Effect of elevated CO2 on the stomatal distribution and leaf physiology of Alnus glutinosa

Variation in stomatal development and physiology of mature leaves from Alnus glutinosa plants grown under reference (current ambient, 360 μmol mol⁻¹ CO₂) and double ambient (720 μmol mol⁻¹ CO₂) carbon dioxide (CO₂) mole fractions is assessed in terms of relative plant growth, stomatal characters (i.e. stomatal index and density) and leaf photosynthetic characters. This is the first study to consider the effects of elevated CO₂ concentration on the distribution of stomata and epidermal cells across the whole leaf and to try to ascertain the cause of intraleaf variation. In general, a doubling of the atmospheric CO₂ concentration enhanced plant growth and significantly increased stomatal index. However, there was no significant change in relative stomatal density. Under elevated CO₂ concentration there was a significant decrease in stomatal conductance and an increase in assimilation rate. However, no significant differences were found for the maximum rate of carboxylation ( V _cmax) and the light saturated rate of electron transport ( J _max) between the control and elevated CO₂ treatment.     

Growth and physiological responses of canola (Brassica napus) to three components of global climate change: temperature, carbon dioxide and drought

Elevated CO₂ appears to be a significant factor in global warming, which will likely lead to drought conditions in many areas. Few studies have considered the interactive effects of higher CO₂, temperature and drought on plant growth and physiology. We grew canola ( Brassica napus cv. 45H72) plants under lower (22/18°C) and higher (28/24°C) temperature regimes in controlled-environment chambers at ambient (370 μmol mol⁻¹) and elevated (740 μmol mol⁻¹) CO₂ levels. One half of the plants were watered to field capacity and the other half at wilting point. In three separate experiments, we determined growth, various physiological parameters and content of abscisic acid (ABA), indole-3-acetic acid and ethylene. Drought-stressed plants grown under higher temperature at ambient CO₂ had decreased stem height and diameter, leaf number and area, dry matter, leaf area ratio, shoot/root weight ratio, net CO₂ assimilation and chlorophyll fluorescence. However, these plants had increased specific leaf weight, leaf weight ratio and chlorophyll concentration. Elevated CO₂ generally had the opposite effect, and partially reversed the inhibitory effects of higher temperature and drought on leaf dry weight accumulation. This study showed that higher temperature and drought inhibit many processes but elevated CO₂ partially mitigate some adverse effects. As expected, drought stress increased ABA but higher temperature inhibited the ability of plants to produce ABA in response to drought.     

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

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

Effects of manganese toxicity on leaf CO2 assimilation of contrasting common bean genotypes

Parameters related to leaf photosynthesis were evaluated in three genotypes of common bean ( Phaseolus vulgaris L.) with contrasting tolerance to Mn toxicity. Two short-term studies in solution culture were used to assess the effect of excess Mn on CO₂ assimilation in mature and immature leaves. Mn toxicity decreased total chlorophyll content only in immature leaves, with a consequent reduction of leaf CO₂ assimilation. Mature leaves that showed brown speckles characteristic of Mn toxicity, did not suffer any detriment in their capacity to assimilate CO₂, at least in a 4-day experiment. Stomatal conductance and transpiration were not affected by the presence of high levels of Mn in leaf tissue. Lower stomatal conductance and transpiration rates were observed only in leaves with advanced chlorosis. Differences among genotypes were detected as increased chlorosis in the more sensitive genotype ZPV-292, followed by A-283 and less chlorosis in the tolerant genotype CALIMA. Since CO₂ assimilation expressed per unit of chlorophyll was not different between high-Mn plants and control plants, we conclude that the negative effect of Mn toxicity on CO₂ assimilation can be explained by a reduction in leaf chlorophyll content.     

Acquisition and allocation of carbon and nitrogen by Danthonia richardsonii in response to restricted nitrogen supply and CO2 enrichment

Dry weight (DW) and nitrogen (N) accumulation and allocation were measured in isolated plants of Danthonia richardsonii (Wallaby Grass) for 37 d following seed imbibition. Plants were grown at ≈ 365 or 735 μ L L^–1 CO₂ with N supply of 0·05, 0·2 or 0·5 mg N plant^–1 d^–1. Elevated CO₂ increased DW accumulation by 28% (low-N) to 103% (high-N), following an initial stimulation of relative growth rate. Net assimilation rate and leaf nitrogen productivity were increased by elevated CO₂, while N concentration was reduced. N uptake per unit root surface area was unaffected by CO₂ enrichment. The ratio of leaf area to root surface area was decreased by CO₂ enrichment. Allometric analysis revealed a decrease in the shoot-N to root-N ratio at elevated CO₂, while the shoot-DW to root-DW ratio was unchanged. Allometric analysis showed leaf area was reduced, while root surface area was unchanged by elevated CO₂, indicating a down-regulation of total plant capacity for carbon gain rather than a stimulation of mineral nutrient acquisition capacity. Overall, growth in elevated CO₂ resulted in changes in plant morphology and nitrogen use, other than those associated simply with changing plant size and non-structural carbohydrate content.     

Ecophysiological responses to simulated canopy gaps of two tree species of contrasting shade tolerance in elevated CO2

1. One-year-old seedlings of shade tolerant Acer rubrum and intolerant Betula papyrifera were grown in ambient and twice ambient (elevated) CO₂, and in full sun and 80% shade for 90 days. The shaded seedlings received 30-min sun patches twice during the course of the day. Gas exchange and tissue–water relations were measured at midday in the sun plants and following 20 min of exposure to full sun in the shade plants to determine the effect of elevated CO₂ on constraints to sun-patch utilization in these species.
2. Elevated CO₂ had the largest stimulation of photosynthesis in B. papyrifera sun plants and A. rubrum shade plants.
3. Higher photosynthesis per unit leaf area in sun plants than in shade plants of B. papyrifera was largely owing to differences in leaf morphology. Acer rubrum exhibited sun/shade differences in photosynthesis per unit leaf mass consistent with biochemical acclimation to shade.
4. Betula papyrifera exhibited CO₂ responses that would facilitate tolerance to leaf water deficits in large sun patches, including osmotic adjustment and higher transpiration and stomatal conductance at a given leaf-water potential, whereas A. rubrum exhibited large increases in photosynthetic nitrogen-use efficiency.
5. Results suggest that species of contrasting successional ranks respond differently to elevated CO₂, in ways that are consistent with the habitats in which they typically occur.     

Elevated atmospheric CO2 lowers leaf litter nutritional quality for stream ecosystem food webs

Up to 99% of the carbon fuelling the food webs of temperate woodland streams is derived from inputs of terrestrial leaf litter. Aquatic bacteria, fungi, and detritivore invertebrates directly utilize these inputs, transferring this energy to other components of the food web. Increases in atmospheric CO₂ could indirectly impact woodland stream food webs by chemically altering leaf litter. This study evaluated CO₂-induced chemical changes in aspen ( Populus tremuloides ) leaf litter, and the corresponding effects on stream bacteria, fungi and leaf-shredding cranefly larvae ( Tipula abdominalis : Diptera). Leaf litter from plants grown under elevated CO₂ had decreased nutritional value to aquatic decomposers and detritivores because of higher levels of structural compounds and lower nitrogen content. Consequently, elevated CO₂-grown leaf litter supported 59% lower bacterial production in a stream than litter grown at ambient CO₂ levels, while not affecting fungal biomass. Larval craneflies fed elevated CO₂-grown microbially colonized leaves consumed less, assimilated less, and grew 12 times slower than their ambient fed counterparts.     

Photosynthesis affects following night leaf conductance in Vicia faba

Night-time stomatal opening in C₃ plants may result in significant water loss when no carbon gain is possible. The objective of this study was to determine if endogenous patterns of night-time stomatal opening, as reflected in leaf conductance, in Vicia faba are affected by photosynthetic conditions the previous day. Reducing photosynthesis with low light or low CO₂ resulted in reduced night-time stomatal opening the following night, irrespective of the effects on daytime stomatal conductance. Likewise, increasing photosynthesis with enriched CO₂ levels resulted in increased night-time stomatal opening the following night. Reduced night-time stomatal opening was not the result of an inability to regulate stomatal aperture as leaves with reduced night-time stomatal opening were capable of greater night-time opening when exposed to low CO₂. After acclimating plants to long or short days, it was found that night-time leaf conductance was greater in plants acclimated to short days, and associated with greater leaf starch and nitrate accumulation, both of which may affect night-time guard cell osmotic potential. Direct measurement of guard cell contents during endogenous night-time stomatal opening will help identify the mechanism of the effect of daytime photosynthesis on subsequent night-time stomatal regulation.     

Photosynthetic acclimation to elevated CO2 is modified by source:sink balance in three component species of chalk grassland swards grown in a free air carbon dioxide enrichment (FACE) experiment

Artificial chalk grassland swards were exposed to either ambient air or air enriched to 600 μ mol mol^–1 CO₂, using free-air CO₂ enrichment technology, and subjected to an 8 week simulated grazing regime. After 14 months of treatment, ribulose-1,5-bisphosphate carboxylase (Rubisco) activity ( V _c,max) and electron transport mediated ribulose-1,5-bisphosphate (RuBP) regeneration capacity ( J _max), estimated from leaf gas exchange, were significantly lower in fully expanded leaves of Anthyllis vulneraria L. (a legume) and Sanguisorba minor Scop. grown in elevated CO₂. After a change in source:sink balance brought about by defoliation, photosynthetic capacity was fully restored in A. vulneraria and S. minor, but acclimation continued in the grass Bromopsis erecta (Hudson) Fourr. Changes in net photosynthesis ( P _n) with growth at elevated CO₂ ranged from a 1·6% reduction in precut leaves of A. vulneraria to a 47·1% stimulation in postcut leaves of S. minor . Stomatal acclimation was observed in leaves of A. vulneraria (reduced stomatal density) and B. erecta (reduced stomatal conductance). The results are discussed in terms of whole-plant resource-use optimization and chalk grassland community competitive interactions at elevated CO₂.     

The interaction of rising CO2 and temperatures with water use efficiency

Abstract. Recent data concerning the impact of elevated atmospheric CO₂ upon water use efficiency (WUE) and the related measure, instantaneous transpiration efficiency (ITE), are reviewed. It is concluded from both short and long-term studies that, at the scale of the individual leaf or plant, an increase in WUE or ITE is generally observed in response to increased atmospheric CO₂ levels. However, the magnitude of this increase may decline with time. The opinion that elevated CO₂ may substantially decrease transpiration at the regional scale is discussed. The mechanisms by which elevated CO₂ may cause a change in these measures are discussed in terms of stomatal conductance, assimilation and respiration responses to elevated CO₂. Finally, recent experimental data and model outputs concerning the impact of the interaction of increased temperature with elevated CO₂ on WUE, ITE and yield are reviewed. It is concluded that substantially more data is required before reliable predictions about the regional scale response of WUE and catchment hydrology can be made.     

The role of temperature in determining the stimulation of CO2 assimilation at elevated carbon dioxide concentration in soybean seedlings

Soybean ( Glycine max cv. Clark) was grown at both ambient (ca 350 μmol mol⁻¹) and elevated (ca 700 μmol mol⁻¹) CO₂ concentration at 5 growth temperatures (constant day/night temperatures of 20, 25, 30, 35 and 40°C) for 17–22 days after sowing to determine the interaction between temperature and CO₂ concentration on photosynthesis (measured as A, the rate of CO₂ assimilation per unit leaf area) at both the single leaf and whole plant level. Single leaves of soybean demonstrated increasingly greater stimulation of A at elevated CO₂ as temperature increased from 25 to 35°C (i.e. optimal growth rates). At 40°C, primary leaves failed to develop and plants eventually died. In contrast, for both whole plant A and total biomass production, increasing temperature resulted in less stimulation by elevated CO₂ concentration. For whole plants, increased CO₂ stimulated leaf area more as growth temperature increased. Differences between the response of A to elevated CO₂ for single leaves and whole plants may be related to increased self-shading experienced by whole plants at elevated CO₂ as temperature increased. Results from the present study suggest that self-shading could limit the response of CO₂ assimilation rate and the growth response of soybean plants if temperature and CO₂ increase concurrently, and illustrate that light may be an important consideration in predicting the relative stimulation of photosynthesis by elevated CO₂ at the whole plant level.     

Interaction of elevated CO2 and O3 on growth, photosynthesis and respiration of three perennial species grown in low and high nitrogen

Seedlings of three species native to central North America, a C₃ tree, Populus tremuloides Michx., a C₃ grass, Agropyron smithii Rybd., and a C₄ grass, Bouteloua curtipendula Michx., were grown in all eight combinations of two levels each of CO₂, O₃ and nitrogen (N) for 58 days in a controlled environment. Treatment levels consisted of 360 or 674 μmol mol^-1 CO₂, 3 or 92 nmol mol^-1 O₃, and 0.5 or 6.0 m M N. In situ photosynthesis and relative growth rate (RGR) and its determinants were obtained at each of three sequential harvests, and leaf dark respiration was measured at the second and third harvests. In all three species, plants grown in high N had significantly greater whole-plant mass, RGR and photosynthesis than plants grown in low N. Within a N treatment, elevated CO₂ did not significantly enhance any of these parameters nor did it affect leaf respiration. However, plants of all three species grown in elevated CO₂ had lower stomatal conductance compared to ambient CO₂-exposed plants. Seedlings of P. tremuloides (in both N treatments) and B. curtipendula (in high N) had significant ozone-induced reductions in whole-plant mass and RGR in ambient but not under elevated CO_2. This negative O₃ impact on RGR in ambient CO₂ was related to increased leaf dark respiration, decreased photosynthesis and/or decreased leaf area ratio, none of which were noted in high O₃ treatments in the elevated CO₂ environment. In contrast, A. smithii was marginally negatively affected by high O_3.     

Acclimation of Arabidopsis thaliana to long-term CO2 enrichment and nitrogen supply is basically a matter of growth rate adjustment

The long-term response of Arabidopsis thaliana to increasing CO₂ was evaluated in plants grown in 800 μl l⁻¹ CO₂ from sowing and maintained, in hydroponics, on three nitrogen supplies: "low,""medium" and "high." The global response to high CO₂ and N-supply was evaluated by measuring growth parameters in parallel with photosynthetic activity, leaf carbohydrates, ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) messenger RNA and protein, stomatal conductance (g_s) and density. CO₂ enrichment was found to stimulate biomass production, whatever the N-supply. This stimulation was transient on low N-supply and persisted throughout the whole vegetative growth only in high N-supply. Acclimation on low N–high CO₂ was not associated with carbohydrate accumulation or with a strong reduction in Rubisco amount or activity. At high N-supply, growth stimulation by high CO₂ was mainly because of the acceleration of leaf production and expansion while other parameters such as specific leaf area, root/shoot ratio and g_s appeared to be correlated with total leaf area. Our results thus suggest that, in strictly controlled and stable growing conditions, acclimation of A. thaliana to long-term CO₂ enrichment is mostly controlled by growth rate adjustment.     

Interaction of enriched CO2 and water stress on the physiology of and biomass production in sweet potato grown in open-top chambers

Abstract. The objective of this study was to investigate the effects of water stress in sweet potato ( Ipomoea batatas L. [Lam] 'Georgia Jet') on biomass production and plant-water relationships in an enriched CO₂ atmosphere. Plants were grown in pots containing sandy loam soil (Typic Paleudult) at two concentrations of elevated CO₂ and two water regimes in open-top field chambers. During the first 12 d of water stress, leaf xylem potentials were higher in plants grown in a CO₂ concentration of 438 and 666 μmol mol⁻¹ than in plants grown at 364 μmol mol⁻¹. The 364 μmol mol⁻¹ CO₂ grown plants had to be rewatered 2 d earlier than the high CO₂-grown plants in response to water stress. For plants grown under water stress, the yield of storage roots and root: shoot ratio were greater at high CO₂ than at 364 μmol mol⁻¹; the increase, however, was not linear with increasing CO₂ concentrations. In well-watered plants, biomass production and storage root yield increased at elevated CO₂, and these were greater as compared to water-stressed plants grown at the same CO₂ concentration.     

Low carbon dioxide concentrations can reverse stomatal closure during water stress

Leaf water potentials below threshold values result in reduced stomatal conductance (g_s). Stomatal closure at low leaf water potentials may serve to protect against cavitation of xylem. Possible control of g_s by leaf water potential or hydraulic conductance was tested by drying the rooting medium in four herbaceous annual species until g_s was reduced and then lowering the [CO₂] to determine whether g_s and transpiration rate could be increased and leaf water potential decreased and whether hydraulic conductance was reduced at the resulting lower leaf water potential. In all species, low [CO₂] could reverse the stomatal closure because of drying despite further reductions in leaf water potential, and the resulting lower leaf water potentials did not result in reductions in hydraulic conductance. The relative sensitivity of g_s to internal [CO₂] in the leaves of dry plants of each species averaged three to four times higher than in leaves of wet plants. Two species in which g_s was reputed to be insensitive to [CO₂] were examined to determine whether high leaf to air water vapor pressure differences (D) resulted in increased stomatal sensitivity to [CO₂]. In both species, stomatal sensitivity to [CO₂] was indeed negligible at low D, but increased with D, and low [CO₂] partly or fully reversed closure caused by high D. In no case did low leaf water potential or low hydraulic conductance during drying of the air or the rooting medium prevent low [CO₂] from increasing g_s and transpiration rate.     

Elevated CO2 concentrations and stomatal density: observations from 17 plant species growing in a CO2 spring in central Italy

Stomatal density (SD) and stomatal conductance ( g _s) can be affected by an increase of atmospheric CO₂ concentration. This study was conducted on 17 species growing in a naturally enriched CO₂ spring and belonging to three plant communities. Stomatal conductance, stomatal density and stomatal index (SI) of plants from the spring, which were assumed to have been exposed for generations to elevated [CO₂], and of plants of the same species collected in a nearby control site, were compared. Stomatal conductance was significantly lower in most of the species collected in the CO₂ spring and this indicated that CO₂ effects on g _s are not of a transitory nature but persist in the long term and through plant generations. Such a decrease was, however, not associated with changes in the anatomy of leaves: SD was unaffected in the majority of species (the decrease was only significant in three out of the 17 species examined), and also SI values did not vary between the two sites with the exception of two species that showed increased SI in plants grown in the CO₂-enriched area. These results did not support the hypothesis that long-term exposure to elevated [CO₂] may cause adaptive modification in stomatal number and in their distribution.     

Fruit effects on photosynthesis in Prunus persica

Seasonal measurements of net CO₂ assimilation, leaf conductance and mesophyll conductance were made in the field on mature, fruiting and defruited Prunus persica L. Batsch trees. During early stages of fruit growth there were no significant differences in leaf gas exchange characteristics between fruiting and defruited trees. During the early part of the last stage of fruit growth, CO₂ assimilation rates were 11–15% higher in fruiting trees than defruited trees. These increased assimilation rates corresponded with approximately 30% increases in leaf conductance and only minor changes in mesophyll conductances or leaf CO₂ assimilation capacity as indicated by leaf nitrogen content. It is concluded that under the field conditions of this study the fruit effect on photosynthesis is primarily related to stomatal behavior.     

CO2 enrichment, drought stress and growth of Alaska pea plants (Pisum sativum)

The interaction of CO₂ enrichment and drought on water status and growth of pea plants was investigated. Pisum sativum L. (cv. Alaska) plants were grown from seeds in growth chambers using 350 and 675 μl I1 CO₂, a photon flux density of 600 μmol M-2 S-1, a 16 h photoperiod and a temperature regime of 20/14°C. The drought treatment was started at the beginning of branch initiation and lasted for 9 or 11 days. The water status of the plants was monitored daily by measuring total leaf water potential and stomatal conductance. The total leaf water potential of well-watered plants was not affected by the CO₂ level. Under draughting conditions total leaf water potential decreased, with a slower decrease under the high CO₂ regime, due, at least in part, to reduced stomatal conductance. Upon rewatering, total leaf water potential and stomatal conductance recovered within one day. High CO₂ counteracted the reduction in height and, to some extent, leaf area that developed in low CO₂ unwatered plants. Additional CO₂ had no effect on branch number and did not prevent the complete inhibition of branch development that resulted from drought stress. Removing the drought conditions resulted in a rapid recovery of the internal water status and also a rapid recovery of most, but not all, plant growth parameters.     

Water vapour fluxes and their impact under elevated CO2 in a C4-tallgrass prairie

We measured leaf-level stomatal conductance, xylem pressure potential, and stomate number and size as well as whole plant sap flow and canopy-level water vapour fluxes in a C4-tallgrass prairie in Kansas exposed to ambient and elevated CO₂. Stomatal conductance was reduced by as much as 50% under elevated CO₂ compared to ambient. In addition, there was a reduction in stomate number of the C4 grass, Andropogon gerardii Vitman, and the C3 dicot herb, Salvia pitcheri Torr., under elevated CO₂ compared to ambient. The result was an improved water status for plants exposed to elevated CO₂ which was reflected by a less negative xylem pressure potential compared to plants exposed to ambient CO₂. Sap flow rates were 20 to 30% lower for plants exposed to elevated CO₂ than for those exposed to ambient CO₂. At the canopy level, evapotranspiration was reduced by 22% under elevated CO₂. The reduced water use by the plant canopy under elevated CO₂ extended the photosynthetically-active period when water became limiting in the ecosystem. The result was an increased above- and belowground biomass production in years when water stress was frequent.     

The response of nodulated alfalfa to water supply, temperature and elevated CO2: photosynthetic downregulation

Plants grown in an environment of elevated CO₂ and temperature often show reduced CO₂ assimilation capacity, providing evidence of photosynthetic downregulation. The aim of this study was to analyse the downregulation of photosynthesis in elevated CO₂ (700 µmol mol⁻¹) in nodulated alfalfa plants grown at different temperatures (ambient and ambient + 4°C) and water availability regimes in temperature gradient tunnels. When the measurements were taken in growth conditions, a combination of elevated CO₂ and temperature enhanced the photosynthetic rate; however, when they were carried out at the same CO₂ concentration (350 and 700 µmol mol⁻¹), elevated CO₂ induced photosynthetic downregulation, regardless of temperature and drought. Intercellular CO₂ concentration measurements revealed that photosynthetic acclimation could not be accounted for by stomatal limitations. Downregulation of plants grown in elevated CO₂ was a consequence of decreased carboxylation efficiency as a result of reduced rubisco activity and protein content; in plants grown at ambient temperature, downregulation was also induced by decreased quantum efficiency. The decrease in rubisco activity was associated with carbohydrate accumulation and depleted nitrogen availability. The root nodules were not sufficiently effective to balance the source–sink relation in elevated CO₂ treatments and to provide the required nitrogen to counteract photosynthetic acclimation.