controlled environment glasshouses

Stands of groundnut (Arachis hypogaea L. cv. Kadiri‐3) were grown in controlled environment glasshouses at mean atmospheric CO₂ concentrations of 375 or 700 μmol mol^?1 and daily mean air temperatures of 28 or 32°C on irrigated or drying soil profiles. Leaf water (Ψ_l) and solute potential (Ψ_s), relative water content (RWC), stomatal conductance (g_l) and net photosynthesis (P_n) were measured at midday for the youngest mature leaf throughout the growing season. Elevated CO₂ and temperature had no detectable effect on the water relations of irrigated plants, but higher values of RWC, Ψ_l and Ψ_s were maintained for longer under elevated CO₂ during progressive drought. Turgor potential (Ψ_p) reached zero when Ψ_l declined to ?1.6 to ?1.8 MPa in all treatments; turgor was lost sooner when droughted plants were grown under ambient CO₂. A 4°C increase in mean air temperature had no effect on Ψ_s in droughted plants, but elicited a small increase in Ψ_l; midday g_l values were lower under elevated than under ambient CO₂, and Ψ_l and g_l declined below ?1.5 MPa and 0.25 cm s^?1, respectively, as the soil dried. Despite the low g_l values recorded for droughted plants late in the season, P_n was maintained under elevated CO₂, but declined to zero 3 weeks before final harvest under ambient CO₂. Concurrent reductions in g_l and increases in water use efficiency under elevated CO₂ prolonged photosynthetic activity during drought and increased pod yields relative to plants grown under ambient CO₂. The implications of future increases in atmospheric CO₂ for the productivity of indeterminate C₃ crops grown in rainfed subsistence agricultural systems in the semi‐arid tropics are discussed.     

The effect of elevated CO2 on diel leaf growth cycle, leaf carbohydrate content and canopy growth performance of Populus deltoides

Image sequence processing methods were applied to study the effect of elevated CO₂ on the diel leaf growth cycle for the first time in a dicot plant. Growing leaves of Populus deltoides, in stands maintained under ambient and elevated CO₂ for up to 4 years, showed a high degree of heterogeneity and pronounced diel variations of their relative growth rate (RGR) with maxima at dusk. At the beginning of the season, leaf growth did not differ between treatments. At the end of the season, final individual leaf area and total leaf biomass of the canopy was increased in elevated CO₂. Increased final leaf area at elevated CO₂ was achieved via a prolonged phase of leaf expansion activity and not via larger leaf size upon emergence. The fraction of leaves growing at 30–40% day^?1 was increased by a factor of two in the elevated CO₂ treatment. A transient minimum of leaf expansion developed during the late afternoon in leaves grown under elevated CO₂ as the growing season progressed. During this minimum, leaves grown under elevated CO₂ decreased their RGR to 50% of the ambient value. The transient growth minimum in the afternoon was correlated with a transient depletion of glucose (less than 50%) in the growing leaf in elevated CO₂, suggesting diversion of glucose to starch or other carbohydrates, making this substrate temporarily unavailable for growth. Increased leaf growth was observed at the end of the night in elevated CO₂. Net CO₂ exchange and starch concentration of growing leaves was higher in elevated CO₂. The extent to which the transient reduction in diel leaf growth might dampen the overall growth response of these trees to elevated CO₂ is discussed.     

Acclimation of photosynthesis to elevated CO2 and temperature in five British native species of contrasting functional type

Acclimation of photosynthesis to growth at elevated CO₂ concentration varies markedly between species. Species functionally classified as stress-tolerators (S) and ruderals (R), are thought to be incapable, or the least capable, of responding positively in terms of growth to elevated [CO₂]. Is this pattern of response also apparent in leaf photosynthesis of wild S- and R-strategists? Acclimatory loss of a photosynthetic and growth response to elevated [CO₂] is assumed to reflect limitation on capacity to utilize additional photosynthate. The doubling of pre-industrial global [CO₂] is expected to coincide with a 3 °C increase in mean temperature which could stimulate growth; will photosynthetic capacity at elevated [CO₂] be greater when the concurrent temperature increase is simulated? Five species from natural grassland of NW Europe and of contrasting ecological strategy were grown in hemispherical greenhouses, environmentally controlled to track the external microclimate. Within a replicated design, plants were grown at (i) current ambient [CO₂] and temperature, (ii) elevated [CO₂] (ambient + 340 μmol mol^–1) and ambient temperature, (iii) ambient [CO₂] and elevated temperature (ambient + 3 °C), or (iv) elevated [CO₂] and elevated temperature. After 75–104 days, the CO₂ response of light-saturated rates of photosynthesis (A_sat) was analysed in controlled-environment cuvettes in a field laboratory. There was no acclimatory loss of photosynthetic capacity with growth in elevated [CO₂] or elevated temperature over this period in Poa alpina (S), Bellis perennis (R) or Plantago lanceolata (mixed C-S-R strategist), and a significant (P ? ? bl 0.05) increase in capacity in Helianthemum nummularium (S) and Poa annua (R). Photosynthetic rates of leaves grown and measured in elevated [CO₂] were therefore significantly higher than rates for leaves grown and measured in ambient [CO₂], for all species. With the exception of Poa alpina, stomatal conductance and stomatal limitation on A_sat showed no acclimatory response to growth in elevated [CO₂]. Carboxylation efficiency, determined from the initial slope of the response of A_sat to intercellular CO₂ concentration was significantly increased by elevated [CO₂] and elevated temperature in H.nummularium, implying a possible increase in in vivo RubisCO activity. Increased carboxylation efficiency of this species was also reflected by an increase in the CO₂- and light-saturated rates of photosynthesis, indicating an increased capacity for regeneration of the primary CO₂ acceptor in photosynthesis. The results show that R-strategists and slow-growing S-strategists, are inherently capable of large increases in leaf photosynthetic capacity with growth in elevated [CO₂] in contrast to expectations from growth studies. With the exception of P.annua, where there was a significant negative interaction between CO₂ and temperature, concurrent increase in growth temperature had little effect on this pattern of response.     

Carbon balance of young birch trees grown in ambient and elevated atmospheric CO2 concentrations

We constructed a carbon budget for young birch trees grown in ambient and elevated CO₂ concentrations over their fourth year of growth. The annual total of net leaf photosynthesis was 110% more in elevated CO₂ than in ambient CO₂. However, the trees in elevated CO₂ grew only 59% more biomass than the trees in ambient CO₂ over the year. Modelling studies showed that larger loss of carbon from fine-root production and growth of the root-associated mycorrhiza by the trees in elevated CO₂ probably accounted for all the remaining difference in net photosynthesis between the two treatments. Our modelling also showed that the fraction of net photosynthate consumed by respiration of nonleaf tissue was similar in the two CO₂ treatments, and was 26% and 24% for trees in ambient and elevated CO₂, respectively. Trees in elevated CO₂ had 43% more leaves, and produced 110% more net photosynthate than trees in ambient CO₂, even though the maximum rate of carboxylation per unit leaf nitrogen decreased by 21%. Sensitivity studies showed that down-regulation reduced the annual net photosynthetic production of the trees in elevated CO₂ by only 6%. Direct effects of higher CO₂ on photosynthesis and greater leaf area of the trees in elevated CO₂ increased the net photosynthesis of the trees by 68% and 60%, respectively; and together accounted for most of the difference in net photosynthesis between the two treatments.     

Effects of elevated CO2 on leaf gas exchange in beech and oak at two levels of nutrient supply: consequences for sensitivity to drought in beech

Beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.) were grown from seed for two whole seasons at two CO₂ concentrations (ambient and ambient + 250 μmol mol^?1) with two levels of soil nutrient supply. Measurements of net leaf photosynthetic rate (A) and stomatal conductance (g_s) of well-watered plants were taken over both seasons; a drought treatment was applied in the middle of the second growing season to a separate sample of beech drawn from the same population. The net leaf photosynthetic rate of well-watered plants was stimulated in elevated CO₂ by an average of 75% in beech and 33% in oak; the effect continued through both growing seasons at both nutrient levels. There were no interactive effects of CO₂ concentration and nutrient level on A or g_s in beech or oak. Stomatal conductance was reduced in elevated CO₂ by an average of 34% in oak, but in beech there were no significant reductions in g_s except under cloudy conditions (–22% in elevated CO₂). During drought, there was no effect of CO₂ concentration on g_s in beech grown with high nutrients, but for beech grown with low nutrients, g_s was significantly higher in elevated CO₂, causing more rapid soil drying. With high nutrient supply, soil drying was more rapid at elevated CO₂ due to increased leaf area. It appears that beech may substantially increase whole-plant water consumption in elevated CO₂, especially under conditions of high temperature and irradiance when damage due to high evaporative demand is most likely to occur, thereby putting itself at risk during periods of drought.     

Seasonal response of photosynthesis to elevated CO2 in loblolly pine (Pinus taeda L.) over two growing seasons

Trees growing in natural systems undergo seasonal changes in environmental factors that generate seasonal differences in net photosynthetic rates. To examine how seasonal changes in the environment affect the response of net photosynthetic rates to elevated CO₂, we grew Pinus taeda L. seedlings for three growing seasons in open-top chambers continuously maintained at either ambient or ambient + 30 Pa CO₂. Seedlings were grown in the ground, under natural conditions of light, temperature nd nutrient and water availability. Photosynthetic capacity was measured bimonthly using net photosynthetic rate vs. intercellular CO₂ partial pressure (A-C_i) curves. Maximum Rubisco activity (Vc_max) and ribulose 1,5-bisphosphate regeneration capacity mediated by electron transport (J_max) and phosphate regeneration (PiRC) were calculated from A-C_i curves using a biochemically based model. Rubisco activity, activation state and content, and leaf carbohydrate, chlorophyll and nitrogen concentrations were measured concurrently with photosynthesis measurements. This paper presents results from the second and third years of treatment. Mean leaf nitrogen concentrations ranged from 13.7 to 23.8 mg g^?1, indicating that seedlings were not nitrogen deficient. Relative to ambient CO₂ seedlings, elevated CO₂ increased light-saturated net photosynthetic rates 60–110% during the summer, but < 30% during the winter. A relatively strong correlation between leaf temperature and the relative response of net photosynthetic rates to elevated CO₂ suggests a strong effect of leaf temperature. During the third growing season, elevated CO₂ reduced Rubisco activity 30% relative to ambient CO₂ seedlings, nearly completely balancing Rubisco and RuBP-regeneration regulation of photosynthesis. However, reductions in Rubisco activity did not eliminate the seasonal pattern in the relative response of net photosynthetic rates to elevated CO₂. These results indicate that seasonal differences in the relative response of net photosynthetic rates to elevated CO₂ are likely to occur in natural systems.     

Effects of elevated CO2 concentration and increased temperature on winter wheat: test of ARCWHEAT1 simulation model

Winter wheat (Triticum aestivum L., ev. Mercia) was grown in a controlled-environment facility at two CO₂ concentrations (targets 350 and 700 μmol mol^?1), and two temperature regimes (tracking ambient and ambient + 4°C). Observations of phenology, canopy growth, dry matter production and grain yield were used to test the ARCWHEAT1 simulation model. Dry-matter production and grain yield were increased at elevated CO₂ concentration (27 and 39%, respectively) and reduced at increased temperature (?16 and ?35%, respectively). ARCWHEAT1 substantially underestimated canopy growth for all treatments. However, differences in the facility environment from field conditions over the winter, indicated by the unusually rapid canopy growth observed in this period, meant that empirical model relationships were being used outside the conditions for which they were developed. The ARCWHEAT productivity submodel, given observed green area indices as inputs, overestimated the effect of CO₂ on productivity. An alternative, more mechanistic submodel of productivity, based on the SUCROS87 and Farquhar & von Caemmerer models, simulated observed crop biomass very closely. When these productivity simulations were inputed into the ARCWHEAT1 partitioning and grain-fill submodels, grain yield was predicted poorly, mainly as a result of the assumption that the number of grains is proportional to total growth during a short pre-anthesis phase. While yield was not correlated with growth in this phase, it was correlated with growth in longer pre-anthesis phases, indicating that ARCWHEAT1 could be improved by taking into account the contribution of earlier growth in determining yield.     

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.     

Effects of long-term exposure to elevated CO2 and N fertilization on the development of photosynthetic capacity and biomass accumulation in Quercus suber L.

The effects of long‐term (4 year) CO₂ enrichment (70 Pa versus 35 Pa) and nitrogen nutrition (8 mm versus 1 mm NO₃^–) on biomass accumulation and the development of photosynthetic capacity in leaves of cork oak (Quercus suber L., a Mediterranean evergreen tree) were studied. The evolution of photosynthetic parameters with leaf development was estimated by fitting the biochemical model of Farquhar et al. (Planta 149, 78–90, 1980) with modifications by Sharkey (Botanical Review 78, 71–75, 1985) to A–C_i response curves. CO₂ enrichment had a small reduction effect on the development of the maximum CO₂ fixation capacity by Rubisco (V_Cmax), and no effect over maximum electron transport capacity (J_max), day‐time respiration (R_d) and Triose‐P utilization (TPU). However, there was a statistically significant effect of N fertilization and the interaction CO₂ × N over the evolution of V_Cmax, J_max and TPU. Relative stomatal limitation (estimated from A–C_i curves) was higher (+20%) for plants grown under ambient CO₂ than for plants grown under elevated CO₂. There was a significant effect of CO₂ and N fertilization over total biomass accumulation as well as leaf area. Plants grown at elevated CO₂ had 27% more biomass than plants grown at ambient CO₂ when given high N. However, for plants grown under low N there was no significant effect of CO₂ enrichment on biomass accumulation. Plants grown under low N also had significantly higher root : shoot ratios whereas there were no differences between CO₂ treatments. The larger biomass accumulation of Q. suber under elevated CO₂ is attributable to a higher availability of CO₂ coupled to a larger leaf area, with no significant decrease in photosynthetic capacity under CO₂ enrichment and elevated N fertilization. For low N fertilization, the effects of CO₂ enrichment over leaf area and biomass accumulation are lost, suggesting that in native ecosystems with low N availability, the effects of CO₂ enrichment may be insignificant.     

Photosynthesis and conductance of spring-wheat leaves: field response to continuous free-air atmospheric CO2 enrichment

Spring wheat was grown from emergence to grain maturity in two partial pressures of CO₂ (pCO₂): ambient air of nominally 37 Pa and air enriched with CO₂ to 55 Pa using a free-air CO₂ enrichment (FACE) apparatus. This experiment was the first of its kind to be conducted within a cereal field without the modifications or disturbance of microclimate and rooting environment that accompanied previous studies. It provided a unique opportunity to examine the hypothesis that continuous exposure of wheat to elevated pCO₂ will lead to acclimatory loss of photosynthetic capacity. The diurnal courses of photosynthesis and conductance for upper canopy leaves were followed throughout the development of the crop and compared to model-predicted rates of photosynthesis. The seasonal average of midday photosynthesis rates was 28% greater in plants exposed to elevated pCO₂ than in contols and the seasonal average of the daily integrals of photosynthesis was 21% greater in elevated pCO₂ than in ambient air. The mean conductance at midday was reduced by 36%. The observed enhancement of photosynthesis in elevated pCO₂ agreed closely with that predicted from a mechanistic biochemical model that assumed no acclimation of photosynthetic capacity. Measured values fell below predicted only in the flag leaves in the mid afternoon before the onset of grain-filling and over the whole diurnal course at the end of grain-filling. The loss of enhancement at this final stage was attributed to the earlier senescence of flag leaves in elevated pCO₂. In contrast to some controlled-environment and field-enclosure studies, this field-scale study of wheat using free-air CO₂ enrichment found little evidence of acclimatory loss of photosynthetic capacity with growth in elevated pCO₂ and a significant and substantial increase in leaf photosynthesis throughout the life of the crop.     

Effects of Elevated CO2 and Defoliation on Compensatory Growth and Photosynthesis of Seedlings in a Tropical Tree,Copaifera aromatica1

After defoliation by herbivores, some plants exhibit enhanced rates of photosynthesis and growth that enable them to compensate for lost tissue, thus maintaining their fitness relative to competing, undefoliated plants. Our aim was to determine whether compensatory photosynthesis and growth would be altered by increasing concentrations of atmospheric CO₂. Defoliation of developing leaflets on seedlings of a tropical tree, Copaifera aromatica, caused increases in photosynthesis under ambient CO₂, but not under elevated CO₂. An enhancement in the development of buds in the leaf axils followed defoliation at ambient levels of CO₂. In contrast, under elevated CO₂, enhanced development of buds occurred in undefoliated plants with no further enhancement in bud development due to exposure to elevated CO₂. Growth of leaf area after defoliation was increased, particularly under elevated CO₂. Despite this increase, defoliated plants grown under elevated CO₂ were further from compensating for tissue lost during defoliation after 5¹/₂ weeks than those grown under ambient CO₂ concentrations.     

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.     

NMR imaging of root water distribution in intact Vicia faba L plants in elevated atmospheric CO2

The effect of elevated atmospheric CO₂ on water distribution in the intact roots of Vicia faba L. bean seedlings grown in natural soil was studied noninvasively with proton (¹H) nuclear magnetic resonance (NMR) imaging. Exposure of 24-d-old plants to atmospheric CO₂-enriched air at 650 cm³ m^?3 produced significant increases in water imaged in upper roots, hypogeal cotyledons and lower stems in response to a short-term drying-stress cycle. Above ground, drying produced negligible stem shrinkage and stomatal resistance was unchanged. In contrast, the same drying cycle caused significant depletion of water imaged in the same upper root structures in control plants subject to ambient CO₂ (350 m³ m^?3), and stem shrinkage and increased stomatal resistance. The results suggest that inhibition of transpiration caused by elevated CO₂ does not necessarily result in attenuation of water transport from lower root structures. Inhibition of water loss from upper roots and lower stem in elevated CO₂ environments may be a mitigating factor in assessing deleterious effects of greenhouse changes on crops during periods of dry climate.