Homeostatic gas-exchange parameters inferred from 13C/12C in tree rings of conifers

The CO₂ concentration of the atmosphere has increased by almost 30% in the past two centuries, with most of the increase (>5 Pa) during the past 60 years. Controlled environment studies of crop plants dependent on the C₃ photosynthetic pathway indicate that an increase of this magnitude would enhance net photosynthesis, reduce stomatal conductance, and increase the difference in CO₂ concentration across the stomata, i.e., CO₂ concentration outside the leaf to that within (c _a-c _i). Here we report evidence, based on stable isotope composition of tree rings from three species of field-grown, native conifer trees, that the trees have indeed responded. However, rather than increasing c _a-c _i, intercellular CO₂ concentrations have shifted upward to match the rise in atmospheric concentrations, holding c _a-c _i constant. No differences were detected among Douglas-fir (Pseudotsuga menziesii), ponderosa pine (Pinus ponderosa), or western white pine (Pinus monticola). The values of c _a-c _i were inferred from stable carbon isotope ratio (¹³C) of tree ring holocellulose adjusted for the 0.6–2.6 difference between holocellulose and whole sapwood. The cellulose extraction removed contaminants deposited in the tree ring after it formed and the adjustment corrected for the enrichment of cellulose relative to whole tissue. The whole sapwood values were then adjusted for bublished estimates of past atmospheric ¹³CO₂ and CO₂ concentrations. To avoid confounding tree age with CO₂, cellulose deposited by saplings in the 1980s was compared to cellulose deposited in the inner rings of nature trees when the mature trees were saplings, between 1910–1929 and 1941–1970; thus saplings were compared to saplings. In a separate analysis, the juvenile effect, which describes the tendency for ¹³C to increase in the first decades of a tree's life, was quantified independent of source CO₂ effects. This study provides evidence that conifers have undergone adjustments in the intercellular CO₂ concentration that have maintained c _a-c _i constant. Based on these results and others, we suggest that c _a-c _i, which has also been referred to as the intrinsic water-use efficiency, should be considered a homeostatic gas-exchange set point for these conifer species.     

Effect of changes in water content on photosynthesis,transpiration and discrimination against 13CO2 and C18O16O in Pleurozium and Sphagnum

Photosynthetic gas exchange characteristics of two common boreal forest mosses, Sphagnum (section acutifolia) and Pleurozium schreberi, were measured continuously during the time required for the moss to dry out from full hydration. Similar patterns of change in CO₂ assimilation with variation in water content occurred for both species. The maximum rates of CO₂ assimilation for Sphagnum (approx. 7 mol m^–2 s^–1) occurred at a water content of approximately 7 (fresh weight/dry weight) while for Pleurozium the maximum rate (approx. 2 mol m^–2 s^–1) occurred at a water content of approximately 6 (fresh weight/dry weight). Above and below these water contents CO₂ assimilation declined. In both species total conductance to water vapour (expressed as a percentage of the maximum rates) remained nearly constant at a water content above 9 (fresh weight/dry weight), but below this level declined in a strong linear manner. Short-term, on-line ¹³CO₂ and C¹⁸O¹⁶O discrimination varied substantially with changes in moss water content and associated changes in the ratio of chloroplast CO₂ to ambient CO₂ partial pressure. At full hydration (maximum water content) both Sphagnum and Pleurozium had similar values of ¹³CO₂ discrimination (approx. 15). Discrimination against ¹³CO₂ increased continuously with reductions in water content to a maximum of 27 in Sphagnum and 22 in Pleurozium. In a similar manner C¹⁸C¹⁶O discrimination increased from approximately 30 at full hydration in both species to a maximum of 150 in Sphagnum and 90 in Pleurozium, at low water content. The observed changes in C¹⁸O¹⁶O were strongly correlated to predictions of a mechanistic model of discrimination processes. Field measurements of moss water content suggested that photosynthetic gas exchange by moss in the understory of a black spruce forest was regularly limited by low water content.     

Effect at the ecosystem level of elevated atmospheric CO2in an aquatic microcosm

We studied the responses of an aquatic microcosm in two different eutrophic conditions to elevated atmospheric CO₂concentration. We used microcosms, consisting of Escherichia coli(bacteria), Tetrahymena thermophila(protozoa) and Euglena gracilis(algae), in salt solution with 50 and 500 mg l^–1of proteose peptone (eutrophic and hypereutrophic conditions, respectively) under ambient and elevated CO₂(1550±100 l l^–1) conditions. The density of E. gracilisincreased significantly under elevated CO₂in both eutrophic and hypereutrophic microcosms. In the eutrophic microcosm, the other elements were not affected by elevated CO₂. In the hypereutrophic microcosm, however, the concentrations of ammonium and phosphate decreased significantly under elevated CO₂. Furthermore, the density of T. thermophilawas maintained in higher level than that in the microcosm with ambient CO₂and the density of E. coliwas decreased by CO₂enrichment. Calculating the carbon biomasses of T. thermophilaand E. colifrom their densities, the changes in their biomasses by CO₂enrichment were little as compared with large increase of E. graciliscarbon biomass converted from chlorophyll a. From the responses to elevated CO₂in the subsystems of the hypereutrophic microcosm consisting of either one or two species, the increase of E. graciliswas a direct effect of elevated CO₂, whereas the changes in the density of E. coliand T. thermophilaand the decreases in the concentration of ammonium and phosphate are considered to be indirect effects rather than direct effects of elevated CO₂. The indirect effects of elevated CO₂were prominent in the hypereutrophic microcosm.     

Leaf and canopy responses to elevated CO2 in a pine forest under free-air CO2 enrichment

Physiological responses to elevated CO₂ at the leaf and canopy-level were studied in an intact pine (Pinus taeda) forest ecosystem exposed to elevated CO₂ using a free-air CO₂ enrichment (FACE) technique. Normalized canopy water-use of trees exposed to elevated CO₂ over an 8-day exposure period was similar to that of trees exposed to current ambient CO₂ under sunny conditions. During a portion of the exposure period when sky conditions were cloudy, CO₂-exposed trees showed minor (7%) but significant reductions in relative sap flux density compared to trees under ambient CO₂ conditions. Short-term (minutes) direct stomatal responses to elevated CO₂ were also relatively weak (5% reduction in stomatal aperture in response to high CO₂ concentrations). We observed no evidence of adjustment in stomatal conductance in foliage grown under elevated CO₂ for nearly 80 days compared to foliage grown under current ambient CO₂, so intrinsic leaf water-use efficiency at elevated CO₂ was enhanced primarily by direct responses of photosynthesis to CO₂. We did not detect statistical differences in parameters from photosynthetic responses to intercellular CO₂ (A _net-C _i curves) for Pinus taeda foliage grown under elevated CO₂ (550 mol mol^–1) for 50–80 days compared to those for foliage grown under current ambient CO₂ from similar-sized reference trees nearby. In both cases, leaf net photosynthetic rate at 550 mol mol^–1 CO₂ was enhanced by approximately 65% compared to the rate at ambient CO₂ (350 mol mol^–1). A similar level of enhancement under elevated CO₂ was observed for daily photosynthesis under field conditions on a sunny day. While enhancement of photosynthesis by elevated CO₂ during the study period appears to be primarily attributable to direct photosynthetic responses to CO₂ in the pine forest, longer-term CO₂ responses and feedbacks remain to be evaluated.     

The specific radioactivity of glycolic acid in relation to the specific activity of carbon dioxide evolved in light in photosynthesizing sunflower leaves

Attached leaves of sunflower (Helianthus annuus L.) were exposed to ¹⁴CO₂ during steady-state photosynthesis for 2 to 30 min in 345 l/l CO₂ and 21% O₂ at 29° C and a light intensity of 1300 E m^-2s^-1. Glycolic acid was extracted with water and diethyl ether, and was determined in the aqueous residue by high-pressure liquid column chromatography. The relative specific radioactivity of the glycolic acid synthesized during photosynthesis reached about 100% after 30 min of photosynthesis and was almost equal to that of the CO₂ evolved during photorespiration, their ratio at all times being nearly one. These results provide strong in-vivo evidence that the glycolic acid is the substrate for CO₂ evolved by sunflower leaves in light.     

Phototrophic growth in the lumostat: a photo-bioreactor with on-line optimization of light intensity

A novel approach is described for the growth of phototrophic microorganisms in batch culture in laboratory-scale photo-bioreactors. Pure CO₂ is added separately to the aeration gas in a closed loop and the rate of photosynthetic activity is monitored continuously by recording the amount of CO₂ added in order to maintain constant pH. These activity measurements are used to control the intensity of illumination by varying the voltage applied to a bank of fluorescent tubes. The intensity of illumination is maintained at the value giving the maximal rate of photosynthesis while photoinhibition due to excess light is avoided. Since the light intensity received by the individual cells is maintained at the optimal value we term the device a lumostat.Measurements of photosynthetic activity by monitoring CO₂ addition were in excellent agreement with off-line measurements of cell carbon as long as corrections were made for diffusion loss through the walls of the tubing. Exponential growth of a thermophilic strain of the cyanobacteriumSynechococcus was obtained for 7 generations with maximum cell densities of 8 × 10⁷ cells mL^–1. The productivity of the lumostat is superior to that of batch cultures at any fixed light intensity.     

Long-term photosynthetic response in single leaves of A C3 and C4 salt marsh species grown at elevated atmospheric CO2 in situ

Summary Mono-specific communities of the C₃ sedge, Scirpus olneyi and the C₄ grass, Spartina patens, were exposed to normal ambient or elevated CO₂, (ca. 680 l l^–1) throughout the 1987 and 1988 growing seasons in open-top field chambers located on a tidal marsh. Single stems of C₃ plants grown in ambient or elevated CO₂ showed an increased photosynthetic rate when tested at elevated CO₂ for both seasons. This increase in photosynthetic response in the C₃ species was maintained throughout the 1987 and 1988 growing season. The stimulation of photosynthesis with elevated CO₂ appeared to increase as temperature increased and decreased as photosynthetic photon flux (PPF) increased. Analysis of the photosynthetic response of the C₃ species during the 1988 season indicated that significant differences in light-saturated photosynthetic rate between ambient and elevated CO₂ conditions continued until October. In contrast to the C₃ sedge, the C₄ grass showed no significant photosynthetic increase to elevated CO₂ except at the beginning of the 1988 season.     

Inorganic-carbon transport in some marine eukaryotic microalgae

Inorganic-carbon transport was investigated in the eukaryotic marine microalgaeStichococcus minor, Nannochloropsis oculata and aMonallantus sp. Photosynthetic O₂ evolution at constant inorganic-carbon concentration but varying pH showed thatS. minor had a greater capacity for CO₂ rather than HCO ₃ ^– utilization but forN. oculata andMonallantus HCO ₃ ^– was the preferred source of inorganic carbon. All three microalgae had a low affinity for CO₂ as shown by the measurement of inorganic-carbon-dependent photosynthetic O₂ evolution at pH 5.0. At pH 8.3, where HCO ₃ ^– is the predominant form of inorganic carbon, the concentration of inorganic carbon required for half-maximal rate of photosynthetic O₂ evolution [K _0.5 (CO₂)] was 53 M forMonallantus sp. and 125 M forN. oculata, values compatible with HCO ₃ ^– transport. Neither extra- nor intracellular carbonic anhydrase was detected in these three microalgal species. It is concluded that these microalgae lack a specific transport system for CO₂ but that HCO ₃ ^– transport occurs inN. oculata andMonallantus, and in the absence of intracellular carbonic anhydrase the conversion of HCO ₃ ^– to CO₂ may be facilitated by the internal pH of the cell.     

Carbon monoxide pathway enzyme activities in a thermophilic anaerobic bacterium grown acetogenically and in a syntrophic acetate-oxidizing coculture

We recently isolated an acetate-oxidizing rodshaped eubacterium (AOR) which was capable of oxidizing acetate to CO₂ when grown in coculture with the hydrogenotrophic methanogen Methanobacterium sp. strain THF. The AOR was also capable of growing axenically on H₂CO₂ which it converted to acetate. Previous results for the acetate oxidizing coculture showed isotopic exchange between acetate and CO₂, suggesting that the AOR was using a pathway for acetate oxidation resembling a reveral of the acetogenic (carbon monoxide) pathway. In this study, it was found that production of ¹⁴CO₂ from ¹⁴CH₃COO^- by the coculture was inhibited by 200 M cyanide, while methanogenesis from H₂–CO₂ was unaffected, implying the involvement of carbon monoxide dehydrogenase (CODH) in acetate oxidation. CODH was present at 0.055 mol methyl viologen reduced min^-1 mg^-1 protein in extracts of Methanobacterium sp. strain THF, but was present in higher levels in the acetate oxidizing coculture and in the AOR grown axenically and on H₂–CO₂ (2.0 and 6.4 mol min^-1 mg^-1 protein respectively). Anaerobic activity stains for CODH in native polyacrylamide gels from the AOR coculture showed components co-migrating with bands from both organisms, as well as an additional band in extracts of the coculture. Formate dehydrogenase (FDH) was present in both the AOR coculture and monoculture but not in extracts of H₂–CO₂ grown cells of Methanobacterium sp. strain THF. Formyltetrahydrofolate (FTHF) synthetase was not detectable in extracts of the AOR monoculture or coculture, although it was found in high amounts in extracts of H₂–CO₂ grown cells of the thermophilic acetogen Acetogenium kivui. Extracts of H₂–CO₂ grown cells of the AOR showed a fluorescence spectrum typical of pterin derivatives. Bioassay for folates showed levels to be at anabolic rather than catabolic levels. It is possible that the AOR uses pterins distinct from folate for catabolism. Isocitrate dehydrogenase, a citric acid cycle enzyme, was also present in the AOR, but at anabolic levels and -ketoglutarate dehydrogenase was not detectable.Abbreviations (AOR) acetate-oxidizing rod - (CODH) carbon monoxide dehydrogenase - (FDH) formate dehydrogenase - (FTHF) formyltetrahydrofolate     

Gas exchange of ears of cereals in response to carbon dioxide and light

One cultivar each of spring wheat (Triticum aestivum L. cv. Arkas), oat (Avena sativa L. cv. Lorenz), and barley (Hordeum vulgare L. cv. Aramir) was chosen in order to study the relative contributions of individual bracts to the gas exchange of whole ears. The distribution and frequency of the stomata on the bracts were examined. Gas exchange was measured at normal atmospheric CO₂ (330 bar) and at high CO₂ (2000 bar) on intact ears and on ears from which glumes or lemmas and pleae (wheat and oat) or awns (barley) had been removed.The relative contribution to the gas exchange of the whole organ is highest for the awns of barley ears. In wheat, the contribution of the glumes is slightly higher than that of the inner bracts before anthesis. Two weeks after anthesis the inner bracts contribute more than the glumes. This tendency of increasing importance of the inner bracts is also found in oat ears, but the relative amount of CO₂ uptake by the glumes is higher than in wheat. These changes during ontogeny result from the better supply of light to the inner bracts caused by opening of the ears' structures during grain filling, which in part compensates for the decreasing photosynthetic capacity.The ratio of the photosynthesis rate at high CO₂ to that at normal CO₂ is lower for the glumes of oat and for the awns of barley than for the other bracts.Abbreviations A³³⁰, A²⁰⁰⁰ net photosynthesis rate, A³³⁰ at normal atmospheric CO₂ (330 bar), A²⁰⁰⁰ at high CO₂ (2000 bar) - PPFD photosynthetic photon flux density - p^c intercellular partial pressure of CO₂     

Interactions of glycolate-, HCO 3 - -, Cl--, and H+-balance of Scenedesmus obliquus

Excretion and absorption of glycolate by young cells of Scenedesmus obliquus (Turp.) Krüger strain D₃ grown synchronously with 2% CO₂ was compared after no pretreatment with air (CO₂-adapted) or after a 2 h adaptation to normal air (0.03% CO₂) (air-adapted). At 21% O₂, excretion occurred only from CO₂-adapted cells at high pH (pH 8.0). Under conditions where no excretion occurred, external glycolate (0.2 mM) was taken up by both air-and CO₂-adapted cells at a much faster rate at pH 5 than at pH 8. The uptake was accompanied by an apparent stoichiometric uptake of H⁺. CO₂-adapted algae exhibited high uptake rates that were even higher in the dark than in the light. Air-adapted algae showed high uptake rates in the light but only minimal uptake in the dark. The uptake rate was decreased to about 1/3 with 5% CO₂, except with CO₂-adapted cells in the light, in which a slight stimulation occurred. Cl^- ions inhibited glycolate uptake by air-adapted cells in the light; conversely, light-stimulated Cl^- uptake of these cells was inhibited by glycolate. A hypothesis is discussed according to which the internal pH regulates the uptake and release of Cl^-, HCO ₃ ^- , and glycolate.Abbreviations DCMU 3-(3,4 dichlorophenyl)-1, 1-dimethyl urea - FCCP carbonyl cyanide p-trifluoro-methoxyphenylhydrazone - HEPES 2-(4-(2-hydroxyethyl)-piperazinyl) ethanesulfonic acid - HPMS -hydroxypyridinemethanesulfonate - MES 2-morpholinoethanesulfonic acid - PCV packed cell volume     

Developmental Change in CO2 Compensation Concentrations in Spartina alterniflora Results from Sigmoidal Photosynthetic CO2 Responses

We investigated seasonal patterns of photosynthetic responses to CO₂ concentrations in Spartina alterniflora Loisel, an aerenchymous halophyte grass, from a salt marsh of the Bay of Fundy (NB, Canada), and from plants grown from rhizome in controlled-environment chambers. From late May to August, CO₂ compensation concentrations () of field-grown leaves varied between 2.5–10.7 cm³(CO₂) m^–3, with a mean of 5.4 cm³(CO₂) m^–3. From September onwards field leaves showed CO₂ compensation concentrations from 6.6–21.1 cm³(CO₂) m^–3, with a mean of 13.1 cm³ m^–3 well into the C₃–C₄ intermediate range. The seasonal variability in did not result from changing respiration, but rather from a sigmoidal response of net photosynthetic rate (P _N) to applied CO₂ concentration, found in all tested leaves but which became more pronounced late in the season. One explanation for the sigmoidal response of P _N to external CO₂ concentration could be internal delivery of CO₂ from roots and rhizomes to bundle sheath cells via the aerenchyma, but the sigmoidal responses in S. alterniflora persisted out to the tips of leaves, while the aerenchyma extend only to mid-leaf. The sigmoidicity persisted when CO₂ response curves were measured from low to high CO₂, or from high to low CO₂, and even when prolonged acclimation times were used at each CO₂ concentration.     

Growth dynamics and population development in an alpine grassland under elevated CO2

Leaf expansion, population dynamics and reproduction under elevated CO₂ were studied for two dominant and four subdominant species in a high alpine grassland (2500 above sea level, Swiss Central Alps). Plots of alpine heath were exposed to 335 l l^-1 and 680 l l^-1 CO₂ in open-top chambers over three growing seasons. Treatments also included natural and moderately improved mineral nutrient supply (40 kg N ha^-1 year^-1 in an NPK fertilizer mix). Seasonal dynamics of leaf expansion, which was studied for the dominant graminoid Carex curvula only, were not affected by elevated CO₂ during two warm seasons or during a cool season. Improved nutrient supply increased both the expansion rate and the duration of leaf growth but elevated CO₂ did not cause any further stimulation. Plant and tiller density (studied in all species) increased under elevated CO₂ in the codominant Leontodon helveticus and the subdominant Trifolium alpinum, remained unchanged in two other minor species Poa alpina and Phyteuma globulariifolium, and decreased in Carex curvula. In Potentilla aurea elevated CO₂ compensated for a natural decline in shoot number. By year 3 the number of fertile shoots in Leontodon and individual seed weight in Carex were slightly increased under elevated CO₂, indicating CO₂ effects on sexual reproduction in these two dominant species. The results suggest that the effects of elevated CO₂ on the population dynamics of the species studied were not general, but species-specific and rather moderate effects. However, the reduction of tiller density in Carex curvula, in contrast to the increases observed in Leontodon helveticus and Trifolium alpinum, indicates that elevated CO₂ may negatively affect the abundance of the species most characteristic of this alpine plant community.     

A circadian rhythm in the level of carbon dioxide compensation in Bryophyllum fedtschenkoi with zero values during the transient

Summary Detached shoots of Bryophyllum fedtschenkoi maintained in a closed system in the light exhibited an endogenous circadian rhythm in CO₂ compensation. The rhythm was sensitive to changes in light intensity and temperature. At 15° C it damped rapidly in light of 78 J m^-2 s^-1, but at 10° C a rhythm of considerable amplitude was evident at this same light intensity. During the transient (i.e. the temporary state of the rhythm before it acquired its steady state) low compensation values between 0 and 5 ppm CO₂ were achieved. When the plants were maintained at a higher light intensity prior to the measurements, the period of low compensation during the transient was extended, and zero values were obtained under some conditions.Studies of gas exchange at opposite phases of the rhythm revealed: (i) that the rate of uptake of ¹⁴CO₂ differed, both in light and darkness (the epidermis was removed during these observations to avoid interference from stomatal rhythms); (ii) photorespiration, estimated by extrapolation of the graph relating photosynthetic rate and CO₂ concentration, was highest during the peaks of the rhythm in CO₂ compensation; (iii) estimates of the capacity for photorespiration by the glycine-1-¹⁴C assay indicated highest values during the troughs of the rhythm. These findings are discussed in relation to the C₄-acid metabolism of this species. Low CO₂ compensation is probably due to the activity of phosphoenolpyruvate carboxylase and not to the absence of processes involving CO₂ evolution.     

Combined Effects of CO2 and O3 on Antioxidative and Photoprotective Defense Systems in Needles of Ponderosa Pine

Ponderosa pine (Pinus ponderosa Dougl. ex Laws.) seedlings were exposed to near ambient or elevated CO₂ (average concentrations during the last growing season 446 versus 699 mol mol^–1), combined with low or elevated O₃ for three seasons. Ozone exposure during the last growing season (accumulated dose above threshold 0.06 mol mol^–1) was 0.05 versus 26.13 mol mol^–1 h. Needles of the youngest age class were harvested after the dormancy period. Ozone exposure decreased needle contents of chlorophyll a, chlorophyll b, and ascorbate, and resulted in a more oxidized total ascorbate and a more de-epoxidized xanthophyll cycle pool irrespective of the CO₂ level. Trees under elevated CO₂ had a more oxidized glutathione pool and lower chlorophyll a content. Contents of glutathione, tocopherol, and carotenoids were not affected by the CO₂ or O₃ treatments. There were no interactive effects between elevated CO₂ and elevated O₃ on any of the parameters measured. The results suggest that elevated atmospheric CO₂ concentration does not compensate for ozone stress by increasing antioxidative capacity in ponderosa pine.     

Use of carbon dioxide enrichment to obtain adult morphology of grapevinein vitro

A procedure has been developed forin vitro propagation ofVitis vinifera Pinot noir from lateral-bud cuttings under high CO₂ concentration (1200 µmol mol^–1). Because of inhibition of rooting by CO₂, this procedure requires a rooting pre-culture of explants on medium with sucrose before the CO₂-enriched culture on sucrose-free medium. Shoot growth was enhanced by CO₂ enrichment as a result of both a higher rate of leaf production and greater internode elongation. Leaf expansion and tendril growth were promoted and better rooting was obtained. The more significant effect of CO₂ enrichment was to promote adult morphology with, in particular, the tendril pattern. Thus, for the first time, grapevine plants have been producedin vitro without typical juvenile characteristics. CO₂ enrichment appears to be an interesting process to improve thein vitro propagation of grapevines.     

Variable HCO 3 - affinity of Elodea canadensis Michaux in response to different HCO 3 - and CO2 concentrations during growth

Summary Elodea canadensis grows over a wide range of inorganic carbon, nutrient, and light conditions in lakes and streams. Affinity for HCO ₃ ^- use during photosynthesis ranged from strong to weak in Elodea collected from seven localities with different HCO ₃ ^- and CO₂ concentrations. The response to HCO ₃ ^- was also very plastic in plants grown in the laboratory at high HCO ₃ ^- concentrations and CO₂ concentrations varying from 14.8 to 2,200 M. Bicarbonate affinity was markedly reduced with increasing CO₂ concentrations in the growth medium so that ultimately HCO ₃ ^- use was not detectable. High CO₂ concentrations also decreased CO₂ affinity and induced high CO₂ compensation points (360M CO₂) and tenfold higher half-saturation values (800 M CO₂).The variable HCO ₃ ^- affinity is probably environmentally based. Elodea is a recently introduced species in Denmark, where it reproduces only vegetatively, leaving little opportunity for genetic variation. More important, local populations in the same water system had different HCO ₃ ^- affinities, and a similar variation was created by exposing one plant collection to different laboratory conditions.Bicarbonate use enabled Elodea to photosynthesize rapidly in waters of high alkalinity and enhanced the carbon-extracting capacity by maintaining photosynthesis above pH 10. On the other hand, use of HCO ₃ ^- represents an investment in transport apparatus and energy which is probably not profitable when CO₂ is high and HCO ₃ ^- is low. This explanation is supported by the findings that HCO ₃ ^- affinity was low in field populations where HCO ₃ ^- was low (0.5 and 0.9 m M) or CO₂ was locally high, and that HCO ₃ ^- affinity was suppressed in the laboratory by high CO₂ concentrations.Abbreviations DIC dissolved inorganic carbon (CO₂+ HCO ₃ ^- +CO ₃ ^- ) - CO₂ compensation point - K _1/2 apparent halfsaturation constant - P_{HCO 3} ^– interpolated photosynthesis in pure HCO ₃ ^- and zero CO₂ - P_max photosynthetic rate under carbon and light saturation     

Short-term CO2 exchange response to temperature,irradiance, and CO2 concentration in strawberry

Relative importance of short-term environmental interaction and preconditioning to CO₂ exchange response was examined in Fragaria ananasa (strawberry, cv. Quinault). Tests included an orthogonal comparison of 15 to 60-min and 6 to 7-h exposures to different levels of temperature (16 to 32°C), photosynthetically active radiation (PAR, 200 to 800 E m² s^-1), and CO₂ (300 to 600 l/l) on successive days of study. Plants were otherwise maintained at 21°C, 300 E m² s^-1 PAR and 300–360 l/l CO₂ as standard conditions. Treatment was restricted to the mean interval of 14 h daily illumination and the first 3–4 days of each test week over a 12-week cultivation period. CO₂ exchange rates were followed with each step-change in environmental level including ascending/descending temperature/PAR within a test period, initial response at standard conditions on successive days of testing, and measurement at reduced O₂. Response generally supported prior concepts of leaf biochemical modeling in identifying CO₂ fixation as the major site of environmental influence, while overall patterns of whole plant CO₂ exchange suggested additional effects for combined environmental factors and preconditioning. These included a positive interaction between temperature and CO₂ concentration on photosynthesis at high irradiance and a greater contribution by dark respiration at lower PAR than previously indicated. The further importance of estimating whole plant CO₂ exchange from repetitive tests and measurements was evidenced by a high correlation of response to prior treatment both during the daily test period and on consecutive days of testing.Abbreviations C₃ plant a plant in which the product of CO₂ fixation is a 3-carbon acid (3-phosphoglyceric acid) - IRGA intra-red gas analyzer - PAR photosynthetically active radiation - RH relative humidity - RuBisCO ribulose-1,5-bisphosphate carboxylase/oxygenase Reference to a company and/or product named by the Department is only for purposes of information and does not imply approval or recommendation of the product to the exclusion of others which may also be suitable.     

Growth,photosynthesis and photorespiration of Lemna gibba: response to variations in CO2 and O2 concentrations and photon flux density

Dry weight and Relative Growth Rate of Lemna gibba were significantly increased by CO₂ enrichment up to 6000 l CO₂ l^–1. This high CO₂ optimum for growth is probably due to the presence of nonfunctional stomata. The response to high CO₂ was less or absent following four days growth in 2% O₂. The Leaf Area Ratio decreased in response to CO₂ enrichment as a result of an increase in dry weight per frond. Photosynthetic rate was increased by CO₂ enrichment up to 1500 l CO₂ l^–1 during measurement, showing only small increases with further CO₂ enrichment up to 5000 l CO₂ l^–1 at a photon flux density of 210 mol m^–2 s^–1 and small decreases at 2000 mol m^–1 s^–1. The actual rate of photosynthesis of those plants cultivated at high CO₂ levels, however, was less than the air grown plants. The response of photosynthesis to O₂ indicated that the enhancement of growth and photosynthesis by CO₂ enrichment was a result of decreased photorespiration. Plants cultivated in low O₂ produced abnormal morphological features and after a short time showed a reduction in growth.     

Carbon fixation in eucalypts in the field

Summary The rate of CO₂ assimilation at light saturation and an intercellular CO₂ concentration of 350 l l^-1 (photosynthetic capacity), measured in leaves of Eucalyptus pauciflora, E. behriana, E. delegatensis and Acacia melanoxylon, declined over the course of cloudless days under naturally varying environmental conditions as well as under constant optimal conditions for high CO₂ uptake. Since the capacity did not recover during the light period, it was different from the midday depression of gas exchange. The change appeared to be caused neither by the diurnal variation of total leaf water potential, by photoinhibition of redox-reaction centres in photosystems nor by changes in the intrinsic properties of Ribulose-bisphosphate carboxylase-oxygenase. The decline was more pronounced in winter than in summer. It was related to the duration of illumination or the cumulative carbon gain. It was reversible in the following dark phase, and it did not occur on changeable days with short peaks of high light.Despite the decline in photosynthetic capacity, the initial slope of the CO₂ response of net photosynthesis, as obtained at low intercellular CO₂ concentrations, remained constant during the day, but declined at night when photosynthetic capacity recovered. In all cases stomatal conductance varied in parallel with photosynthetic capacity. The relevance of changes in photosynthetic capacity for the intercellular CO₂ concentration is discussed.Abbreviations and symbols A CO₂ assimilation - ABA abscisic acid - A_c350 photosynthetic capacity at c_i=350l l^-1 - c_i intercellular CO₂ concentration - g leaf conductance to water vapour - I photon flux density (irradiance) - P air pressure - P_i inorganic phosphate - R_d net CO₂ release at _* - Rubisco Ribulose-bisphosphate carboxylase-oxygenase - RuBP Ribulose-bisphosphate - T leaf temperature - w leaf-to-air water vapour concentration difference - A/c_i carboxylation efficiency at low c_i - _* light-independent CO₂ compensation point - total leaf water potential