Carbon isotope ratios in exhaled CO2 can be used to determine not just present, but also past diets in birds

We show that an animal's past and present diet can be distinguished through the delta(13)C of exhaled CO(2). The exhaled delta(13)C of 12 pigeons fed solely corn (a C(4) plant) for 30 days was -13.63 per thousand (+/-0.30). We then fed six pigeons wheat (a C(3) plant) and continued to feed the other six corn. After 48 h the exhaled delta(13)C from the corn-fed pigeons was unchanged; that from the wheat-fed pigeons was -20.5 per thousand. We then fasted three of the wheat-fed pigeons for 3 days, after which their exhaled delta(13)C was -14.96 per thousand, while it was -13.57 per thousand in corn-fed pigeons, and -22.22 per thousand in pigeons that continued on wheat. Thus, we could infer diet from the (13)C/(12)C ratios of exhaled CO(2). Significantly, breath samples from fasted pigeons also revealed that they had eaten corn when their lipid stores were formed. We also showed that the change in the (13)C/(12)C of exhaled CO(2) had a half-life of approximately 3.5 h, and a time constant of approximately 6.7 h. Thus one can infer past and present diet from exhaled delta(13)C alone, if the initial breath sample is followed by a fasted breath sample, without harming the animal or having to recapture it successively.     

13C/12C Ratios of carbon dioxide from peanut and sunflower seedlings and tobacco leaves in light and in darkness

Shoots of intact peanut and sunflower seedlings evolved CO₂in the light which was enriched more than 10 per mille in ¹³Ccompared with simultaneous CO₂ evolution from the roots. Carbondioxide collected from tobacco leaves in the light was 10 permille enriched in ¹³C compared with that collected in the dark.Anaerobic conditions inhibited photorespiration but did notchange isotopic ratios of dark respiration. ¹ Department of Biology, Fresno State College, Fresno, California93710, U. S. A. ² Deceased. (Received February 29, 1972; )     

Carbon isotope ratios of respired CO2 from castor bean, corn, peanut, pea, radish, squash, sunflower and wheat seedlings

During the first month after germination peanut and sunflowerseedlings exhibit a significant shift in the ¹³C/¹²C ratiosof respired CO₂ indicating thatcarbohydrate gives way to lipidas the respiratory substrate. Other species (castor bean, corn,pea,radish, squash and wheat) show no change in the ¹³C/¹²C ratio (Received March 9, 1971; )     

Algal 14C and total carbon metabolisms. 2. Experimental observations with the diatom Skeletonema costatum

Three sets of comparisons of net and gross inorganic carbonassimilation and ¹⁴C uptake were made with an axenic cultureof Skdetonema costatum. The comparisons showed that in the physiologicalwindow studied (10–20% of the intrinsic generation timeand gross photosynthesis/respiration ratios of 2–3), ¹⁴Cuptake into the paniculate plus the dissolved fractions approximatedto net photosynthesis. Rate constants derived from the chemicallydetermined changes were used to parameterize models that accountedfor the respiration of photosynthetic products and for the recyclingof respiratory CO₁₄. The conclusion drawn was that over thetime scale studied, the ¹⁴C technique was measuring net photosynthesis,consistent with essentially 100% recycling of respiratory CO₂.The study has shown that we now possess the basis to make arigorous analysis of net, gross CC₄ fixation and net ¹⁴C uptake,and forms the first step in the development of algorithms forthe interpretation of ¹⁴C field observations.     

Oxygen Exchange in Relation to Carbon Assimilation in Water-stressed Leaves During Photosynthesis

In a study on metabolic consumption of photosynthetic electronsand dissipation of excess light energy under water stress, O₂and CO₂ gas exchange was measured by mass spectrometry in tomatoplants using ¹⁸O₂ and ¹³CO₂. Under water stress, gross O₂ evolution(E_O), gross O₂ uptake (U_O), net CO₂ uptake (P_N), gross CO₂ uptake(TPS), and gross CO₂ evolution (E_C) declined. The ratio P_N/E_Ofell during stress, while the ratios U_O/E_O and E_C/TPS rose.Mitochondrial respiration in the light, which can be measureddirectly by ¹²CO₂ evolution during ¹³CO₂ uptake at 3000 µll^{–1 13}CO₂, is small in relation to gross CO₂ evolutionand CO₂ release from the glycolate pathway. It is concludedthat PSII, the Calvin cycle and mitochondrial respiration aredown-regulated under water stress. The percentages of photosyntheticelectrons dissipated by CO₂ assimilation, photorespiration andthe Mehler reaction were calculated: in control leaves morethan 50 % of the electrons were consumed in CO₂ assimilation,23 % in photorespiration and 13 % in the Mehler reaction. Undersevere stress the percentages of electrons dissipated by CO₂assimilation and the Mehler reaction declined while the percentageof electrons used in photorespiration doubled. The consumptionof electrons in photorespiration may reduce the likelihood ofdamage during water deficit.     

Dynamics of Carbon Photosynthetically Assimilated in Nodulated Soya Bean Plants under Steady-state Conditions 1. Development and Application of 13CO2 Assimilation System at a Constant 13C Abundance

A long-term, steady-state ¹³CO₂ assimilation system at a constantCO₂ concentration with a constant ¹³C abundance was designedand applied to quantitative investigations on the allocationof photoassimilated carbon in nodulated soya bean (Glycine maxL.) plants. The CO₂ concentration in the assimilation chamberand its ¹³C abundance were maintained constant with relativevariances of less than ±0.5 per cent during an 8-h assimilationperiod. At the termination of 8-h ¹³CO₂ assimilation by plantsat early flowering stage, the currently assimilated carbon relativeto total tissue carbon (measured by the degree of isotopic saturation)were for young leaves (including flower buds), 13.9 per cent;mature leaves, 15.7 per cent; stems+petioles, 5.9 per cent;roots, 5.4 per cent and nodules, 6.9 per cent, 48 h after theend of the ¹³CO₂ assimilation period, they were 12.3, 7.5, 7.4,6.8 and 6.1 per cent, respectively. The treatment with a highconcentration of nitrate in the nutrient media significantlydecreased the allocation of ¹³C into nodules. Experiments on¹³CO₂ assimilation by plants at the pod-filling stage were alsoconducted. Labelling by ¹³C was weaker than at the early floweringstage, but an intense accumulation of ¹³C into reproductiveorgans was observed. Glycine max L., nodulated soya bean plants, ¹³CO₂ assimilation, carbon dynamics     

Dynamics of Carbon Photosynthetically Assimilated in Nodulated Soya Bean Plants under Steady-state Conditions 3. Time-course Study on 13C Incorporation into Soluble Metabolites and Respiratory Evolution of 13CO2 from Roots and Nodules

Well-nodulated soya bean (Glycine max L.) plants were allowedto assimilate ¹³CO₂ for 10 h in the light, under steady-stateconditions in which CO₂ concentration and ¹³C abundance wereboth strictly controlled at constant levels. The respiratoryevolution of ¹³CO₂ from roots and nodules and ¹³C incorporationinto various metabolic fractions were measured during the ¹³CO₂feeding and subsequent 48 h chase period. CO₂ respired from nodules was much more rapidly labelled with¹³C than that from roots. The level of labelling (percentageof carbon currently assimilated during the ¹³COM₂ feeding period)of CO₂ respired from nodules reached a maximum of about 87 percent after 4 h of steady-state ^l3CO₂ assimilation and thereafterremained fairly constant. The absolute amount of labelled carbonevolved by the respiration of the nodules during the 10 h ¹³CO₂feeding period was 1·5-fold that of root respiration.These results demonstrated that the currently assimilated (labelled)carbon was preferentially used to support nodule respiration,while root respiration relied considerably on earlier (non-labelled)carbon reserved in the roots. Sucrose pools were mostly composed of currently assimilatedcarbon in all tissues of the plants, since the levels of labellingaccounted for 86–91 per cent at the end of the ¹³CO₂ feeding.In the nodules, the kinetics and levels of sucrose labellingwere in fairly good agreement with those of respired CO₂, whilein the roots, the level of labelling of respired CO₂ was significantlylower than that of sucrose. Succinate and malate were highly labelled in both roots andnodules but they were labelled much more slowly than sucroseand respired CO₂. The kinetics and levels of labelling of theseKrebs cycle intermediates resembled those of major amino acidswhich are derived directly from Krebs cycle intermediates. Itis suggested that large fractions of organic acids in noduleswere physically separate from the respiration site. Glycine max L., Soya bean, ¹³CO₂ assimilation, respiratory evolution of ¹³CO₂, carbon metabolism in root nodules     

Microbial community composition and function beneath temperate trees exposed to elevated atmospheric carbon dioxide and ozone

We hypothesized that changes in plant growth resulting from atmospheric CO₂ and O₃ enrichment would alter the flow of C through soil food webs and that this effect would vary with tree species. To test this idea, we traced the course of C through the soil microbial community using soils from the free-air CO₂ and O₃ enrichment site in Rhinelander, Wisconsin. We added either ¹³C-labeled cellobiose or ¹³C-labeled N-acetylglucosamine to soils collected beneath ecologically distinct temperate trees exposed for 3 years to factorial CO₂ (ambient and 200 µl l^-1 above ambient) and O₃ (ambient and 20 µl l^-1 above ambient) treatments. For both labeled substrates, recovery of ¹³C in microbial respiration increased beneath plants grown under elevated CO₂ by 29% compared to ambient; elevated O₃ eliminated this effect. Production of ¹³C-CO₂ from soils beneath aspen (Populus tremuloides Michx.) and aspen-birch (Betula papyrifera Marsh.) was greater than that beneath aspen-maple (Acer saccharum Marsh.). Phospholipid fatty acid analyses (¹³C-PLFAs) indicated that the microbial community beneath plants exposed to elevated CO₂ metabolized more ¹³C-cellobiose, compared to the microbial community beneath plants exposed to the ambient condition. Recovery of ¹³C in PLFAs was an order of magnitude greater for N-acetylglucosamine-amended soil compared to cellobiose-amended soil, indicating that substrate type influenced microbial metabolism and soil C cycling. We found that elevated CO₂ increased fungal activity and microbial metabolism of cellobiose, and that microbial processes under early-successional aspen and birch species were more strongly affected by CO₂ and O₃ enrichment than those under late-successional maple.     

Effects of NO2 and O3 Alone or in Combination on Kidney Bean Plants (Phaseolus vulgaris L.): Products of 13CO2 Assimilation Detected by 13C Nuclear Magnetic Resonance

The effect of exposure of kidney bean primary leaves to NO₂and O₃, alone or in combination, on the fate of ¹³CO₂ assimilatedby photosynthesis was examined by ¹³C-NMR. There were more than70 peaks appearing in the ¹³C-NMR spectra for substances extractedfrom leaves with 80% ethanol. The 16 relatively well resolvedpeaks corresponded to signals from three sugars, two organicacids and four amino acids. These signals were used to estimatepool sizes and ^l3C incorporation. Exposures to NO₂ and O₃ increased the amounts of sucrose andfructose, but not the incorporation of the ¹³C label during10 min photosynthesis from ¹³CO₂. This suggests the presenceof photo-synthetically inactive pools of sucrose and fructose.Amounts of glycine and serine, and ¹³C incorporation into them,were increased by the exposure to the pollutants. The incorporationof ¹³C into alanine was stimulated by exposure to NO₂, but notby exposure to O₃ alone. The present study shows that with only simple procedures ofsample preparations ¹³C-NMR provides information on the productsof photosynthesis in leaves stressed by the two air pollutants. Key words: NO₂, O₃, Phaseolus vulgaris, CO₂ assimilation, ¹³C-NMR     

Features of Photosynthesis in Haloxylon species of Chenopodiaceae that are Dominant Plants in Central Asian Deserts

Haloxylon aphyllum and H. persicum of Chenopodiaceae are dominantplants in the continental deserts of the Asian Irano-Turanianregion. The photosynthetic organs, assimilating shoots and leaf-likecotyledons of these two species were studied to characterizetheir photosynthetic types. ¹³C/¹²C isotope ratios, the cellularanatomy of as similating organs, primary photosynthetic products,and activities of carbon metabolism enzymes, RUBP carboxylase,PEP carboxylase, malic enzymes, and aspartate aminotransferase,indicate different pathways of CO₂ fixation in the photosyntheticorgans. Assimilating shoots had attributes of the C₄ photosynthesisentirely, while cotyledons lack Kranz-anatomy and incorporatedCO₂ via C₃ photosynthesis. Cotyledons and seeds had lower     

Refixation of respiratory CO2 in the ears of C3 cereals

The spatial arrangement of tissues within the ears of cereals,and gas exchange measurements on intact ears of barley and durumwheat suggest that respiratory CO₂ associated with grain-fillingprocesses, may be refixed close to its site of evolution. Apparentrefixation of respiratory CO₂ in intact ears was compared withthat in flag leaves, by feeding both organs with ¹⁴C-labelledsucrose and trapping ¹⁴CO₂ released by respiration. Apparentrefixation in the ears was twice that measured in flag leafblades of durum wheat genotype Durelle. In ears, the capacityof refixation of respiratory CO₂ at 210 mmol mol^–1 O₂ranged from 55% in barley genotype Roxana to 75% in barley genotypeHatif, and 60% in duwm wheat genotype Bidi 17. A low O₂ concentrationincreased refixation of respiratory CO₂ by up to 90%, 92% and82%, respectively. The occurrence of CO₂ refixation in the field,in a set of 12 barley genotypes grown under irrigated and rainfedMediterranean field conditions, was consistent with observedcarbon isotope ratios (     

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.     

Natural abundance of 13C in CO2 respired from forest soils reveals speed of link between tree photosynthesis and root respiration

Soil respiration from a boreal mixed coniferous forest showed large seasonal variation in natural abundance of ¹³C, ranging from -21.6‰ to -26.5‰. We tested if weather conditions could explain this variation in '¹³C of respired CO₂, and found that the air relative humidity 1-4 days before the days of CO₂ sampling best explained the variation. This suggested that high '¹³C values were caused by effects of air humidity on isotope fractionation during photosynthesis and that it took 1-4 days for the C from canopy photosynthesis of 20-25 m trees to become available for root/rhizosphere respiration. We calculated that these new photoassimilates could account for at least 65% of total soil respiration.     

A Comparative Study of the Influence of Salt Type and Concentration on 14CO2 Fixation in Potato Slices at 25{degrees} C and 0{degrees} C

Dark fixation of ¹⁴CO₂ was followed in potato disks under varyingsalt treatments at 0° C and 25° C. It is shown thatthe specific activity of the ¹⁴CO₂ supplied is heavily dilutedby endogenously produced CO₂ and that the apparently greaterfixation of ¹⁴CO₂, at 0° C as compared with that at 25 °C is due to the lower respiration rate at 0° C, with consequentlyless dilution of the ¹⁴CO₂. supplied. At 25° C organic acidformation in response to different salt treatments fulfils thecommon expectation, ¹⁴CO₂ fixation increasing in the presenceof K₂SO₄ and decreasing in CaCl₂ relative to that in KCl. Therole of organic acids in maintaining ionic balance within thecell at 25° C is thereby indicated but at 0° C organicacid adjustments did not follow the normal pattern. At 25°C but not at o° C increasing external concentration of KCIresulted in an increased level of ¹⁴CO₂ fixation.     

The Influence of Leaf Development on the Expression of C4 Metabolism in Flaveria trinervia, a C4 Dicot

The initial products of ¹⁴CO₂ assimilation were determined understeady state illumination of leaves of Flaveria trinervia, aC₄ dicot of the NADP-mialic enzyme subgroup. Leaf age influencedthe partitioning of ¹⁴CO₂ between the C₄ cycle and the reductivepentose phosphate (RPP) pathway. An estimated 10 to 12%of theCO₂ entered the RPP pathway directly in leaves about 20% fullyexpanded, whereas CO₂ was apparently fixed entirely throughthe C₄ pathway in leaves 75% or more expanded. This partitioningpattern was attributed to the bundle sheath compartment in youngleaves having a relatively high conductance to CO₂ (i.e., beingsomewhat leaky). Of the initially labelled C₄ acids, the proportion that wasmalate, relative to aspartate, increased continuously duringleaf expansion (from 60 : 40 to 87 : 13 at full expansion).Concurrently, there was an increase in the whole leaf activityof NADP malate dehydrogenase and a decrease in the activitiesof aspartate and alanine aminotransferases. Low chlorophylla/b values were observed in young leaves, which may coincidewith an enhanced capacity for non-cyclic electron transportin the bundle sheath chloroplasts of such tissue. Both enhancedaspartate metabolism and direct fixation of CO₂ in the bundlesheath could provide a greater sink for utilization of photochemicallyderived NADPH in the bundle sheath of young leaves. Such metabolicchanges are discussed in relation to a possible decrease inCO₂ conductance of the bundle sheath during leaf development. (Received March 4, 1986; Accepted June 25, 1986)     

Measurements of 14C Incorporation by Illuminated Intact Leaves of Coffee Plants from Gas Mixtures Containing 14CO2

A study was made of the incorporation of ¹⁴C by intact leavesof Coffea arabica (cultivars Mundo Novo, Catuai, 1130–13,and H 6586–2) and Coffea canephora (cultivar Guarini)supplied with gas mixtures containing ¹⁴CO₂ under controlledconditions. Samples of the leaves were combusted and the ¹⁴Cin the CO₂ produced measured using a liquid scintillation counter.The results were used to estimate photosynthetic rates. Theeffects of changing the partial pressures of O₂ and CO₂ on thephotosynthetic rate were studied and estimates made of the CO₂compensation point and photorespiration. The data obtained show differences between the mean net photosyntheticrates of the C. arabica cultivars (6·14 mg CO₂ dm^–2h^–1) and the mean rate for the C. canephora cultivar (3·96mg CO₂ dm^–2 h^–1). The cultivar of the latter speciesphotorespired more rapidly than the cultivar Catuai of C. arabica.Rates of photosynthesis in coffee measured using the ¹⁴CO₂ methodwere similar to rates obtained by others using an infrared gasanalyser. The ¹⁴CO₂ method proved to be reliable for photosyntheticmeasurements and the apparatus is suitable for use in fieldconditions.     

Respiratory Utilization of 13C-Labelled Photosynthate in Nodulated Root Systems of Soybean Plants

Kouchi, H., Akao, S. and Yoneyama, T. 1986. Respiratory utilizationof ¹³C-labelled photosynthate in nodulated root systems of soybeanplants.—J. exp. Bot. 37: 985–993. An improved method for the measurement of respiratory utilizationof current photosynthate in the nodulated root system of water-culturedsoybean (Glycine max L.) plants was developed using a steady-state¹³CO₂ labelling technique. Well-nodulated plants at the latevegetative stage were allowed to assimilate ¹³CO₂ for 10 h incontinuous light at a constant CO₂ concentration with a constant¹³C abundance. The respiratory evolution of ¹³CO₂ from rootsand nodules was measured continuously throughout the periodof ¹³CO₂ assimilation and during a subsequent 36 h chase periodby using a differential infrared ¹³CO₂ analyser. The plantswere grown with nitrogen-free or (15 mmol dm^–3)-containing culture solution for 3 d before¹³CO₂ assimilation. In plants grown without , nodule respiration averaged 69% of the total respiration of the undergroundparts over the full experimental period and the CO₂ respiredreached an apparent isotopic equilibrium at 80–85% labellingafter initiating ¹³CO₂ assimilation. By contrast, the CO₂ respiredfrom the roots did not reach an isotopic equilibrium and labellingwas only 56% at the end of exposure to ¹³CO₂ These findingsdemonstrated that nodule respiration is strongly dependent onrecently assimilated carbon compared with root respiration. Plants supplied with in the culture solution showed a decreased rate of nodule respirationand a slightly increased rate of root respiration. The extentsand time courses of labelling of respired CO₂ from both theroots and nodules were similar in the presence and absence of except that the maximum level of labelling of CO₂ derived from nodule respiration in plantswith was significantly higher (about 91%) than for plants growing without . Key words: Soybean (Glycine max L.), nodule respiration, ¹³CO₂, assimilation, carbon partitioning     

Dynamics of Carbon Photosynthetically Assimilated in Nodulated Soya Bean Plants under Steady-state Conditions 2. The Incorporation of 13C into Carbohydrates, Organic Acids, Amino Acids and some Storage Compounds

Nodulated soya bean (Glycine max L.) plants at the early floweringstage were allowed to assimilate ¹³CO₂ under steady-state conditions,with a constant ¹³C abundance, for 8 h in the light. The plantswere either harvested immediately or 2 d after the end of the¹³CO₂ feeding, divided into young leaves (including flower buds),mature leaves, stems+petioles, roots and nodules; the ¹³C abundancein soluble carbohydrates, organic acids, amino acids, starchand poly-ß-hydroxybutyric acid was determined witha gas chromatography-mass spectrometry. The rapid turnover of ¹³C in the sucrose pools observed in allorgans of the plants showed that sucrose was the principal materialin the translocation stream of primary products of photosynthesis.At the end of the ¹³CO₂ exposure, sucrose in the mature leavesas the major source organs and in the stems+petioles was labelledwith currently assimilated carbon to about 75 per cent, whereasa much higher labelling of sucrose was found in the roots andin the nodules. This suggests the existence of two or more compartmentedpools of sucrose in mature leaves and also in stems+petioles. The relative labelling patterns of individual organic acidsand amino acids were similar in various plant organs. However,the rapid turnover of succinate and glycine was characteristicof nodules. Treatment with a high concentration of nitrate inthe nutrient media increased the turnover rate of amino acidcarbon in shoot organs and roots, while it markedly decreasedthe labelling of amino acids in nodules. The cyclitols, exceptfor D-pinitol, were significantly labelled with assimilated¹³C in mature leaves, but in nodules, the labelling was verymuch less. In the nodules, which were actively fixing atmospheric nitrogen,a large proportion (80–90 per cent) of currently assimilatedcarbon was found as sucrose and starch at the end of the ¹³CO₂feeding. This was also true of the roots. On the other hand,in young growing leaves, the distribution of currently assimilatedcarbon into sucrose, starch and other soluble compounds wasmuch less. This suggests that a large amount of carbon assimilatedby and translocated to young leaves was used to make up structuralmaterials, mainly protein and cell wall polymers synthesis,during the light period. Glycine max L., soya bean, ¹³CO₂ assimilation, carbon metabolism in nodules     

Growth and Partitioning in Pascopyrum smithii (C3) and Bouteloua gracilis(C4) as Influenced by Carbon Dioxide and Temperature

This study investigated how CO₂and temperature affect dry weight(d.wt) accumulation, total nonstructural carbohydrate (TNC)concentration, and partitioning of C and N among organs of twoimportant grasses of the shortgrass steppe,Pascopyrum smithiiRydb. (C₃) andBouteloua gracilis(H.B.K.) Lag. ex Steud. (C₄).Treatment combinations comprised two temperatures (20 and 35°C)at two concentrations of CO₂(380 and 750 µmol mol^-1),and two additional temperatures of 25 and 30°C at 750 µmolmol^-1CO₂. Plants were maintained under favourable nutrient andsoil moisture and harvested following 21, 35, and 49d of treatment.CO₂-induced growth enhancements were greatest at temperaturesconsidered favourable for growth of these grasses. Comparedto growth at 380 µmol mol^-1CO₂, final d.wt of CO₂-enrichedP.smithiiincreased 84% at 20°C, but only 4% at 35°C. Finald.wt ofB. graciliswas unaffected by CO₂at 20°C, but wasenhanced by 28% at 35°C. Root:shoot ratios remained relativelyconstant across CO₂levels, but increased inP. smithiiwith reductionin temperature. These partitioning results were adequately explainedby the theory of balanced root and shoot activity. Favourablegrowth temperatures led to CO₂-induced accumulations of TNCin leaves of both species, and in stems ofP. smithii, whichgenerally reflected responses of above-ground d.wt partitioningto CO₂. However, CO₂-induced decreases in plant tissue N concentrationswere more evident forP. smithii. Roots of CO₂-enrichedP. smithiihadgreater total N content at 20°C, an allocation of N below-groundthat may be an especially important adaptation for C₃plants.Tissue N contents ofB. graciliswere unaffected by CO₂. Resultssuggest CO₂enrichment may lead to reduced N requirements forgrowth in C₃plants and lower shoot N concentration, especiallyat favourable growth temperatures. Acclimation to CO₂; blue grama; Bouteloua gracilis ; carbohydrate; climate change; global change; grass; growth; growth temperature optima; nitrogen; N uptake; Pascopyrum smithii; western wheatgrass     

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