Effect of elevated CO2 and nutrient status on growth, dry matter partitioning and nutrient content of Poa alpina var. vivipara L.

Poa alpina var. vivipara L. was grown in an atmosphere containingeither 340 or 680 µmol CO₂ mol^–1 within controlledenvironment chambers. The available nutrient regime was variedby altering the supply of nitrogen and phosphorus within a completenutrient solution. At a high, but not low, N and P supply regime,elevated CO₂ markedly increased growth. Differences betweennutrient supply, but not atmospheric CO₂ concentration, alteredthe allometric relations between root and shoot. Net photosynthesisof mature leaf blades and leaf N and P concentration were reducedin plants grown at the elevated CO₂ concentration. The question was asked: is it possible to ascribe all of theseeffects to elevated CO₂ or are some due to nutrient deficiencycaused by dilution with excess carbon? Several criteria, includingthe nutrient content of sink tissue, root:shoot allometry andthe use of divalent cations to estimate integrated water flowsare suggested in order to make this distinction. It is concludedthat only at a low supply of N and P₁ and elevated CO₂ concentration,was low leaf N concentration due to induced nutrient deficiency.The data are consistent with a model where the capacity of sinksto use photosynthetically assimilated carbon sets both the rateof import into those sinks (and thus rate of export from sourceleaves) and the rate of photosynthesis of source leaves themselves. Key words: Poa alpina L., growth, photosynthesis, carbohydrate, export, nitrogen, phosphorus     

Effects of elevated CO2 concentrations on three montane grass species: III. Source leaf metabolism and whole plant carbon partitioning

Agrostis capillaris L.⁵, Festuca vivipara L. and Poaalpina L.were grown in outdoor open-top chambers at either ambient (340 3µmol mol^–1) or elevated (6804µmol mol^–1)concentrations of atmospheric carbon dioxide (CO₂) for periodsfrom 79–189 d. Photosynthetic capacity of source leaves of plants grown atboth ambient and elevated CO₂ concentrations was measured atsaturating light and 5% CO₂. Dark respiration of leaves wasmeasured using a liquid phase oxygen electrode with the buffersolution in equilibrium with air (21% O₂, 0.034% CO₂). Photo-syntheticcapacity of P. alpina was reduced by growth at 680 µmolmol^–1 CO₂ by 105 d, and that of F. vivipara was reducedat 65 d and 189 d after CO₂ enrichment began, suggesting down-regulationor acclimation. Dark respiration of successive leaf blades ofall three species was unaltered by growth at 680 relative to340 µmol mol^–1 CO₂. In F. vivipara, leaf respirationrate was markedly lower at 189 d than at either 0 d or 65 d,irrespective of growth CO₂ concentration. There was a significantlylower total non-structural carbohydrate (TNC) concentrationin the leaf blades and leaf sheaths of A. capillaris grown at680µmol mol^–1 CO₂. TNC of roots of A. capillariswas unaltered by CO₂ treatment. TNC concentration was increasedin both leaves and sheaths of P. alpina and F. vivipara after105 d and 65 d growth, respectively. A 4-fold increase in thewater-soluble fraction (fructan) in P. alpina and in all carbohydratefractions in F. vivipara accounted for the increased TNC content. In F. vivipara the relationship between leaf photosyn-theticcapacity and leaf carbohydrate concentration was such that therewas a strong positive correlation between photosynthetic capacityand total leaf N concentration (expressed on a per unit structuraldry weight basis), and total nitrogen concentration of successivemature leaves reduced with time. Multiple regression of leafphotosynthetic capacity upon leaf nitrogen and carbohydrateconcentrations further confirmed that leaf photosynthetic capacitywas mainly determined by leaf N concentration. In P. alpina,leaf photosynthetic capacity was mainly determined by leaf CHOconcentration. Thus there is evidence for down-regulation ofphotosynthetic capacity in P. alpina resulting from increasedcarbohydrate accumulation in source leaves. Leaf dark respiration and total N concentration were positivelycorrelated in P. alpina and F. vivipara. Leaf dark respirationand soluble carbohydrate concentration of source leaves werepositively correlated in A. capillaris. Changes in source leafphotosynthetic capacity and carbohydrate concentration of plantsgrown at ambient or elevated CO₂ are discussed in relation toplant growth, nutrient relations and availability of sinks forcarbon. Key words: Elevated CO₂, Climate change, grasses, carbohydrate partitioning, photosynthesis, respiration     

Is Partitioning of Dry Weight and Leaf Area Within Dactylis glomerata Affected by N and CO2Enrichment?

We examined changes in dry weight and leaf area within Dactylisglomerata L. plants using allometric analysis to determine whetherobserved patterns were truly affected by [CO₂] and N supplyor merely reflect ontogenetic drift. Plants were grown hydroponicallyat four concentrations of in controlled environment cabinets at ambient (360 µll^–1) or elevated (680 µl l^–1) atmospheric[CO₂]. Both CO₂and N enrichment stimulated net dry matter production.Allometric analyses revealed that [CO₂] did not affect partitioningof dry matter between shoot and root at high N supply. However,at low N supply there was a transient increase in dry matterpartitioning into the shoot at elevated compared to ambient[CO₂] during early stages of growth, which is inconsistent withpredictions based on optimal partitioning theory. In contrast,dry matter partitioning was affected by N supply throughoutontogeny, such that at low N supply dry matter was preferentiallyallocated to roots, which is in agreement with optimal partitioningtheory. Independent of N supply, atmospheric CO₂enrichment resultedin a reduction in leaf area ratio (LAR), solely due to a decreasein specific leaf area (SLA), when plants of the same age werecompared. However, [CO₂] did not affect allometric coefficientsrelating dry weight and leaf area, and effects of elevated [CO₂]on LAR and SLA were the result of an early, transient stimulationof whole plant and leaf dry weight, compared to leaf area production.We conclude that elevated [CO₂], in contrast to N supply, changesallocation patterns only transiently during early stages ofgrowth, if at all. Copyright 2000 Annals of Botany Company Allometric growth, carbon dioxide enrichment, Cocksfoot, Dactylis glomerata L., dry weight partitioning, leaf area ratio, nitrogen supply, shoot:root ratio, specific leaf area     

Effects of CO2 enrichment and intraspecific competition on biomass partitioning, nitrogen content and microbial biomass carbon in soil of perennial ryegrass and white clover

Seedlings of perennial ryegrass (Lolium perenne L. cv. Parcour)and white clover (Trifolium repens L. cv. Karina) grown at fivedifferent plant densities were exposed to ambient (390 ppm)and elevated (690 ppm) CO₂ concentrations. After 43 d the effectsof CO₂ enrichment and plant density on growth of shoot and root,nitrogen concentration of tissue, and microbial biomass carbon(C_mic) in soil were determined. CO₂ enrichment of Lolium perenneincreased shoot growth on average by 17% independent of plantdensity, while effects on root biomass ranged between -4% and+ 107% due to an interaction with plant density. Since tilernumber per plant was unaffected by elevated CO₂, the small responseof shoot growth to CO₂ enrichment was atributed to low sinkstrength. A significant correlation between nitrogen concentrationof total plant biomass and root fraction of total plant drymatter, which was not changed by CO₂ enrichment, indicates thatnitrogen status of the plant controls biomass partitioning andthe effect of CO₂ enrichment on root growth. Effects of elevatedCO₂ and plant density on shoot and root growth of Trifoliumrepens were not significantly interacting and mean CO₂-relatedincrease amounted to 29% and 66%, respectively. However, growthenhancement due to elevated CO₂ was strongest when leaf areaindex was lowest. Total amounts of nitrogen in shoots and rootswere bigger at 690 ppm than at 390 ppm CO₂. There was a significantincrease in C_mic in experiments with both species whereas plantdensity had no substantial effect. Key words: CO₂ enrichment, intraspecific competition, biomass partitioning, Lolium perenne, Trifolium repens, grassland     

Growth and Physiology of One-year old Poplar (Populus) Under Elevated Atmospheric CO2 Levels

The effects of elevated atmospheric CO₂ concentrations on theecophysiological responses (gas exchange, chlorophyll a fluorescence,Rubisco activity, leaf area development) as well as on the growthand biomass production of two poplar clones (i.e. Populus trichocarpax P. deltoides clone Beaupré and P. x euramericana cloneRobusta) were examined under open top chamber conditions. Theelevated CO₂ treatment (ambient + 350 µmol mol^-1) stimulatedabove-ground biomass of clones Robusta and Beaupré afterthe first growing season by 55 and 38%, respectively. This increasedbiomass production under elevated CO₂ was associated with asignificant increase in plant height, the latter being the resultof enhanced internode elongation rather than an increased productionof leaves or internodes. Both an increased leaf area index (LAI)and a stimulated net photosynthesis per unit leaf contributedto a significantly higher stem biomass per unit leaf area, andthus to the increased above-ground biomass production underthe elevated CO₂ concentrations in both clones. The larger LAIwas caused by a larger individual leaf size and leaf growthrate; the number of leaves was not altered by the elevated CO₂treatment. The higher net leaf photosynthesis was the resultof an increase in the photochemical (maximal chlorophyll fluorescenceFm and photochemical efficiency Fv/Fm) as well as in the biochemical(increased Rubisco activity) process capacities. No significantdifferences were found in dark respiration rate, neither betweenclones nor between treatments, but specific leaf area significantlydecreased under elevated CO₂ conditions.Copyright 1995, 1999Academic Press Biomass, chlorophyll a fluorescence, elevated CO₂, growth, Populus, poplar, photosynthesis, respiration, Rubisco     

Effects of Elevated Carbon Dioxide on Plant Biomass Production and Competition in a Simulated Neutral Grassland Community

Using open-top chambers, four prominent species (Lolium perenne,Cynosurus cristatus, Holcus lanatusandAgrostis capillaris) ofIrish neutral grasslands were grown at ambient and elevated(700 µmol mol^-1) atmospheric CO₂for a period of 8 months.The effects of interspecific competition on plant responsesto CO₂enrichment were investigated by growing the species ina four-species mixture. The results indicate that the speciesdiffer in their ability to respond to elevated CO₂. CO₂-enrichmenthad the largest effect on the biomass production ofH. lanatus,but substantial stimulations in biomass production were alsofound for the other three species. The CO₂-stimulation of biomassproduction forH. lanatuswas accompanied by increased tillering.In addition, reductions in specific leaf area were found forall species. Exposure to elevated CO₂increased the communitybiomass of the four-species mixture. This increase can be mainlyattributed to a significant increase in the biomass ofH. lanatusatelevated CO₂. No statistically-significant changes in speciescomposition of community biomass were found. However,H. lanatusdidincrease its share of community biomass at each of the harvests,with the other three species, mainlyL. perenne, suffering lossesin their shares at elevated CO₂. The results show that: (1)the species varied in their response to elevated CO₂; and (2)species composition in natural plant communities is likely tochange at elevated CO₂, but these changes may occur rather slowly.Much longer periods of exposure to elevated atmospheric CO₂maybe required to permit detection of significant changes in speciescomposition.Copyright 1998 Annals of Botany Company Carbon dioxide (CO₂) enrichment, competition, Lolium perenne,Cynosurus cristatus, Holcus lanatus, Agrostis capillaris, biomass, specific leaf area, tillering.     

Effect of Elevated CO2 on the Photosynthesis, Respiration and Growth of Perennial Ryegrass

Single, seed-grown plants of ryegrass (Lolium perenne L. cv.Melle) were grown for 49 d from the early seedling stage ingrowth cabinets at a day/night temperature of 20/15 C, witha 12 h photoperiod, and a CO₂ concentration of either 340 or680µI 1^–1 CO₂. Following complete acclimation tothe environmental regimes, leaf and whole plant CO₂ effluxesand influxes were measured using infra-red gas analysis techniques.Elevated CO₂ increased rates of photosynthesis of young, fullyexpanded leaves by 35–46% and of whole plants by morethan 50%. For both leaves and whole plants acclimation to 680µI^–1 CO₂ reduced rates of photosynthesis in bothCO₂ regimes, compared with plants acclimated to 340µll^–1. There was no significant effect of CO₂ regime onrespiration rates of either leaves or whole plants, althoughleaves developed in elevated CO₂ exhibited generally lower ratesthan those developed in 340µI I^–1 CO₂. Initially the seedling plants in elevated CO₂ grew faster thantheir counterparts in 340µI I^–1 CO₂, but this effectquickly petered out and final plant weights differed by onlyc. 10%. Since the total area of expanded and unexpanded laminaewas unaffected by CO₂ regime, specific leaf area was persistently13–40% lower in elevated CO₂ while, similarly, root/shootratio was also reduced throughout the experiment. Elevated CO₂reduced tissue nitrogen contents of expanded leaves, but hadno effect on the nitrogen contents of unexpanded leaves, sheathsor roots. The lack of a pronounced effect of elevated CO₂ on plant growthwas primarily due to the fact that CO₂ concentration did notinfluence tiller (branch) numbers. In the absence of an effecton tiller numbers, any possible weight increment was restrictedto the c. 2.5 leaves of each tiller. The reason for the lackof an effect on tillering is not known. Key words: Lolium perenne, ryegrass, elevated CO₂, photosynthesis, respiration, growth, development     

Response of young barley plants to CO2 enrichment

Barley (Hordeum vulgare L. cv. Digger) was grown for 22 d inenclosed chambers with a CO₂ enrichment of 35, 155, 400 or 675µmol CO₂ mol1. CO₂ enrichment increased photosyntheticcapacity in the plants grown at either of the two highest levelsof pCO₂. A CO₂ enrichment of 675µmol CO₂ caused a significantincrement of shoot dry weight, whereas no changes were observedin fresh weight, chlorophyll or protein levels. At a light intensityof 860µmol m^–2s^–1 CO₂ enrichment caused photosyntheticcapacity to increase by 250%, whereas no effect was observedat 80 µmol m^–2 s^–1. Over time, photosynthesisdecreased by 70% independent of CO₂. A time-dependent increasein the level of extractable fructose was observed whereas totalextractable carbohydrate only changed slightly. Key words: Carbohydrates, CO₂ enrichment, Hordeum vulgare, photosynthesis, respiration     

Effects of elevated carbon dioxide on three montane grass species: I. Growth and dry matter partitioning

Upland grasslands are a major component of natural vegetationwithin the UK. Such grasslands support slow growing relativelystable plant communities. The response of native montane grassspecies to elevated atmospheric carbon dioxide concentrationshas received little attention to date. Of such studies, mosthave only focused on short-term (days to weeks) responses, oftenunder favourable controlled environment conditions. In thisstudy Agrostis caplllaris L.⁵, Festuca vivipara L. and Poa alpinaL. were grown under semi-natural conditions in outdoor open-topchambers at either ambient (340µmol mol^–1) or elevated(680µmol mol^–1) concentrations of atmospheric carbondioxide (CO₂ for periods from 79 to 189 d, with a nutrient availabilitysimilar to that of montane Agrostis-Fescue grassland in Snowdonia,N. Wales. Whole plant dry weight was increased for A. capillarisand P. alpina, but decreased for F. vivipara, at elevated CO₂.Major components of relative growth rate (RGR) contributingto this change at elevated CO₂ were transient changes in specificleaf area (SLA) and leaf area ratio (LAR). Despite changes ingrowth rate at 680 µmol mol^–1 CO₂, partitioningof dry weight between shoot and root in plants of A. capillarisand P. alpina was unaltered. There was a significant decreasein shoot relative to root growth at elevated CO₂ in F. viviparawhich also showed marked discoloration of the leaves and increasedsenescence of the foliage. Key words: Allometry, growth analysis, elevated CO₂, grasses     

The Greenhouse Effect: Acclimation of Tomato Plants Growing in High CO2, Photosynthesis and Ribulose-1, 5-Bisphosphate Carboxylase Protein

Tomato plants were grown in solution culture in a controlledenvironment at 20 ?C with a 12 h photoperiod of 400 µmolquanta m^–2 s^–1 PAR with either normal ambient CO₂,approximately 340 vpm, or with 1000 vpm CO₂. The short- andlong-term effects of CO₂ enrichment on photosynthesis were determinedtogether with the levels of ribulose-1, 5-bisphosphate carboxylase(RuBPco) E.C. 4.1.1.39 [EC] protein and activity throughout leafdevelopment of the unshaded 5th leaf above the cotyledons. Thehigh CO₂ concentration during growth did not appreciably affectthe rate of leaf expansion or final leaf area but did increasethe fresh weight per unit area of leaf. With short-term CO₂enrichment, i.e. only during the photosynthesis measurements,the light-saturated photosynthetic rate (P_max) of young leavesdid not increase while those reaching full expansion more thandoubled their net rate of CO₂ fixation. However, with longerterm CO₂ enrichment, i.e. growing the crop in high CO₂, theplants did not maintain this photosynthetic gain. While theCO₂ concentration during growth did not affect the peak in P_maxmeasured in 300 vpm CO₂ or P_max in 1000 vpm CO₂, RuBPco proteinor its activity, the subsequent ontogenetic decline in theseparameters was greatly accelerated by the high CO₂ treatment.Compared with plants grown in normal ambient CO₂ the high CO₂grown leaves, when almost fully expanded, contained only approximatelyhalf as much RuBPco protein and P_max in 300 vpm CO₂ and P_maxin1000 vpm CO₂ were similarly reduced. The loss of RuBPco proteinmay be a major factor associated with the accelerated fall inP_max since it was close to that predicted from the amount andkinetics of RuBPco assuming RuBP saturation. In the oldest leavesexamined grown in high CO₂ additional factors may be limitingphotosynthesis since RuBPco kinetics marginally overestimatedP_max in 300 vpm CO₂ and the initial slope of photosynthesisin response to intercellular CO₂ was also less than expectedfrom the extractable RuBPco. Key words: Lycopersicon esculentum (Mill.) cv. Findon Cross, CO₂ enrichment, acclimation to high CO₂, photosynthesis, RuBPco protein and activity     

Photosynthetic acclimation to elevated CO2 in poplar grown in glasshouse cabinets or in open top chambers depends on duration of exposure

The effects of elevated CO₂ were studied on the photosyntheticgas exchange behaviour and leaf physiology of two contrastingpoplar (Populus) hybrids grown and treated in open top chambers(OTCs in Antwerp, Belgium) and in closed glasshouse cabinets(GHCs in Sussex, UK). The CO₂ concentrations used in the OTCswere ambient and ambient +350 µmol mol^–1 while inthe GHCs they were c. 360 µmol mol^–1 versus 719µmol mol^–1. Measurements of photosynthetic gas exchangewere made for euramerican and interamerican poplar hybrids incombination with measurements of dark respiration rate and Rubiscoactivity. Significant differences in the leaf anatomy and structure(leaf mass per area and chlorophyll content) were observed betweenthe leaves grown in the OTCs and those grown in the GHCs. ElevatedCO₂ stimulated net photosynthesis in the poplar hybrids after1 month in the GHCs and after 4 months in the OTCs, and therewas no evidence of downward acclimation (or down-regulation)of photosynthesis when the plants in the two treatments weremeasured in their growth CO₂ concentration. There was also noevidence of down-regulation of Rubisco activity and there wereeven examples of increases in Rubisco activity. Rubisco exerteda strong control over the light-saturated rate of photosynthesis,which was demonstrated by the close agreement between observednet photosynthetic rates and those that were predicted fromRubisco activities and Michaelis-Menten kinetics. After 17 monthsin elevated CO₂ in the OTCs there was a significant loss ofRubisco activity for one of the hybrid clones, i.e. Beaupr,but not for clone Robusta. The effect of the CO₂ measurementconcentration (i.e. the short-term treatment effect) on netphotosynthesis was always larger than the effect of the growthconcentration in both the OTCs or GHCs (i.e. the longterm growthCO₂ effect), with one exception. For the interamerican hybridBeaupr dark respiration rates in the OTCs were not significantlyaffected by the elevated CO₂ concentrations. The results suggestthat for rapidly growing tree species, such as poplars, thereis little evidence for downward acclimation of photosynthesiswhen plants are exposed to elevated CO₂ for up to 4 months;longer term exposure reveals loss of Rubisco activity. Key words: Elevated CO₂, Populus, Rubisco, photosynthesis, chlorophyll content     

Effects of elevated carbon dioxide on three grass species from montane pasture II. Nutrient uptake, allocation and efficiency of use

Agrostis capillaris L.⁴ Festuca vivipara L. and Poa alpinaL.were grown in outdoor open-top chambers at either ambient (340µmol mol^–1) or elevated (680 µmol^–1)CO² for periods from 79 to 189 d. Under these conditions thereis increased growth of A. caplllarls and P. alpina, but reducedgrowth of F. vivipara. Nutrient use efficiency, nutrient productivity(total plant dry weight gain per unit of nutrient) and nutrientallocation of all three grass species were measured in an attemptto understand their individual growth responses further andto determine whether altered nutrient-use efficiencies and productivitiesenable plants exposed to an elevated atmospheric CO₂ environmentto overcome potential limitations to growth imposed by soilfertility. Total uptake of nutrients was, in general, greater in plantsof A. capillaris and P. alpina (with the exception of N andK in the latter) when grown at 680 µmol mol^–1 CO₂.In F. vivipara, however, uptake was considerably reduced inplants grown at the higher CO₂ concentration. Overall, a doubling of atmospheric CO₂ concentration had littleeffect on the nutrient use efficiency or productivity of A.capillaris. Reductions in tissue nutrient content resulted fromincreased plant growth and not altered nutrient use efficiency.In P. alpina, potassium, magnesium and calcium productivitieswere significantly reduced and photosynthetic nitrogen and phosphorususe efficiencies were doubled at elevated CO₂ with respect toplants grown at ambient CO₂ F. vivipara grown for 189 d showedthe most marked changes in nutrient use efficiency and nutrientproductivity (on an extracted dry weight basis) when grown atelevated CO₂, F. vivipara grown at elevated CO₂ however, showedlarge increases in the ratio of non-structural carbohydrateto nitrogen content of leaves and reproductive tissues, indicatinga substantial imbalance between the production and utilizationof assimilate. Key words: Nutrient, allocation, nutrient use efficiency, grasses, nutrient productivity, elevated CO₂, cliniate change     

The Effects of CO2 Enrichment and Nutrient Supply on Growth Morphology and Anatomy of Phaseolus vulgaris L. Seedlings

Plants of Phaseolus vulgaris were grown from seed in open-topgrowth chambers at the present (P, 350 µmol mol^–1)atmospheric CO₂ concentration and at an elevated (E, 700 µmolmol^–1) CO₂ concentration, and at low (L, without additionalnutrient solution) and high (H, with additional nutrient solution)nutrient supply for 28 d The effects of CO₂ and nutrient availabilitywere examined on growth, morphological and biochemical characteristics Leaf area and dry mass were significantly increased by CO₂ enrichmentand by high nutrient supply Stomatal density, stomatal indexand epidermal cell density were not affected by elevated CO₂concentration or by nutrient supply Leaf thickness respondedpositively to CO₂ increasing particularly in mesophyll areaas a result of cell enlargement Intercellular air spaces inthe mesophyll decreased slightly in plants grown in elevatedCO₂ Leaf chlorophyll content per unit area or dry mass was significantlylower in elevated CO₂ grown plants and increased significantlywith increasing nutrient availability The content of reducingcarbohydrates of leaves, stem, and roots was not affected byCO₂ but was significantly increased by nutrient addition inall plant parts Starch content in leaves and stem was significantlyincreased by elevated CO₂ concentration and by high nutrientsupply Phaseolus vulgaris, elevated atmospheric CO₂, CO₂-nutrient interaction, stomatal density, leaf anatomy, chlorophyll, carbohydrates, starch     

Plant responses to atmospheric carbon dioxide enrichment: interactions with some soil and atmospheric conditions

In general, C₃ plant species are more responsive to atmospheric carbon dioxide (CO₂) enrichment than C₄-plants. Increased relative growth rate at elevated CO₂ primarily relates to increased Net Assimilation Rate (NAR), and enhancement of net photosynthesis and reduced photorespiration. Transpiration and stomatal conductance decrease with elevated CO₂, water use efficiency and shoot water potential increase, particularly in plants grown at high soil salinity. Leaf area per plant and leaf area per leaf may increase in an early growth stage with increased CO₂, after a period of time Leaf Area Ratio (LAR) and Specific Leaf Area (SLA) generally decrease. Starch may accumulate with time in leaves grown at elevated CO₂. Plants grown under salt stress with increased (dark) respiration as a sink for photosynthates, may not show such acclimation to increased atmospheric CO₂ levels. Plant growth may be stimulated by atmospheric carbon dioxide enrichment and reduced by enhanced UV-B radiation but the limited data available on the effect of combined elevated CO₂ and ultraviolet B (280–320 nm) (UV-B) radiation allow no general conclusion. CO₂-induced increase of growth rate can be markedly modified at elevated UV-B radiation. Plant responses to elevated atmospheric CO₂ and other environmental factors such as soil salinity and UV-B tend to be species-specific, because plant species differ in sensitivity to salinity and UV-B radiation, as well as to other environmental stress factors (drought, nutrient deficiency). Therefore, the effects of joint elevated atmospheric CO₂ and increased soil salinity or elevated CO₂ and enhanced UV-B to plants are physiologically complex.     

CO2浓度增加和不同氮肥水平对冬小麦根系呼吸及生物量的影响

 依托FACE(Free-air CO₂ enrichment)研究平台, 利用特制分根集气生长箱, 采用静态箱-GC(Gas chromatography)法, 连续两年研究 了大气CO₂浓度升高和不同氮肥水平对冬小麦拔节期、孕穗抽穗期和灌浆末期的根系呼吸及生物量的影响。两季结果表明, CO₂浓度升高和高氮 肥量均不同程度地增加了3个阶段的地上部和地下部的生物量, 这有利于增加根茬的还田量; CO₂浓度升高对冬小麦不同生长阶段的根系呼吸影 响不同, 在拔节期影响较小;孕穗抽穗期显著增加了根系呼吸, 2004~2005季分别增加33.8%(148.1 mg N&;#8226;kg^-1 干土, HN)和43.9%(88.9 mg N&;#8226;kg^-1 干土, LN), 2005~2006季分别为23.8%(HN)和28.9%(LN); 而灌浆末期显著降低了根系呼吸, 2004~2005季分别降低31.4%(HN)和23.3% (LN), 2005~2006季分别为25.1%(HN)和18.5%(LN); 高施氮量比低施氮量促进了根系呼吸; 随着作物生长根系呼吸与地下生物量呈显著线性负相 关, 高CO₂环境中的R²变小,表明随着作物生长发育高CO₂浓度降低了作物根系呼吸与地下部生物量积累间的相关性.     

Impact of elevated atmospheric CO2 and O3 on gas exchange and chlorophyll content in spring wheat (Triticum aestivum L.)

Stands of spring wheat grown in open-top chambers (OTCs) wereused to assess the individual and interactive effects of season-longexposure to elevated atmospheric carbon dioxide (CO₂ and ozone(O₃) on the photosynthetic and gas exchange properties of leavesof differing age and position within the canopy. The observedeffects were related to estimated ozone fluxes to individualleaves. Foliar chlorophyll content was unaffected by elevatedCO₂ but photosynthesis under saturating irradiances was increasedby up to 100% at 680 µmol mol^–1 CO₂ relative tothe ambient CO₂ control; instantaneous water use efficiencywas improved by a combination of increased photosynthesis andreduced transpiration. Exposure to a seasonal mean O₃ concentration(7 h d^–1) of 84 nmol mol^–1 under ambient CO₂ acceleratedleaf senescence following full expansion, at which time chlorophyllcontent was unaffected. Stomatal regulation of pollutant uptakewas limited since estimated O₃ fluxes to individual leaves werenot reduced by elevated atmospheric CO₂, A common feature ofO₃-treated leaves under ambient CO₂ was an initial stimulationof photosynthesis and stomatal conductance for up to 4 d and10 d, respectively, after full leaf expansion, but thereafterboth variables declined rapidly. The O₃-induced decline in chlorophyllcontent was less rapid under elevated CO₂ and photosynthesiswas increased relative to the ambient CO₂ treatment. A/C_i analysessuggested that an increase in the amount of in vivo active RuBisCOmay be involved in mitigating O₃-induced damage to leaves. Theresults obtained suggest that elevated atmospheric CO₂ has animportant role in restricting the damaging effects of O₃ onphotosynthetic activity during the vegetative growth of springwheat, and that additional direct effects on reproductive developmentwere responsible for the substantial reductions in grain yieldobtained at final harvest, against which elevated CO₂ providedlittle or no protection. Key words: Elevated CO₂ and O₃, gas exchange, O₃ flux, stomata, chlorophyll, Triticum aestivum     

Influence of Elevated CO2 and Temperature on the Photosynthesis and Respiration of White Clover Dependent on N2 Fixation

Single clonal plants of white clover (Trifolium repens L) grownfrom explants in a Perlite rooting medium, and dependent fornitrogen on N₂ fixation in root nodules, were grown for severalweeks in controlled environments which provided two regimesof CO₂, and temperature 23/18 °C day/night temperaturesat 680 µmol mol^–1 CO₂, (C680), and 20/15 °Cday/night temperatures at 340 µmol mol^–1 CO₂ (C340)After 3–4 weeks of growth, when the plants were acclimatedto the environmental regimes, leaf and whole-plant photosynthesisand respiration were measured using conventional infra-red gasanalysis techniques Elevated CO₂ and temperature increased ratesof photosynthesis of young, fully expanded leaves at the growthirradiance by 17–29%, despite decreased stomatal conductancesand transpiration rates Water use efficiency (mol CO₂ mol H₂O^–1)was also significantly increased Plants acclimated to elevatedCO₂, and temperature exhibited rates of leaf photosynthesisvery similar to those of C340 leaves ‘instantaneously’exposed to the C680 regime However, leaves developed in theC680 regime photosynthesised less rapidly than C340 leaves whenboth were exposed to a normal CO₂, and temperature environmentIn measurements where irradiance was varied, the enhancementof photosynthesis in elevated CO₂ at 23 °C increased graduallyfrom approx 10 % at 100 µmol m^–1 s^–1 to >27 % at 1170 µmol m^–2 s^–1 In parallel, wateruse efficiency increased by 20–40 % at 315 µmolm^–2 s^–1 In parallel, water use efficiency increasedby 20–40 % at 315 µmol m^–2 s^–1 In parallel,water use efficiency increased by 20–40 % at 315 µmolm^–2 s^–1 In parallel, water use efficiency increasedby 20–40 % at 315 µmol m^–2 s^–1 to approx100 % at the highest irradiance Elevated CO₂, and temperatureincreased whole-plant photosynthesis by > 40 %, when expressedin terms of shoot surface area or shoot weight No effects ofelevated CO₂ and temperature on rate of tissue respiration,either during growth or measurement, were established for singleleaves or for whole plants Dependence on N₂, fixation in rootnodules appeared to have no detrimental effect on photosyntheticperformance in elevated CO₂, and temperature Trifolium repens, white clover, photosynthesis, respiration, elevated CO₂, elevated temperature, water use efficiency, N₂ fixation     

Growth at elevated CO2 leads to down-regulation of photosynthesis and altered response to high temperature in Quercus suber L. seedlings

The effects of growth at elevated CO₂ on the response to hightemperatures in terms of carbon assimilation (net photosynthesis,stomatal conductance, amount and activity of Rubisco, and concentrationsof total soluble sugars and starch) and of photochemistry (forexample, the efficiency of excitation energy captured by openphotosystem II reaction centres) were studied in cork oak (Quercussuber L.). Plants grown in elevated CO₂ (700 ppm) showed a down-regulationof photosynthesis and had lower amounts and activity of Rubiscothan plants grown at ambient CO₂ (350 ppm), after 14 monthsin the greenhouse. At that time plants were subjected to a heat-shocktreatment (4 h at 45C in a chamber with 80% relative humidityand 800–1000 mol m^–2 s^–1 photon flux density).Growth in a CO₂-enriched atmosphere seems to protect cork oakleaves from the short-term effects of high temperature. ElevatedCO₂ plants had positive net carbon uptake rates during the heatshock treatment whereas plants grown at ambient CO₂ showed negativerates. Moreover, recovery was faster in high CO₂-grown plantswhich, after 30 min at 25C, exhibited higher net carbon uptakerates and lower decreases in photosynthetic capacity (A_max aswell as in the efficiency of excitation energy captured by openphotosystem II reaction centres (F_vJF_m than plants grown atambient CO₂. The stomata of elevated CO₂ plants were also lessresponsive when exposed to high temperature. Key words: Elevated CO₂, temperature, acclimation, photosynthesis, Quercus suber L.     

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     

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