The Inhibition of Photosynthesis after Exposure of Bean Leaves to Various Low Levels of CO2

Mature first leaves of Phaseolus vulgaris L. were exposed tolow partial pressures of CO₂ (7, 6 and 0 Pa CO₂) for 24 h. Afterexposure of leaves to 6 Pa CO₂ for 24 h, there was a reductionin the carbon exchange rate (CER) at all partial pressures ofCO₂ at which measurements were made. After exposure to 7 PaCO₂, the CER decreased only at high partial pressures of CO₂.The rates of electron transport from water to methyl viologen,through the whole chain, decreased in parallel with the decreasein CER measured at 90 Pa CO₂. One site of inhibition in leavesexposed for 24 h to 6 Pa CO₂ appeared to be the intersystemelectron-transport chain since there were no significant changesin the activities of PSI and PSII, as determined from the levelof P-700 and measurement of fluorescence, respectively. Anotherinhibitory phenomenon appeared to be a negative change in theactivation state of Rubisco, while the level of Rubisco wasunaffected by the exposure to 6 Pa CO₂. These decreases in photosyntheticactivity caused by depletion of CO₂ explains at least in part,the inhibition of photosynthesis that is caused by rain treatment[Ishibashi and Terashima (1995) Plant Cell Environ. 18: 431]. (Received September 19, 1996; Accepted March 10, 1997)     

Effect of CO2 Concentration During Growth on a CO2 Concentrating Mechanism in White Clover as Predicted from Differential 14CO2/12CO2Uptake

Lehnherr, B. M?chler, F. and N?sberger, J. 1985. Effect of CO₂concentration during growth on a CO₂ concentrating mechanismin white clover as predicted from differential ¹⁴CO₂/¹²CO² uptake.-J. exp. Bot. 36: 1835-1841. White clover was grown at 20 and100 Pa p(CO₂). The CO₂ response of net photosynthesis and differentialuptake of ¹⁴CO₂ and ¹²CO₂ by leaves were measured at varioustemperatures and at various O₂ and CO₂ partial pressures andcompared with predictions from ribulose bisphosphate carboxylase/oxygenasekinetics. Discrepancies between the observed gas exchange characteristicsfor the leaves and those predicted from the enzyme kineticswere interpreted as being due to a CO₂ concentrating mechanism.Plants grown at 20 Pa p(CO₂) showed a higher affinity for CO₂than plants grown at 100 Pa p(CO₂) when measured at 10 ?C. Nodifference in affinity was found at 30 ?C. The postulated CO₂concentrating effect was greater in plants grown at low CO₂than in plants grown at high CO₂ concentration and occurredonly at low temperature and low CO₂ partial pressure. It issuggested that plants grown at the lower CO₂ partial pressurehave a higher affinity for CO₂ due to a more efficient CO₂ concentratingsystem than plants grown at the higher CO₂ partial pressure. Key words: Photosynthesis, CO₂, concentration, RuBP carboxylase/oxygenase     

Humidity Pretreatment Affects the Responses of Stomata and CO2 Assimilation to Vapor Pressure Difference in C3 and C4 Plants

Experiments were carried out to investigate the long-term influenceof humidity on the short-term responses of stomata and CO₂ assimilationto vapor pressure difference in Oryza sativa (rice, C₃ species)and Panicum maximum (green panic, C₄ species). Plants were grownfor four weeks in growth chambers set at 35% and 85% relativehumidity at 25C air temperature, 38+2 Pa CO₂ partial pressureand 1,700µmol m^-2s^-1 photon flux density. Soil was saturatedwith water in both humidity treatments. Low humidity pretreatmentscaused low leaf conductance and low rates of transpiration andCO₂ assimilation in O. sativa, but small changes in stomatalresponses to humidity and in CO₂ assimilation were found inP. maximum. From the short-term gas exchange experiments, itwas noted that the responsiveness of leaf conductance to vaporpressure difference were affected by humidity pretreatmentsin O. sativa, whereas unaffected in P. maximum. In O. sativameasurements of CO₂ assimilation as a function of internal CO₂partial pressure (A-Ci curve) indicated that low humidity pretreatmentsreduced the CO₂ assimilation at high internal CO₂ partial pressure,but the initial slope of the A-Ci curve was unaffected. Furthermore,plant characteristics such as total dry weight and leaf areaof plants subjected to low umidity were lower than plants subjectedto high humidity. The reductions in O. sativa, however, werelarger than in P. maximum. Stomatal frequency from low humiditygrown plant was higher than that from high humidity grown plantsin both species although there is no significant difference.The data indicated that if the short term inhibition of netCO₂ assimilation at a high vapor pressure difference was imposedduring vegetative growth, the photosynthetic biochemistry andthe resultant plant growth were largely depressed in O. sativa,a C₃ species. (Received May 26, 1992; Accepted November 2, 1992)     

Variations of stomatal density and carbon isotope values of<Emphasis Type="Italic"> Picea crassifolia</Emphasis> at different altitudes in the Qilian Mountains

The stomatal characteristics, length and dry weight as well as stable carbon isotope composition (i¹³C) of needles and tree rings of Qinghai spruce (Picea crassifolia) at different altitudes in the Qilian mountains were investigated. The results showed that stomatal density, distribution pattern on epidermis, and length and dry weight of needles all increased with altitude below 3,000 m. In contrast, these parameters all decreased with increasing altitude above 3,000 m. Furthermore an assay of tree rings showed that near 3,000 m in altitude was the optimum zone for growth and development of Qinghai spruce. Although atmospheric CO₂ concentration influences stomatal density, the effects of many environmental factors, such as temperature, light and rainfall, could obscure the effects of changes in atmospheric CO₂ concentration on stomatal density at altitudes higher than the optimum for growth. The correlation of stomatal density with atmospheric CO₂ concentration and i¹³C of Qinghai spruce needles was significant below 3,000 m, but was insignificant above 3,000 m altitude. The i¹³C value of needles, however, was negatively correlated with atmospheric CO₂ concentrations. Therefore, the influence of altitude should be considered in research on plant physiological ecology and the relationship of stomatal density with i¹³C value or atmospheric CO₂ concentration.     

Separation of Direct and Indirect Responses of Stomata to Light: Results from a Leaf Inversion Experiment at Constant Intercellular CO2 Molar Fraction

Previous work has shown that stomata respond directly to light,but it was not clear whether the only additional response isthrough CO₂, or whether some other metabolite is involved inthis response. Gas exchange experiments were done with normallypositioned and inverted leaves of Hedera helix to investigatethis problem. The macroscopic optical properties of the leavesand their anatomical structure were also studied. These experimentssnowed that there is no need to postulate the existence of amessenger other than CO₂ to explain the indirect response ofstomata to light. The experiments also showed that leaf inversionaffects both stomatal conductance and photosynthesis, and highlightthe difficulties involved in the interpretation of the effectof leaf inversion on stomata when stomatal conductance measurementsare not done concurrently with measurements of CO₂ flux densityand intercellular CO₂ molar fraction Key words: Hedera helix, ivy, gas exchange, leaf inversion, stomatal conductance, light, CO₂ flux density, photosynthesis     

Stomatal Responses of Variegated Leaves to CO2 Enrichment

The responses of stomatal density and stomatal index of fivespecies of ornamental plants with variegated leaves grown attwo mole fractions of atmospheric CO₂ (350 and 700 µmolmol^-1) were measured. The use of variegated leaves allowed anypotential effects of mesophyll photosynthetic capacity to beuncoupled from the responses of stomatal density to changesin atmospheric CO₂ concentration. There was a decrease in stomataldensity and stomatal index with CO₂ enrichment on both white(unpigmented) and green (pigmented) leaf areas. A similar responseof stomatal density and index was also observed on areas ofleaves with pigmentation other than green indicating that anydifferences in metabolic processes associated with colouredleaves are not influencing the responses of stomatal densityto CO₂ concentrations. Therefore the carboxylation capacityof mesophyll tissue has no direct influence on stomatal densityand index responses as suggested previously (Friend and Woodward1990 Advances in Ecological Research 20: 59-124), instead theresponses were related to leaf structure. The stomatal characteristics(density and index) of homobaric variegated leaves showed agreater sensitivity to CO₂ on green portions, whereas heterobaricleaves showed a greater sensitivity on white areas. These resultsprovide evidence that leaf structure may play an important rolein determining the magnitude of stomatal density and index responsesto CO₂ concentrations.Copyright 1995, 1999 Academic Press Leaf structure, photosynthesis, stomatal conductance, CO₂, stomatal density, stomatal index     

Effect of Elevated CO2 on Stomatal Density and Distribution in a C4 Grass and a C3 Forb under Field Conditions

Two common tallgrass prairie species, Andropogon gerardii, thedominant C₄ grass in this North American grassland, and Salviapitcheri, a C₃ forb, were exposed to ambient and elevated (twiceambient) CO₂ within open-top chambers throughout the 1993 growingseason. After full canopy development, stomatal density on abaxialand adaxial surfaces, guard cell length and specific leaf mass(SLM; mg cm^-2) were determined for plants in the chambers aswell as in adjacent unchambered plots. Record high rainfallamounts during the 1993 growing season minimized water stressin these plants (leaf xylem pressure potential was usually >-1·5 MPa in A. gerardii) and also minimized differencesin water status among treatments. In A. gerardii, stomatal densitywas significantly higher (190 ± 7 mm^-2; mean ±s.e.) in plants grown outside of the chambers compared to plantsthat developed inside the ambient CO₂ chambers (161 ±5 mm^-2). Thus, there was a significant 'chamber effect' on stomataldensity. At elevated levels of CO₂, stomatal density was evenlower (P < 0·05; 121 ± 5 mm^-2). Most stomatawere on abaxial leaf surfaces in this grass, but the ratio ofadaxial to abaxial stomatal density was greater at elevatedlevels of CO₂. In S. pitcheri, stomatal density was also significantlylower when plants were grown in the open-top chambers (235 ±10 mm^-2 outside vs. 140 ± 6 mm^-2 in the ambient CO₂ chamber).However, stomatal density was greater at elevated CO₂ (218 ±12 mm^-2) compared to plants from the ambient CO₂ chamber. Theratio of stomata on adaxial vs. abaxial surfaces did not varysignificantly in this herb. Guard cell lengths were not significantlyaffected by growth in the chambers or by elevated CO₂ for eitherspecies. Growth within the chambers resulted in lower SLM inS. pitcheri, but CO₂ concentration had no effect. In A. gerardii,SLM was lower at elevated CO₂. These results indicate that stomataland leaf responses to elevated CO₂ are species specific, andreinforce the need to assess chamber effects along with treatmenteffects (CO₂) when using open-top chambers.Copyright 1994, 1999Academic Press Andropogon gerardii, elevated CO₂, Salvia pitcheri, stomatal density, tallgrass prairie     

Stomatal Density and Index of Fossil Plants Track Atmospheric Carbon Dioxide in the Palaeozoic

It has been demonstrated that the leaves of a range of foresttree species have responded to the rising concentration of atmosphericCO₂ over the last 200 years by a decrease in both stomatal densityand stomatal index. This response has also been demonstratedexperimentally by growing plants under elevated CO₂ concentrations.Investigation of Quaternary fossil leaves has shown a correspondingstomatal response to changing CO₂ concentrations through a glacial-interglacialcycle, as revealed by ice core data. Tertiary leaves show asimilar pattern of stomatal density change, using palynologicalevidence of palaeo-temperature as a proxy measure of CO₂ concentration.The present work extends this approach into the Palaeozoic fossilplant record. The stomatal density and index of Early Devonian,Carboniferous and Early Permian plants has been investigated,to test for any relationship that they may show with the changesin atmospheric CO₂ concentration, derived from physical evidence,over that period. Observed changes in the stomatal data givesupport to the suggestion from physical evidence, that atmosphericCO₂ concentrations fell from an Early Devonian high of 10-12times its present value, to one comparable to that of the presentday by the end of the Carboniferous. These results suggest thatstomatal density of fossil leaves has potential value for assessingchanges in atmospheric CO₂ concentration through geologicaltime.Copyright 1995, 1999 Academic Press Aglaophyton major, Sawdonia ornata, Swillingtonia denticulata, Lebachia frondosa, Juncus effusus, Psilotum nudum, Araucaria heterophylla, stomatal density, stomatal index, Palaeozoic CO₂     

Comparison of Palaeobotanical Observations with Experimental Data on the Leaf Anatomy of Durmast Oak [Quercus petraea(Fagaceae)] in Response to Environmental Change

To test whether stomatal density measurements on oak leaf remainsare reliable tools for assessing palaeoatmospheric carbon dioxideconcentration [CO₂], under changing Late Miocene palaeoenvironmentalconditions, young seedings of oak (Quercus petraea,Liebl.) weregrown at elevatedvs.ambient atmospheric [CO₂] and at high humiditycombined with an increased air temperature. The leaf anatomyof the young oaks was compared with that of fossil leaves ofthe same species. In the experiments, stomatal density and stomatalindex were significantly decreased at elevated [CO₂] in comparisonto ambient [CO₂]. Elevated [CO₂] induced leaf cell expansionand reduced the intercellular air space by 35%. Leaf cell sizeor length were also stimulated at high air humidity and temperature.Regardless of a temperate or subtropical palaeoclimate, leafcell size in fossil oak was not enhanced, since neither epidermalcell density nor length of the stomatal apparatus changed. Theabsence of these effects may be attributed to the phenologicalresponse of trees to climatic changes that balanced temporalchanges in environmental variables to maintain leaf growth underoptimal and stable conditions.Quercus petraea,which evolvedunder recurring depletions in the palaeoatmospheric [CO₂], maypossess sufficient phenotypic plasticity to alter stomatal frequencyin hypostomatous leaves allowing high maximum stomatal conductanceand high assimilation rates during these phases of low [CO₂].Copyright1998 Annals of Botany Company Atmospheric CO₂, high humidity, elevated temperature,Quercus petraea,durmast oak, Late Miocene, palaeoclimates, leaf anatomy, stomatal density, stomatal index     

Elevated CO2and Temperature have Different Effects on Leaf Anatomy of Perennial Ryegrass in Spring and Summer

Mature second leaves of Lolium perenne L. cv. Vigor, were sampledin a spring and summer regrowth period. Effects of CO₂enrichmentand increased air temperature on stomatal density, stomatalindex, guard cell length, epidermal cell density, epidermalcell length and mesophyll cell area were examined for differentpositions on the leaf and seasons of growth. Leaf stomatal density was smaller in spring but greater in summerin elevated CO₂and higher in both seasons in elevated temperatureand in elevated CO₂xtemperature relative to the respective controls.In spring, leaf stomatal index was reduced in elevated CO₂butin summer it varied with position on the leaf. In elevated temperature,stomatal index in both seasons was lower at the tip/middle ofthe leaf but slightly higher at the base. In elevated CO₂xtemperature,stomatal index varied with position on the leaf and betweenseasons. Leaf epidermal cell density was higher in all treatmentsrelative to controls except in elevated CO₂(spring) and elevatedCO₂xtemperature (summer), it was reduced at the leaf base. Inall treatments, stomatal density and epidermal cell densitydeclined from leaf tip to base, whilst guard cell length showedan inverse relationship, increasing towards the base. Leaf epidermalcell length and mesophyll cell area increased in elevated CO₂inspring and decreased in summer. In elevated CO₂xtemperatureleaf epidermal cell length remained unaltered in spring comparedto the control but decreased in summer. Stomatal conductancewas lower in all treatments except in summer in elevated CO₂itwas higher than in the ambient CO₂. These contrasting responses in anatomy to elevated CO₂and temperatureprovide information that might account for differences in seasonalleaf area development observed in L. perenne under the sameconditions. Lolium perenne ; perennial ryegrass; elevated CO₂and temperature; stomatal density; stomatal index; cell size     

Stomatal Characteristics of Four Native Herbs Following Exposure to Elevated CO2

Contrasting effects on the stomatal index (SI), stomatal density,epidermal cell size and number were observed in four chalk grasslandherbs (Sanguisorba minor Scop., Lotus corniculatus L., Anthyllisvulneraria L. and Plantago media L.) following exposure to elevatedcarbon dioxide concentrations (CO₂) in controlled environmentgrowth cabinets. SI of S. minor increased for both leaf surfaces,whilst in A. vulneraria and P. media SI decreased on one surfaceonly. In L. corniculatus , no differences in SI were observedas epidermal cell density changed in parallel with stomataldensity. In L. corniculatus and S. minor stomatal density increasedon both surfaces, whereas in P. media it decreased; in A. vulnerariastomatal density decreased on the abaxial leaf surface alonefollowing exposure to elevated CO₂. In the latter three species,SI changed because stomatal density did not change in parallelwith epidermal cell density. The results suggest elevated CO₂is either directly or indirectly affecting cell differentiationand thus stomatal initiation in the meristem. In S. minor and P. media leaf growth increased in elevated CO₂,because of increased cell expansion of epidermal cells, whereasin L. corniculatus, epidermal cell size decreased and greaterleaf growth was because of an increase in epidermal cell divisions.In A. vulneraria, leaf size did not change, but increased cellexpansion on the adaxial surface suggests CO₂ affects leaf surfacesdifferently, either directly or indirectly at the cell differentiationstage or as the leaf grows. These results suggest component species of a plant communitymay differ in their response to elevated CO₂. Predicting theeffect of environmental change is therefore difficult.Copyright1994, 1999 Academic Press Elevated CO₂, Sanguisorba minor (salad burnet), Lotus corniculatus (birdsfoot trefoil), Anthyllis vulneraria (kidney vetch), Plantago media (hoary plantain), stomatal index, stomatal density, epidermal cell size     

Stomatal Density Responses of Egyptian Olea europaea L. Leaves to CO2 Change Since 1327 BC

We have attempted to separate the effects of CO₂ and temperaturechange on stomatal density by examining ancient leaf materialof Olea europaea L. The distribution of this species is confinedto a Mediterranean type climate, so that O. europaea leavesof different ages will have formed under similar temperaturesbut different CO₂ levels over the last 3000 years. Stomataldensity measurements have been made upon leaves of O. europaeaoriginating from King Tutankhamun's tomb dating from 1327 BC,and have been compared with values obtained from Egyptian O.europaea material dating from pre-332 BC, 1818 and 1978 AD.Together, the four dates provide a record of how the plant hasresponded to increases in atmospheric CO₂ concentration duringthat time. The results demonstrate that in accordance with similarstudies examining the stomatal density response of plants overthree time scales (hundreds, thousands and tens of thousandsof years) stomatal density falls as CO₂ levels increase. Sincewe have examined a natural system with leaves developing undersimilar environmental temperatures the results confirm observationsfrom experimental studies in which plants were grown under thesame temperature but different CO₂ regimes.Copyright 1993, 1999Academic Press Olea europaea, stomatal density, atmospheric CO₂, temperature, climate change     

In situ Observations of Stomatal Movements

Kappen, L., Andresen, G. and L?sch, R. 1987. In situ observationsof stomatal movements.—J. exp. Bot. 38: 126–141. A device is described by which stomatal movements in situ canbe observed and recorded continuously in light and in darkness.It is mounted in a conditioned CO₂ exchange measuring chamberso that stomatal movements can be observed whilst CO₂ exchange(photosynthesis and respiration) of the same leaf is measured.Advantages and limitations are discussed. By this method itwas shown that stomata of Vicia faba although responding inthe same direction to environmental stimuli exhibited a widerange of pore widths. Responses to changes of air humidity andof CO₂ content were clearly evident when the leaves were exposedto light. Before stomata closed due to decreasing water vapourpressure differences between leaf and air they showed a markedwidening of the pore. An inverse response occurred when watervapour pressure deficit decreased. In darkness stomata did notrespond to such changes. Key words: Stomata, leaf gas exchange, microscopic observation     

Nocturnal Accumulation of Acid in Leaves of Wall Pennywort (Umbilicus rupestris) Following Exposure to Water Stress

Physiological responses to water stress (drought) have beeninvestigated in Umbilicus rupestris (wall pennywort) by comparingcontrol (well-watered) and draughted plants with respect to(i) diurnal fluctuations in the acid content of the leaves,(ii) CO₂ exchange patterns and (iii) stomatal conductance. Controlplants show no diurnal fluctuations in acid content, whereasafter 6 d of drought a clear CAM-type pattern (nocturnal acidificationfollowed by deacidification in the light) is observed. In controlplants, the CO₂ exchange pattern over a 24 h period is of atypical C-3 ‘square-wave’ type, with extensive CO₂uptake in the light and CO₂ output in the dark. In droughtedplants the day-time CO₂ uptake is confined to a morning ‘burst’,whilst night-time CO₂ output is markedly reduced. There is howeverno net noctural uptake of CO₂. In control plants, stomatal conductanceis high during the day (especially in the first half of theday) falling to a low level at the onset of darkness, and thenrising slowly through the remainder of the night. In droughtedplants, stomatal conductance is very low, except that thereis morning ‘burst’ of high conductance and a periodduring the night when conductance is higher than in controlplants. These results are discussed in relation to the response of U.rupestris to drought both in laboratory and in field conditions. Umbilicus rupestris, wall pennywort, CO₂ exchange, Crassulacean acid metabolism, drought, stomatal conductance, water stress     

Changes in N and S Leaf Content, Stomatal Density and Specific Leaf Area of 14 Plant Species during the Last Three Centuries of CO2 Increase

Parallel to the increase in atmospheric CO² from 278 µmolmol^–1 in AD 1750 to the current ambient level of 348 µmolmol^–1, there have been overall decreases in leaf nitrogencontent and stomatal density from 144% and 121%, respectively,in AD 1750 to 100% today of herbarium specimens of 14 trees,shrubs, and herbs collected over the last 240 years in Catalonia,a Mediterranean climate area. These decreases were steeper duringthe initial slower increases in CO₂ atmospheric levels as comparedwith the relatively faster CO₂ increases in recent years. Thedeclines in leaf N content and stomatal density have also beenreported in experimental studies on leaves of plants grown underenriched CO₂ environments. Meanwhile, the stomatal index andoverall carbon and sulphur leaf contents have not changed significantly.Leaf S content was higher in the 1940s samples coinciding withthe burning of increased quantities of sulphur-rich coal. Consequently,the epidermal cell density has decreased parallel to the stomataldensity and the C/N ratio of leaves has increased, implyingpossible important consequences on herbivores, decomposers,and ecosystems. An overall decrease in the specific leaf area(SLA) from 184% in the 18th century to 100% today has also beenfound, as would be expected under CO₂ enrichment, but whichmight also be an artifact of prolonged storage. Key words: Carbon dioxide increase, leaf nitrogen content, leaf sulphur content, stomatal density, last centuries     

Very high CO2 partially restores photosynthesis in sunflower at low water potentials

We re-examined the question of whether the stomata limit photosynthesis in dehydrated sunflower (Helianthus annuus L.) plants having low leaf water potentials. A gas-exchange apparatus was modified to operate at external CO₂ partial pressures as high as 3000 Pa (3%), which were much higher than previously achieved. This allowed photosynthesis and stomatal behavior to be monitored simultaneously at very high CO₂ in the same leaf. The data were compared with those from leaves treated with abscisic acid (ABA) where effects on photosynthesis are entirely stomatal. Photosynthesis was inhibited at low water potential and was only slightly enhanced by increasing the external CO₂ partial pressure from 34 Pa (normal air) to 300 Pa. Photosynthesis in ABA-treated leaves was similarly inhibited but recovered fully at 300 Pa. In both cases, the stomata closed to the same extent as judged from the average conductance of the leaves. Because the ABA effect resulted from diffusion limitation for CO₂ caused by stomatal closure, the contrasting data show that most of the dehydration effect was nonstomatal at low water potentials. When CO₂ partial pressures were raised further to 3000 Pa, photosynthesis increased somewhat at low water potentials but not in ABA-treated leaves. This indicates that some nonstomatal component of photosynthesis responded differently in leaves at low water potential and leaves treated with ABA. Because this component was only partially restored by very high CO₂, it was likely to be metabolic and was an important source of photosynthetic inhibition.Abbreviations and Symbol ABA abscisic acid - Chl chlorophyll - p_a external partial pressure of CO₂ - P_i intercellular partial pressure of CO₂ - _w water potential This work was supported by grant DE-FG02-87ER13776 from the Department of Energy and a grant from E.I. DuPont de Nemours and Company.     

Energetic Costs of Tissue Construction in Yellow-poplar and White Oak Trees Exposed to Long-term CO2Enrichment

Two methods were used to estimate construction costs for leaves,stems, branches and woody roots of yellow-poplar (LiriodendrontulipiferaL.) trees grown at ambient (35 Pa) and elevated (65Pa) CO₂for 2.7 years and trees of white oak (Quercus albaL.)grown at these same CO₂partial pressures for 4 years. Samplecombustion in a bomb calorimeter combined with measurementsof ash and nitrogen content provided the primary method of estimatingtissue construction costs (W_G; g glucose g^-1dry mass). Thesevalues were compared with a second, simpler method in whichcost estimates were derived from tissue ash, carbon and nitrogencontent (V_G). Estimates of W_Gwere lower for leaves, branchesand roots of yellow-poplar and for leaves of white oak grownat elevated compared with ambient CO₂partial pressures. TheseCO₂-induced differences in W_Granged from 3.7% in yellow-poplarroots to 2.1% in white oak leaves. Only in the case of yellow-poplarleaves, however, were differences in V_Gobserved between CO₂treatments.Leaf V_Gwas 1.46 g glucose g^-1dry mass in ambient-grown treescompared with 1.41 g glucose g^-1dry mass for CO₂-enriched trees.Although paired-estimates of W_Gand V_Gclustered about a 1:1 linefor leaves and branches, estimates of V_Gwere consistently lowerthan W_Gfor stems and roots. Construction costs per unit leafarea were 95 g glucose m^-2for yellow-poplar trees grown at ambientCO₂and 106 g glucose m^-2for trees grown at elevated CO₂partialpressures. No differences in area-based construction costs wereobserved for white oak. Whole-plant energy content was 1220g glucose per tree in ambient-grown white oak compared with2840 g glucose per tree for those grown at elevated CO₂partialpressures. These differences were driven largely by CO₂-inducedchanges in total biomass. We conclude that while constructioncosts were lower at elevated CO₂partial pressures, the magnitudeof this response argues against an increased efficiency of carbonuse in the growth processes of trees exposed to CO₂enrichment. Bomb calorimeter; construction costs; elevated CO₂; energy allocation; global change; growth respiration; heat of combustion; respiration; Liriodendron tulipifera; Quercus alba     

The Responses of Net Photosynthesis to Vapour Pressure Deficit and CO2 Concentration in Spartina townsendii (sensu lato), a C4 Species from a Cool Temperate Climate

As the initial part of a detailed study of photosynthetic CO₂assimilation in the temperate C₄ grass, Spartina townsendii,the responses of net photosynthesis to the leaf-air vapour pressuredeficit (VPD) and to CO₂ concentration are examined. Water vapourand CO₂ exchange for single attached leaves were measured undercontrolled-environment conditions in an open gas-exchange system.The responses of net photosynthesis, stomatal resistance (r_s),and residual resistance (r_r) to vapour pressure deficit(VPD)and CO₂ concentration under a range of light and temperatureconditions are reported. Net photosynthesis was insensitiveto increase in the VPD up to 1.0 kPa, but beyond this valuenet photosynthesis decreased with further increase in VPD asa result of an increase in r_s. The residual resistance was notaffected by VPD under any of the conditions examined. Net photosynthesisresponded linearly to increase in the CO₂ concentration in theexternal air (C_a up to the normal atmospheric concentrationwhere there was a sharp change in the response, net photosynthesisbeing independent of any further increase in C_a. Differencesbetween the response curves observed here and in other studiesare discussed and the possible reasons for these differencesare considered.     

The effect of elevated atmospheric CO2 and drought onstomatal frequency in groundnut (Arachis hypogaea (L.))

The effects of elevated atmospheric CO₂, alone or in combinationwith water stress, on stomatal frequency in groundnut (Arachishypogaea (L.) cv. Kadiri-3) were investigated. CO₂ exerted significanteffects on stomatal frequency only in irrigated plants. Theeffects of drought on leaf development out weighed the smallereffects of CO₂ concentration, although reductions in stomatalfrequency induced by elevated atmo-spheric CO₂ were still observed.When stands of groundnut were grown under irrigated conditionswith unrestricted root systems, an increase in atmospheric CO₂from 375 to 700 ppmv decreased stomatal frequency on both leafsurfaces by up to 16% in droughted plants, stomatal frequencywas reduced by 8% on the adaxial leaf surface only. Elevatedatmospheric CO₂ promoted larger reductions in leaf conductancethan the changes in stomatal frequency, indicating partial stomatalclosure. As a result, the groundnut stands grown at elevatedCO₂ utilized the available soil moisture more slowly than thosegrown under ambient CO₂, there by extending the growing period.Despite the large variations in cell frequencies induced bydrought, there was no treatment effect on either stomatal indexor the adaxial/abaxial stomatalfrequency ratio. The data suggestthat the effects of future increases in atmospheric CO₂ concentrationon stomatal frequency in groundnut are likely to be small, especiallyunder conditions of water stress, but that the combination ofassociated reductions in leaf con-ductance and enhanced assimilationat elevated CO₂ will be important in semi-arid regions Key words: Arachis hypogaea L, Leguminosae, groundnu, stomatal frequency, CO₂, drought     

Effects of Different CO2 Environments on the Photosynthesis-Yield Relationship and the Carbon and Water Balance of a White Clover (Trifolium repens L. cv. Blanca) Sward

Effects of atmospheric CO₂ enrichment to a level above 600 parts10^–6 on leaf and canopy gas exchange characteristics wereinvestigated in Trifolium repens, using an open system for gasexchange measurement. The cuvettes of the system served as growthchambers, allowing continuous measurement in a semi-controlledenvironment of ±350 and ±600 parts 10^–6CO₂, respectively. Carbon balance data were compared with cropyield and effects on the canopy level were compared with measuredleaf responses of photosynthesis and stomatal behaviour. Photosyntheticstimulation by high CO₂ was stronger at the canopy level (103%on average) than for leaves (90% in full light), as a consequenceof accelerated foliage area development. The latter increasedabsolute water consumption by 16%, despite strong stomatal closure.The overall result was a 63% improvement in canopy water useefficiency (WUE), while leaf WVE increased almost 3-fold insaturating light. The stomatal response was such that, whilethe internal CO₂ concentration in the leaf, ch increased withrising atmospherical CO₂ concentration, c_a, ci/c_a was somewhatdecreased. Total canopy resistance, R_c, was generally lowerat high CO₂ levels, despite higher leaf resistance. Higher canopyCO₂ loss at night and faster light extinction in a larger-sizedhigh CO₂ canopy were major drawbacks which prevented a furtherincrease in dry matter production (the harvest index was increasedby a factor 1.83). Key words: CO₂ enrichment, canopy CO₂ exchange, carbon balance, water use efficiency, leaf and canopy resistance