Effects of elevated CO2 on flowering phenology and nectar production of nectar plants important for butterflies of calcareous grasslands

Effects of elevated CO₂ on flowering phenology and nectar production were investigated in Trifolium pratense, Lotus corniculatus, Scabiosa columbaria, Centaurea jacea and Betonica officinalis, which are all important nectar plants for butterflies. In glasshouse experiments, juvenile plants were exposed to ambient (350 μl l⁻¹) and elevated (660 μl l⁻¹) CO₂ concentrations for 60–80 days. Elevated CO₂ significantly enhanced the development of flower buds in C. jacea. B. officinalis flowered earlier and L. corniculatus produced more flowers under elevated CO₂. In contrast, the number of flowers decreased in T. pratense. The amount of nectar per flower was not affected by elevated CO₂ in the tested legumes (T. pratense and L. corniculatus), but was significantly reduced (!) in the other forbs. Elevated CO₂ did not significantly affect nectar sugar concentration and composition. However, S. columbaria and C. jacea produced significantly less total sugar under elevated CO₂. The nectar amino acid concentration remained unaffected in all investigated plant species, whereas the total of amino acids produced per flower was reduced in all non-legumes. In addition, the amino acid composition changed significantly in all investigated species except for C. jacea. The observed effects are unexpected and are a potential threat to flower visitors such as most butterflies which have no alternative food resources to nectar. Changes in nectar production due to elevated CO₂ could also have generally detrimental effects on the interactions of flowers and their pollinators. Received: 12 September 1996 / Accepted: 9 September 1997     

在高CO2浓度下生长的小麦对棉铃虫生长发育和繁殖的影响

通过室内饲养实验研究了在高CO₂浓度（738.8±25.7μL/L）中生长的小麦对棉铃虫 Helicoverpa armigera (Hübner)生长发育,繁殖以及营养效应的影响。结果表明： （1）取食高CO₂浓度大气中生长的麦粒的棉铃虫对食料的取食量和粪便排泄量增加,与对照相比,取食量和粪便排泄量分别增加46.3%（P<0.05）和37.8%;（2）大气CO₂浓度增加影响了麦粒中的营养成分的含量,其中,可溶性蛋白、游离氨基酸、葡萄糖和总糖的含量及碳氮比(C∶N)都显著增加,果糖和粗蛋白的含量都显著降低;（3）大气 CO₂浓度升高所导致的麦粒营养成分的变化影响了棉铃虫幼虫的食物利用效率,与对照组相比,棉铃虫幼虫对食物的毛转化率和净转化率分别降低27.2%和25.4%,对食物的相对取食率则显著提高58.8%（P < 00.1）。据此推测,未来高CO₂浓度的大气环境会降低春小麦的营养价值,从而影响棉铃虫的生长发育,加重其对小麦的危害。     

Above- and Below-ground Production of Young Scots Pine (Pinus sylvestris L.) Trees after Three Years of Growth in the Field under Elevated CO2

Scots pine (Pinus sylvestris L.) seedlings were grown for 3years in the ground in open top chambers and exposed to twoconcentrations of atmospheric CO₂(ambient or ambient + 400 µmol mol^-1) without addition of nutrients and water. Biomassproduction (above-ground and below-ground) and allocation, aswell as canopy structure and tissue nitrogen concentrationsand contents, were examined by destructive harvest after 3 years.Elevated CO₂increased total biomass production by 55%, reducedneedle area and needle mass as indicated, respectively, by lowerleaf area ratio and leaf mass ratio. A relatively smaller totalneedle area was produced in relation to fine roots under elevatedCO₂. The proportion of dry matter in roots was increased byelevated CO₂, as indicated by increased root-to-shoot ratioand root mass ratio. Within the root system, there was a significantshift in the allocation towards fine roots. Root litter constituteda much higher fraction of fine roots in trees grown in the elevatedCO₂than in those grown in ambient CO₂. Growth at elevated CO₂causeda significant decline in nitrogen concentration only in theneedles, while nitrogen content significantly increased in branchesand fine roots (with diameter less than 1 mm). There were nochanges in crown structure (branch number and needle area distribution).Based upon measurements of growth made throughout the 3 years,the greatest increase in biomass under elevated CO₂took placemainly at the beginning of the experiment, when trees grownin elevated CO₂had higher relative growth rates than those grownunder ambient CO₂; these differences disappeared with time.Symptoms of acclimation of trees to growth in the elevated CO₂treatmentwere observed and are discussed. Copyright 2000 Annals of BotanyCompany Elevated CO₂, Pinus sylvestris, biomass production, allocation, fine roots, root litter, crown structure, nitrogen, C/N ratio     

Photosynthesis and photorespiration in soybean [Glycine max (L.) Merr.] chronically exposed to elevated carbon dioxide and ozone

The effects of elevated carbon dioxide (CO₂ and ozone (O₃) onsoybean (Glycine max (L.) Merr.] photosynthesis and photorespiration-relatedparameters were determined periodically during the growing seasonby measurements of gas exchange, photorespiratory enzyme activitiesand amino acid levels. Plants were treated in open-top fieldchambers from emergence to harvest maturity with seasonal meanconcentrations of either 364 or 726 µmol mol^–1 CO₂in combination with either 19 or 73 nmol mol^–1 O₃ (12h daily averages). On average at growth CO₂ concentrations,net photosynthesis (A) increased 56% and photorespiration decreased36% in terminal mainstem leaves with CO₂ enrichment. Net photosynthesisand photorespiration were suppressed 30% and 41%, respectively,by elevated O₃ during late reproductive growth in the ambientCO₂ treatment, but not in the elevated CO₂ treatment. The ratioof photorespiration to A at growth CO₂ was decreased 61% byelevated CO₂ There was no statistically significant effect ofelevated O₃ on the ratio of photorespiration to A. Activitiesof glycolate oxidase, hydroxypyruvate reductase and catalasewere decreased 10–25% by elevated CO₂ and by 46–66%by elevated O₃ at late reproductive growth. The treatments hadno significant effect on total amino acid or glycine levels,although serine concentration was lower in the elevated CO₂and O₃ treatments at several sampling dates. The inhibitoryeffects of elevated O₃ on photorespiration-related parameterswere generally commensurate with the O₃-induced decline in A.The results suggest that elevated CO₂ could promote productivityboth through increased photoassimilation and suppressed photorespiration. Key words: Photorespiration, CO₂-enrichment, ozone, climate change, air pollution     

Compensatory Roles of Nitrogen Uptake and Photosynthetic N-use Efficiency in Determining Plant Growth Response to Elevated CO2: Evaluation Using a Functional Balance Model

We used a modified functional balance (FB) model to predictgrowth response of Helianthus annuus L. to elevated CO₂. Modelpredictions were evaluated against measurements obtained twiceduring the experiment. There was a good agreement between modelpredictions of relative growth rate (RGR) responses to elevatedCO₂and observations, particularly at the second harvest. Themodel was then used to compare the relative effects of biomassallocation to roots, nitrogen (N) uptake and photosyntheticN-use efficiency (PNUE) in determining plant growth responseto elevated CO₂. The model predicted that a rather substantialincrease in biomass allocation to root growth had little effecton whole plant growth response to elevated CO₂, suggesting thatplasticity in root allocation is relatively unimportant in determininggrowth response. Average N uptake rate at elevated comparedto ambient CO₂was decreased by 21–29%. In contrast, elevatedCO₂increased PNUE by approx. 50% due to a corresponding risein the CO₂-saturation factor for carboxylation at elevated CO₂.The model predicted that the decreased N uptake rate at elevatedCO₂lowered RGR modestly, but this effect was counterbalancedby an increase in PNUE resulting in a positive CO₂effect ongrowth. Increased PNUE may also explain why in many experimentselevated CO₂enhances biomass accumulation despite a significantdrop in tissue nitrogen concentration. The formulation of theFB model as presented here successfully predicted plant growthresponses to elevated CO₂. It also proved effective in resolvingwhich plant properties had the greatest leverage on such responses.Copyright 2000 Annals of Botany Company Elevated CO₂, functional balance model, Helianthus annuus L., N uptake, photosynthetic nitrogen use efficiency, root:shoot ratio     

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.     

Phloem Exudate Collected via Scale Insect Stylets for the CAM Species Opuntia ficus-indica under Current and Doubled CO2 Concentrations

A method was devised for collecting phloem sap from the CAMspecies Opuntia ficus-indica using severed stylets of a scaleinsect (Dactylopius opuntiae), for which exudation could continuefor up to 5 d. For both basal (planted) cladodes and first-orderdaughter cladodes, the concentrations of sucrose and total aminoacids in the phloem exudate were virtually constant over 24-hperiods whereas the chlorenchyma osmolality had sizeable increasesduring the night under both current and doubled atmosphericCO₂ concentrations. Sucrose, total amino acids, and potassiumaccounted for 56, 21, and 9%, respectively, of the osmolalityof the phloem exudate, which was about 350 mOsm at the two CO₂concentrations; valine, isoleucine, leucine, tyrosine, glutamine,and lysine accounted for about 70% of the total amino acids.Doubling the CO₂ concentration led to approx. 5% more sucrose,560% more mannose and 17% less amino acids in the phloem exudateand also significantly increased mannose, starch and glucomannanin the chlorenchyma. Atmospheric CO₂ concentrations thus affectedvarious solute properties in the phloem and the chlorenchymaof O. ficus-indica.Copyright 1995, 1999 Academic Press Dactylopius opuntiae, Opuntia ficus-indica, cladode, CO₂ concentrations, Crassulacean acid metabolism, phloem exudate     

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     

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     

Chloroplast ultrastructure of sugar beet (Beta vulgaris L.) cultivated in normal and elevated CO2 concentrations with two contrasted nitrogen supplies

Sugar beet (Beta vulgaris L., cultivar Celt) plants were grownunder simulated field conditions in pots and supplied with adequateor deficient nitrogen (HN and LN, respectively) combined withtwo CO₂ concentrations, ambient (c. 350µmol mol^–1C0₂—AC), or elevated CO₂ (c. 600 µmol mol^–1CO₂—HC). Chloroplast structure in mesophyll palisade cellsof mature leaves (leaf number 19 in HN and 9 in LN), sampledat midday on 16 August 1993 was studied by transmission electronmicroscopy and quantified stereologically. The ultrastructureof palisade parenchyma chloroplasts was affected by the elevatedCO₂ concentration and strikingly affected by nitrogen supply.Chloroplast diameter (cross-sectional length) was slightly,but not significantly, greater in HC than AC treatments withinan N treatment, but was smaller in LN than HN; chloroplast cross-sectionalarea also increased with HC in both N treatments, but only significantlyso in LN. Elevated CO₂ reduced the proportion of total thylakoids(significant at 5% and 0.1% in HN and LN, respectively) dueto decreased granal thylakoids, but the proportion of inter-granal(stromal) thylakoid membranes was not affected compared to chloroplastsfrom plants grown with ambient CO₂. Chloroplast stroma increasedas a proportion of chloroplast volume with elevated comparedto ambient CO₂ with HN but not LN. Starch inclusions were notsignificantly different with elevated compared to ambient CO₂at HN, but the proportion of starch increased considerably atelevated compared to ambient C0₂ at LN, indicating an over-productionof assimilates. Plastoglobuli in chloroplasts increased withdeficient N, but decreased with elevated CO₂. Larger chloroplastswith a greater proportion of stroma, but a smaller proportionof granal thylakoids, suggest increased CO² assimilating capacityand decreased light harvesting/PSII capacity with elevated CO₂. Key words: Chloroplast, ultrastructure, elevated CO₂ concentration, nitrogen deficiency, sugar beet, Beta vulgaris     

The Interactive Effects of Carbon Dioxide Enrichment and Daylength on Growth and Development inPetunia hybrida

Plants were grown at either 350 or 1000 µl l^-1CO₂and inone of three photoperiod treatments: continuous short days (SD),continuous long days (LD), or short switched to long days atday 41 (SD–LD). All plants received 9 h of light at 450µmol m^-2s^-1and LD plants received an additional 4 h oflight at 8 µmol m^-2s^-1. Growth of SD plants respondedmore positively to elevated CO₂than did LD plants, due largelyto differences in the effect of CO₂on unit leaf rate. High CO₂increasedheight and decreased branching under SD conditions, but hadno effect under LD conditions. Elevated CO₂also increased thenumber of buds and open flowers, the effect for flower numberbeing greater in short than in long days. The specific leafarea of plants grown at 1000 µl l^-1CO₂was reduced regardlessof daylength. High CO₂also decreased leaf and increased reproductiveallocation, the magnitude of these effects being greater underSD conditions. Bud formation and flower opening was advancedunder high CO₂conditions in SD plants but bud formation wasdelayed and there was no effect on flower opening under LD conditions.The effects of CO₂on plants switched from SD to LD conditionswere largely intermediate between the two continuous treatments,but for some parameters, more closely resembled one or the other.The results illustrate that daylength is an important factorcontrolling response of plants to elevated CO₂. Petunia hybridaHort. ex Vilm; carbon dioxide; photoperiod; functional growth analysis; daylength; global change; development; phenology     

Non-Autotrophic CO2 Fixation and Drought Tolerance in Mosses

Cratoneuron filicinum, a drought-sensitive moss, and Tortularuralis, a drought-tolerant moss, fix CO₂ non-autotrophicallyat a rate of about 1.2 and 2.2 µmol h^–1 g^–1dry wt. respectively. During drying, T. ruralis fixes CO₂ atan undiminished rate until the tissue loses about 60% of theinitial fresh weight. Thereafter, CO₂ fixation rapidly declinesto zero. Dark CO₂ fixation by C.filicinum declines steadilyduring the dehydration period. On rehydration, dark CO₂ fixationis resumed immediately in T. ruralis but not in C.filicinum.When dried T. ruralis is equilibrated with an atmosphere ofnearly 100% relative humidity, its weight increases to about40% of the original fresh weight and dark CO₂ fixation resumesat a rate about 60% of the fresh moss. In C.filicinum thereis only a small increase in weight and little CO₂ fixation inthe dark. The non-autotrophically fixed carbon, in both mossesstudied, is incorporated into amino acids (more than 60% ofthe total, mainly into aspartate, alanine and glutamate) andorganic acids (less than 40% of the total, mainly into malate).It is suggested that on rehydration immediate availability ofNADPH, known to be produced by transhydrogenation from NADHduring dark CO₂ fixation, may be an important factor in therepair of drought-induced cellular damage by reductive biosynthesisof membrane components and other cellular constituents. Key words: Mosses, Dehydration, Rehydration, Dark CO₂ fixation, Amino acids, Organic acids, NADPH, Drought tolerance.     

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     

Garden Flowers: Insect Visits and the Floral Reward of Horticulturally-modified Variants

Garden flowers can be valuable to wildlife if they produce nectar,pollen and/or seeds. To provide information needed by gardenersto select wildlife-friendly plants, we investigated nectar productionand insect visits toTropaeolum majus, Consolidasp.,Antirrhinummajus, Violaxwittrockiana, Tagetes patulaandAlcea rosea, ineach case comparing a near-original flower type with a cultivarthat had spurless, doubled, peloric or enlarged flowers. Allspecies showed high secretion rates and standing crops of nectar.In most cases the horticultural modifications affected the numbersor species composition of the assemblage of insect visitors,and they generally reduced the value of the floral reward toinsects, often affecting accessibility. Effects on seed yieldwere not investigated directly here, but are likely to furtherreduce the wildlife value of modified variants.Copyright 1999Annals of Botany Company. Garden plants, horticultural modification, nectar, pollen, coevolution, insects, pollinators,Bombus, Apis mellifera,foraging.     

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.     

Soil O2 and CO2 Effects on Apparent Cell Viability for Roots of Desert Succulents

Roots of desert succulents occupy the upper layers of porous,well-aerated soils. However, roots of Agave deserti, Ferocactusacanthodes, and Opuntia ficus-indica all tolerated many daysof soil anoxia; 0% O₂ in the soil gas phase for 30 d reducedthe fraction of cells taking up the vital stain neutral red,an average of only 18% for the cortex and 6% for parenchymacells within the stele of perennial established roots. Ephemeralrain roots, induced by watering as branches on the establishedroots, were more susceptible to 0% O₂ in the soil gas phase;19 d abolished stain uptake for cortical cells and 32 d forstelar parenchyma cells. Soil CO₂ levels above the 0.1% observedin the root zone in the field rapidly reduced uptake of neutralred; the fraction of cortical cells taking up the stain decreased30% in 10 h at 0.5% CO₂ and was abolished in 9 h at 2% and 7h at 10% CO₂ averaged for the three species. Rain roots weresomewhat more susceptible than established roots to elevatedsoil CO₂ levels, and stelar parenchyma cells were much lesssusceptible than were cortical cells. When uptake of the vitalstain was abolished by elevated soil CO₂, no anatomical evidenceof cellular damage was observed. For A. deserti exposed to 2%CO₂, the pH of macerated root tissue decreased about 0.35 pHunit over 10 h; CO₂ apparently entered the cells, lowered theintracellular and/or cell wall pH, and prevented the accumulationof neutral red. Elevated soil CO₂ also inhibits root respirationfor the three desert succulents considered. Hence, the restrictionof such species to porous soils may reflect the relatively rapidinhibiting effects of elevated soil CO₂ levels rather than arequirement for high soil O₂ levels, consistent with the observationthat desert soils tend to have low gas-phase CO₂ levels near0.1% compared with 1% or more in the root zone of non-desertspecies. Key words: Agave deserti, Ferocactus acanthodes, neutral red, Opuntia ficus-indica, pH     

Contrasting CO2 and temperature effects on leaf growth of perennial ryegrass in spring and summer

The effects of increased atmospheric carbon dioxide (CO₂) of700 µmol mol^–1 and increased air temperature of+ 4C were examined in Lolium perenne L. cv. Vigor, growingin semi-controlled greenhouses. Leaf growth, segmental elongationrates (SER), water relations, cell wall (tensiometric) extensibility(%P) and epidermal cell lengths (ECL) were measured in expandingleaves in spring and summer. In elevated CO₂, shoot dry weight (SDW) increased in mid-summer.In both seasons, SDW decreased in elevated air temperatureswith this reduction being greater in summer as compared to spring.Specific leaf area (SLA) decreased in elevated CO₂ and in CO₂ temperature in both seasons. In spring, increased leaf extensionand SER in elevated CO₂ were linked with increased ECL, %P andfinal leaf size whilst in summer all were reduced. In high temperature,leaf extension, SER, %P and final leaf size were reduced inboth seasons. In elevated CO₂ temperature, leaf extension,SER, %P, and ECL increased in spring, but final leaf size remainedunaltered, whilst in summer all decreased. Mid-morning waterpotential did not differ with CO₂ or temperature treatments.Leaf turgor pressure increased in elevated CO₂ in spring andremained similar to the control in summer whilst solute potentialdecreased in spring and increased in summer. Contrasting seasonalgrowth responses of L. perenne in response to elevated CO₂ andtemperature suggests pasture management may change in the future.The grazing season may be prolonged, but whole season productivitymay become more variable than today. Key words: Lolium perenne, ryegrass, CO₂ and temperature, leaf extension, cell wall rheology     

Limitation to Photosynthesis in Water-stressed Leaves: Stomata vs. Metabolism and the Role of ATP

Decreasing relative water content (RWC) of leaves progressivelydecreases stomatal conductance (g_s), slowing CO₂ assimilation(A) which eventually stops, after which CO₂ is evolved. In somestudies, photosynthetic potential (A_pot), measured under saturatingCO₂, is unaffected by a small loss of RWC but becomes progressivelymore inhibited, and less stimulated by elevated CO₂, below athreshold RWC (Type 1 response). In other studies, A_pot andthe stimulation of A by elevated CO₂ decreases progressivelyas RWC falls (Type 2 response). Decreased A_pot is caused byimpaired metabolism. Consequently, as RWC declines, the relativelimitation of A by g_s decreases, and metabolic limitation increases.Causes of decreased A_pot are considered. Limitation of ribulosebisphosphate (RuBP) synthesis is the likely cause of decreasedA_pot at low RWC, not inhibition or loss of photosynthetic carbonreduction cycle enzymes, including RuBP carboxylase/oxygenase(Rubisco). Limitation of RuBP synthesis is probably caused byinhibition of ATP synthesis, due to progressive inactivationor loss of Coupling Factor resulting from increasing ionic (Mg²⁺)concentration, not to reduced capacity for electron or protontransport, or inadequate trans-thylakoid proton gradient (pH).Inhibition of A_pot by accumulation of assimilates or inadequateinorganic phosphate is not considered significant. DecreasedATP content and imbalance with reductant status affect cellmetabolism substantially: possible consequences are discussedwith reference to accumulation of amino acids and alterationsin protein complement under water stress.     

Elevated atmospheric CO2 concentration ameliorates effects of NaCl salinity on photosynthesis and leaf structure of Aster tripolium L.

This study investigated the interaction of NaCl-salinity andelevated atmospheric CO₂ concentration on gas exchange, leafpigment composition, and leaf ultrastructure of the potentialcash crop halophyte Aster tripolium. The plants were irrigatedwith five different salinity levels (0, 25, 50, 75, 100% seawatersalinity) under ambient and elevated (520 ppm) CO₂. Under salineconditions (ambient CO₂) stomatal and mesophyll resistance increased,leading to a significant decrease in photosynthesis and wateruse efficiency (WUE) and to an increase in oxidative stress.The latter was indicated by dilations of the thylakoid membranesand an increase in superoxide dismutase (SOD) activity. Oxidativestress could be counteracted by thicker epidermal cell wallsof the leaves, a thicker cuticle, a reduced chlorophyll content,an increase in the chlorophyll a/b ratio and a transient declineof the photosynthetic efficiency. Elevated CO₂ led to a significantincrease in photosynthesis and WUE. The improved water and energysupply was used to increase the investment in mechanisms reducingwater loss and oxidative stress (thicker cell walls and cuticles,a higher chlorophyll and carotenoid content, higher SOD activity),resulting in more intact thylakoids. As these mechanisms canimprove survival under salinity, A. tripolium seems to be apromising cash crop halophyte which can help in desalinizingand reclaiming degraded land. Key words: Aster tripolium, cash crop halophyte, elevated CO₂, gas exchange, oxidative stress, photosynthesis, salt tolerance, ultrastructure, water use efficiency Received 29 July 2008; Revised 8 October 2008 Accepted 9 October 2008     

Effects of nitrogen on Pinus palustris foliar respiratory responses to elevated atmospheric CO2 concentration

Indirect effects of atmospheric CO₂ concentration [CO₂], onlongleaf pine (Pinus palustris Mill.) foliage respiration werestudied by growing trees in a factorial arrangement of low andhigh [CO₂] (369 and 729µmol CO₂ mol^–1) and low andhigh N (40 and 400 kg ha^–1 yr^–1). Direct effectsof [CO₂] on leaf respiration were tested by measuring respirationrates of foliage from all treatments at two CO₂ levels (360and 720µmol CO₂mol^–1) at the time of measurement.Elevated CO₂ did not directly or indirectly affect leaf respirationwhen expressed on a leaf area or mass basis, but a significantincrease in respiration per unit leaf N was observed in treesgrown in elevated [CO₂] (indirect response to elevated [CO₂]).The lack of a [CO₂] effect on respiration, when analysed onan area or mass basis, may have resulted from combined effectsof [CO₂] on factors that increase respiration (e.g. greateravailability of non-structural carbohydrates stimulating growthand carbon export from leaves) and on factors that decreaserespiration (e.g. lower N concentration leading to lower constructioncosts and maintenance requirements). Thus, [CO₂] affected factorsthat influence respiration, but in opposing ways. Key words: Pinus palustris, elevated CO₂, nitrogen, foliar, respiration