Economy of water, carbon, and nitrogen in the developing cowpea fruit

The nutritional economy of the fruit of cowpea (Vigna unguiculata (L.) Walp cv Vita 3) was assessed quantitatively from intake and utilization of carbon, nitrogen, and water. Fruits failed to make net gains of CO₂ from the atmosphere during daytime, although pod photosynthesis did play a role in the fruit's carbon economy by refixing a proportion of the fruit's respired CO₂. Of every 100 units by weight of carbon entering the fruit, 70.4 were finally incorporated into seeds, 10.3 remained as nonmobilizable material in pod walls, and the remaining 19.3 were lost in fruit respiration. Phloem supplied 97% of the fruit's carbon and 72% of its nitrogen. The xylem contribution of nitrogen occurred mainly in early growth. Ninety-six% of the fruit's nitrogen was incorporated into seeds, approximately 10% of this mobilized from the senescing pod. The mean transpiration ratio of the fruit was very low—8 milliliters water transpired per gram dry matter accumulated. Models of carbon, nitrogen, and water flow were constructed for the two consecutive 11 day periods of fruit development, and indicated a considerably greater entry of water through xylem and phloem than could be accounted for in changes in fruit tissue water and transpiration loss. This discrepancy was greater in the second half of fruit growth and was interpreted as evidence that a significant fraction of the water entering the fruit through phloem cycled back to the parent plant via the xylem.     

Photosynthesis by inflated pods of a desert shrub,Isomeris arborea

Summary The photosynthetic capacity and carbon metabolism of the fruits of Isomeris arborea (Capparidaceae), an evergreen shrub endemic to the desert and coastal habitats of Southern California and Baja California, are described. The inflated structure of the pods of I. arborea provides a model system for experimental studies of fruit photosynthesis in native plants since the gas concentration of the internal space can be manipulated and monitored separately from the external pod environment. CO₂ released by seed respiration is partially contained in the inner gas space of the pods, resulting in an elevated CO₂ environment inside the fruit (500 to 4000 mol mol^–1 depending on the stage of fruit development). A portion of this CO₂ is assimilated by the inner layers of the pericarp, but a larger fraction leaks out. The photosynthetic layers of the pericarp use two different sources of CO₂: the exocarp fixes exogenous CO₂ while the endocarp fixes CO₂ released by seed respiration into the pod cavity. Even though the total weight of the fruit increases during development, the combined rates of fixation of externally and internally supplied CO₂ remained constant (10–11 mol CO₂ pod^–1 h^–1). After the pods attain maximum volume, the major change in gas exchange that takes place during fruit growth is a gradual increase in the amount of respiratory CO₂ released by the seeds. This shifts the CO₂ balance of the fruit from positive, in young fruits, to negative in mature fruits. Pericarp photosynthesis helped support not only the cost of fruit maintenance, but also the cost of fruit growth, particularly during the first stages of fruit development. During later fruiting stages insufficient carbon is fixed to fully supply either respiration or growth.     

Interaction of carbon dioxide and ethylene in overcoming thermodormancy of lettuce seeds

The combination of ethylene with CO₂ will completely overcome the thermodormancy of lettuce (Lactuca sativa L.) seeds at 35 C. This combination is effective if it is added to seeds either at the start or after several days of imbibition. The action of ethylene is dependent upon the CO₂ level present in the atmosphere surrounding the seeds. When CO₂ is trapped by KOH the ethylene effect is essentially nil.     

Growth,reproductive phenology and yield responses of a potential biofuel plant,Jatropha curcas grown under projected 2050 levels of elevated CO2

Jatropha (Jatropha curcas) is a non‐edible oil producing plant which is being advocated as an alternative biofuel energy resource. Its ability to grow in diverse soil conditions and minimal requirements of essential agronomical inputs compared with other oilseed crops makes it viable for cost‐effective advanced biofuel production. We designed a study to investigate the effects of elevated carbon dioxide concentration ([CO₂]) (550 ppm) on the growth, reproductive development, source‐sink relationships, fruit and seed yield of J. curcas. We report, for the first time that elevated CO₂ significantly influences reproductive characteristics of Jatropha and improve its fruit and seed yields. Net photosynthetic rate of Jatropha was 50% higher in plants grown in elevated CO₂ compared with field and ambient CO₂‐grown plants. The study also revealed that elevated CO₂ atmosphere significantly increased female to male flower ratio, above ground biomass and carbon sequestration potential in Jatropha (24 kg carbon per tree) after 1 year. Our data demonstrate that J. curcas was able to sustain enhanced rate of photosynthesis in elevated CO₂ conditions as it had sufficient sink strength to balance the increased biomass yields. Our study also elucidates that the economically important traits including fruit and seed yield in elevated CO₂ conditions were significantly high in J. curcas that holds great promise as a potential biofuel tree species for the future high CO₂ world.     

Photosynthetic Carbon Assimilation in a Shootless Orchid, Chiloschista usneoides (DON) LDL: A Variant on Crassulacean Acid Metabolism

Photosynthetic carbon assimilation in the roots of a shootless orchid Chiloschista usneoides (DON) LDL involves the synthesis and accumulation of malic acid from CO₂ in darkness. Malic acid is consumed in the light.

The roots do not possess stomata or any means of diurnally regulating the diffusive conductance of the pathway between the internal gas phase of the plant and the atmosphere. Regulation of internal CO₂ concentration near to atmospheric levels avoids a large net loss of CO₂ to the atmosphere during malic acid consumption in the light.

The water-absorbing function of the velamen conflicts with the photosynthetic function of the roots. Plants with water-saturated velamina do not acquire CO₂ from the atmosphere at night.

     

Elevated CO2 atmosphere enhances production of defense-related flavonoids in soybean elicited by NO and a fungal elicitor

Increased atmospheric pollutants including carbon dioxide (CO₂) and nitric oxide (NO) have a large impact on vegetation, with detrimental or beneficial influences on plant growth and metabolism. Here, we evaluated the effect of an elevated CO₂ atmosphere on the production of soybean defensive secondary chemicals induced by NO and a fungal elicitor. We hypothesized that an excess of carbon may alter the production of specific flavonoids that were previously shown to be induced by NO in soybean cotyledons. Pots containing soybean seeds (Glycine max [L.] Merr.) were submitted to 380 and 760 μmol mol^?1 of atmospheric CO₂ in open-top chambers. After nine days, plantlets grown under these conditions were assessed for biochemical and physiological parameters. Defense-related flavonoids were evaluated in detached cotyledon diffusates elicited with two different NO donors and with the β-glucan elicitor from Phytophthora sojae. A CO₂-enriched atmosphere stimulated initial growth, photosynthetic assimilation, and an altered C/N ratio in soybean plantlets resulting in allocation of precursors into different branches of the phenylpropanoid pathway in the cotyledons. Under elevated CO₂, the biotic elicitor caused accumulation of phytoalexins (glyceollins) as the natural end products of the phenylpropanoid pathway. In contrast, elevated CO₂ combined with NO resulted in an increase of intermediates and diverted end products (daidzein—127%, coumestrol—93%, genistein—93%, luteolin—89% and apigenin—238%) with a concomitant increase of 1.5–3.0 times in the activity of enzymes related to their biosynthetic routes. These observations point to changes in the pool of defense-related flavonoids that are related to increased carbon availability in soybeans. This may alter the responsiveness of soybean plants to pathogens when they are grown in CO₂ atmospheric concentrations close to those predicted for the upcoming several decades.     

Photosynthetic Pod Wall of Pea (Pisum sativum L.): Distribution of Carbon Dioxide-fixing Enzymes in Relation to Pod Structure

The pod wall of pea (Pisum sativum L.) was shown to contain two distinct photosynthetic layers. The outer, comprising chlorenchyma of the mesocarp, captured CO₂ from the outside atmosphere; the inner, a chloroplast-containing epidermis lining the pod gas cavity, was involved in photoassimilation of the CO₂ released from respiring seeds.     

Banana ripening: implications of changes in glycolytic intermediate concentrations, glycolytic and gluconeogenic carbon flux, and fructose 2,6-bisphosphate concentration

In ripening banana (Musa sp. [AAA group, Cavendish subgroup] cv Valery) fruit, the concentration of glycolytic intermediates increased in response to the rapid conversion of starch to sugars and CO₂. Glucose 6-phosphate (G-6-P), fructose 6-phosphate (Fru 6-P), and pyruvate (Pyr) levels changed in synchrony, increasing to a maximum one day past the peak in ethylene synthesis and declining rapidly thereafter. Fructose 1,6-bisphosphate (Fru 1,6-P₂) and phosphoenolpyruvate (PEP) levels underwent changes dissimilar to those of G 6-P, Fru 6-P, and Pyr, indicating that carbon was regulated at the PEP/Pyr and Fru 6-P/Fru 1,6-P₂ interconversion sites. During the climacteric respiratory rise, gluconeogenic carbon flux increased 50- to 100-fold while glycolytic carbon flux increased only 4- to 5-fold. After the climacteric peak in CO₂ production, gluconeogenic carbon flux dropped dramatically while glycolytic carbon flux remained elevated. The steady-state fructose 2,6-bisphosphate (Fru 2,6-P₂) concentration decreased to ½ that of preclimacteric fruit during the period coinciding with the rapid increase in gluconeogenesis. Fru 2,6-P₂ concentration increased thereafter as glycolytic carbon flux increased relative to gluconeogenic carbon flux. It appears likely that the initial increase in respiration in ripening banana fruit is due to the rapid influx of carbon into the cytosol as starch is degraded. As starch reserves are depleted and the levels of intermediates decline, the continued enhancement of respiration may, in part, be maintained by an increased steady-state Fru 2,6-P₂ concentration acting to promote glycolytic carbon flux at the step responsible for the interconversion of Fru 6-P and Fru 1,6-P₂.     

Contribution of stem CO2 fixation to whole-plant carbon balance in nonsucculent species

In many plant species that remain leafless part of the year, CO₂ fixation occurring in green stems represents an important carbon gain. Traditionally, a distinction has been made between stem photosynthesis and corticular photosynthesis. All stem photosynthesis is, sensu stricto, cortical, since it is carried out largely by the stem cortex. We proposed the following nomenclature: stem net photosynthesis (SNP), which includes net CO₂ fixation by stems with stomata in the epidermis and net corticular CO₂ fixation in suberized stems, and stem recycling photosynthesis (SRP), which defines CO₂ ling in suberized stems. The proposed terms should reflect differences in anatomical and physiological traits. SNP takes place in the chlorenchyma below the epidermis with stomata, where the net CO₂ uptake occurs, and it resembles leaf photosynthesis in many characteristics. SRP is found in species where the chlorenchyma is beneath a well-developed stomata-free periderm and where reassimilation of internally respired CO₂ occurs. SNP is common in plants from desert ecosystems, rates reaching up to 60% of the leaf photosynthetic rate. SRP has been demonstrated in trees from temperate forests and it offsets partially a carbon loss by respiration of stem nonphotosynthetic tissues. Reassimilation can vary between 7 and 123% of respired CO₂, the latter figure implying net CO₂ uptake from the atmosphere. Both types of stem photosynthesis contribute positively to the carbon economy of the species, in which they occur; they are advantageous to the plant because they allow the maintenance of physiological activity during stress, an increase of integrated water use efficiency, and they provide the carbon source used in the production of new organs.     

Carbon Dioxide Fixation in the Carbon Economy of Developing Seeds of Lupinus albus (L.)

The effects of CO₂ concentration and illumination on net gas exchange and the pathway of ¹⁴CO₂ fixation in detached seeds from developing fruits of Lupinus albus (L.) have been studied.

Increasing the CO₂ concentration in the surrounding atmosphere (from 0.03 to 3.0% [v/v] in air) decreased CO₂ efflux by detached seeds either exposed to the light flux equivalent to that transmitted by the pod wall (500 to 600 micro-Einsteins per square meter per second) in full sunlight or held in darkness. Above 1% CO₂ detached seeds made a net gain of CO₂ in the light (up to 0.4 milligrams of CO₂ fixed per gram fresh weight per hour) but ¹⁴CO₂ injected into the gas space of intact fruits (containing around 1.5% CO₂ naturally) was fixed mainly by the pod and little by the seeds.

Throughout development seeds contained ribulose-1,5-bisphosphate carboxylase activity (EC 4.1.1.39), especially in the embryo (up to 99 micromoles of CO₂ fixed per gram fresh weight per hour) and phosphoenolpyruvate carboxylase (EC 4.1.1.31) in both testa (up to 280 micromoles of CO₂ fixed per gram fresh weight per hour) and embryo (up to 355 micromoles of CO₂ fixed per gram fresh weight per hour).

In kinetic experiments the most significant early formed product of ¹⁴CO₂ fixation in both light and dark was malate but in the light phosphoglyceric acid and sugar phosphates were also rapidly labeled. ¹⁴CO₂ fixation in the light was linked to the synthesis of sugars and amino acids but in the dark labeled sugars were not formed.

     

Air-sea carbon dioxide equilibrium: Will it be possible to use seaweeds for carbon removal offsets?

To limit global warming below 2°C by 2100, we must drastically reduce greenhouse gas emissions and additionally remove ~100–900 Gt CO₂ from the atmosphere (carbon dioxide removal, CDR) to compensate for unavoidable emissions. Seaweeds (marine macroalgae) naturally grow in coastal regions worldwide where they are crucial for primary production and carbon cycling. They are being considered as a biological method for CDR and for use in carbon trading schemes as offsets. To use seaweeds in carbon trading schemes requires verification that seaweed photosynthesis that fixes CO₂ into organic carbon results in CDR, along with the safe and secure storage of the carbon removed from the atmosphere for more than 100 years (sequestration). There is much ongoing research into the magnitude of seaweed carbon storage pools (e.g., as living biomass and as particulate and dissolved organic carbon in sediments and the deep ocean), but these pools do not equate to CDR unless the amount of CO₂ removed from the atmosphere as a result of seaweed primary production can be quantified and verified. The draw-down of atmospheric CO₂ into seawater is via air-sea CO₂ equilibrium, which operates on time scales of weeks to years depending upon the ecosystem considered. Here, we explain why quantifying air-sea CO₂ equilibrium and linking this process to seaweed carbon storage pools is the critical step needed to verify CDR by discrete seaweed beds and nearshore and open ocean aquaculture systems prior to their use in carbon trading.     

Photosynthetic and Respiratory Activity of Fruiting Forms within the Cotton Canopy

The supply of photosynthates by leaves for reproductive development in cotton (Gossypium hirsutum L.) has been extensively studied. However, the contribution of assimilates derived from the fruiting forms themselves is inconclusive. Field experiments were conducted to document the photosynthetic and respiratory activity of cotton leaves, bracts, and capsule walls from anthesis to fruit maturity. Bracts achieved peak photosynthetic rates of 2.1 micromoles per square meter per second compared with 16.5 micromoles per square meter per second for the subtending leaf. However, unlike the subtending leaf, the bracts did not show a dramatic decline in photosynthesis with increased age, nor was their photosynthesis as sensitive as leaves to low light and water-deficit stress. The capsule wall was only a minor site of ¹⁴CO₂ fixation from the ambient atmosphere. Dark respiration by the developing fruit averaged −18.7 micromoles per square meter per second for 6 days after anthesis and declined to −2.7 micromoles per square meter per second after 40 days. Respiratory loss of CO₂ was maximal at −158 micromoles CO₂ per fruit per hour at 20 days anthesis. Diurnal patterns of dark respiration for the fruit were age dependent and closely correlated with stomatal conductance of the capsule wall. Stomata on the capsule wall of young fruit were functional, but lost this capacity with increasing age. Labeled ¹⁴CO₂ injected into the fruit interior was rapidly assimilated by the capsule wall in the light but not in the dark, while fiber and seed together fixed significant amounts of ¹⁴CO₂ in both the light and dark. These data suggest that cotton fruiting forms, although sites of significant respiratory CO₂ loss, do serve a vital role in the recycling of internal CO₂ and therein, function as important sources of assimilate for reproductive development.     

Pollen performance of Raphanus sativus (Brassicaceae) declines in response to elevated [CO2]

Although increases in atmospheric [CO₂] are known to affect plant physiology, growth and reproduction, understanding of these effects is limited because most studies of reproductive consequences focus solely on female function. Therefore, we examined the effects of CO₂ enrichment on male function in the annual Raphanus sativus. Pollen donors grown under elevated [CO₂] initially sired a higher proportion of seeds per fruit than ambient [CO₂]-grown plants when each was tested against two different standard competitors; however, by the end of the 5-month experiment, these pollen donors sired fewer seeds than ambient [CO₂]-grown plants and produced a lower proportion of viable pollen grains. The results of this experiment confirm that elevated [CO₂] can alter reproductive success. Additionally, the change in response to elevated [CO₂] over time varied among pollen donor families; thus, changes in [CO₂] could act as a selective force on this species.     

Leaf Photosynthesis and Respiration of High CO(2)-Grown Tobacco Plants Selected for Survival under CO(2) Compensation Point Conditions

Four self-pollinated, doubled-haploid tobacco, (Nicotiana tabacum L.) lines (SP422, SP432, SP435, and SP451), selected as haploids by survival in a low CO₂ atmosphere, and the parental cv Wisconsin-38 were grown from seed in a growth room kept at high CO₂ levels (600-700 parts per million). The selected plants were much larger (especially SP422, SP432, and SP451) than Wisconsin-38 nine weeks after planting. The specific leaf dry weight and the carbon (but not nitrogen and sulfur) content per unit area were also higher in the selected plants. However, the chlorophyll, carotenoid, and alkaloid contents and the chlorophyll a/b ratio varied little. The net CO₂ assimilation rate per unit area measured in the growth room at high CO₂ was not higher in the selected plants. The CO₂ assimilation rate versus intercellular CO₂ curve and the CO₂ compensation point showed no substantial differences among the different lines, even though these plants were selected for survival under CO₂ compensation point conditions. Adult leaf respiration rates were similar when expressed per unit area but were lower in the selected lines when expressed per unit dry weight. Leaf respiration rates were negatively correlated with specific leaf dry weight and with the carbon content per unit area and were positively correlated with nitrogen and sulfur content of the dry matter. The alternative pathway was not involved in respiration in the dark in these leaves. The better carbon economy of tobacco lines selected for low CO₂ survival was not apparently related to an improvement of photosynthesis rate but could be related, at least partially, to a significantly reduced respiration (mainly cytochrome pathway) rate per unit carbon.     

Why are East Asian ecosystems important for carbon cycle research?

The global carbon cycle is one of the most important bio-geochemical cycles. Through photosynthesis, green plants absorb CO₂ from the atmosphere to produce organic matters, such as sugars, and covert solar energy into chemical energy. The organic matters are then used by all other life forms including humans. When ecosystems and atmosphere are in dynamic equilibrium, the flow of CO₂ from the atmosphere into the biosphere because of photosynthesis should be equivalent to the flow of CO₂ released back into the atmosphere by respiration. However, during the past century atmospheric CO₂ concentration has increased substantially because of the burning of fossil fuels. It is highly likely that the atmospheric increase has resulted in global warming and sea level rise, as suggested by the Intergovernmental Panel on Climate Change (IPCC) .     

Combinatorial optimization of cyanobacterial 2,3-butanediol production

A vital goal of renewable technology is the capture and re-energizing of exhausted CO₂ into usable carbon products. Cyanobacteria fix CO₂ more efficiently than plants, and can be engineered to produce carbon feedstocks useful for making plastics, solvents, and medicines. However, fitness of this technology in the economy is threatened by low yields in engineered strains. Robust engineering of photosynthetic microorganisms is lagging behind model microorganisms that rely on energetic carbon, such as Escherichia coli, due in part to slower growth rates and increased metabolic complexity. In this work we show that protein expression from characterized parts is unpredictable in Synechococcus elongatus sp. strain PCC 7942, and may contribute to slow development. To overcome this, we apply a combinatorial approach and show that modulation of the 5'-untranslated region (UTR) can produce a range of protein expression sufficient to optimize chemical feedstock production from CO₂.     

Synthesis, Excretion, and Metabolism of Glycolate under Highly Photorespiratory Conditions in Euglena gracilis Z

Glycolate was excreted from the 5% CO₂-grown cells of Euglena gracilis Z when placed in an atmosphere of 100% O₂ under illumination at 20,000 lux. The amount of excreted glycolate reached 30% of the dry weight of the cells during incubation for 12 hours. The content of paramylon, the reserve polysaccharide of E. gracilis, was decreased during the glycolate excretion, and of the depleted paramylon carbon, two-thirds was excreted to the outside of cells and the remaining metabolized to other compounds, both as glycolate. The paramylon carbon entered Calvin cycle probably as triose phosphate or 3-phosphoglycerate, but not as CO₂ after the complete oxidation through the tricarboxylic acid cycle. The glycolate pathway was partially operative and the activity of the pathway was much less than the rate of the synthesis of glycolate in the cells under 100% O₂ and 20,000 lux; this led the cells to excrete glycolate outside the cells. Exogenous glycolate was metabolized only to CO₂ but not to glycine and serine. The physiologic role of the glycolate metabolism and excretion under such conditions is discussed.     

Seasonal CO2 exchange patterns of developing peach (Prunus persica) fruits in response to temperature, light and CO2 concentration

CO₂ exchange rates per unit dry weight, measured in the field on attached fruits of the late-maturing Cal Red peach cultivar, at 1200 μmol photons m^?2S^?1 and in dark, and photosynthetic rates, calculated by the difference between the rates of CO₂ evolution in light and dark, declined over the growing season. Calculated photosynthetic rates per fruit increased over the season with increasing fruit dry matter, but declined in maturing fruits apparently coinciding with the loss of chlorophyll. Slight net fruit photosynthetic rates ranging from 0. 087 ± 0. 06 to 0. 003 ± 0. 05 nmol CO₂ (g dry weight)^?1 S^?1 were measured in midseason under optimal temperature (15 and 20°C) and light (1200 μmol photons m^?2 S^?1) conditions. Calculated fruit photosynthetic rates per unit dry weight increased with increasing temperatures and photon flux densities during fruit development. Dark respiration rates per unit dry weight doubled within a temperature interval of 10°C; the mean seasonal O₁₀ value was 2. 03 between 20 and 30°C. The highest photosynthetic rates were measured at 35°C throughout the growing season. Since dark respiration rates increased at high temperatures to a greater extent than CO₂ exchange rates in light, fruit photosynthesis was apparently stimulated by high internal CO₂ concentrations via CO₂ refixation. At 15°C, fruit photosynthetic rates tended to be saturated at about 600 μmol photons m^?2 S^?1. Young peach fruits responded to increasing ambient CO₂ concentrations with decreasing net CO₂ exchange rates in light, but more mature fruits did not respond to increases in ambient CO₂. Fruit CO₂ exchange rates in the dark remained fairly constant, apparently uninfluenced by ambient CO₂ concentrations during the entire growing season. Calculated fruit photosynthetic rates clearly revealed the difference in CO₂ response of young and mature peach fruits. Photosynthetic rates of younger peach fruits apparently approached saturation at 370 μl CO₂1^?2. In CO₂ free air, fruit photosynthesis was dependent on CO₂ refixation since CO₂ uptake by the fruits from the external atmosphere was not possible. The difference in photosynthetic rates between fruits in CO₂-free air and 370 μl CO₂ 1^?1 indicated that young peach fruits were apparently able to take up CO₂ from the external atmosphere. CO₂ uptake by peach fruits contributed between 28 and 16% to the fruit photosynthetic rate early in the season, whereas photosynthesis in maturing fruits was supplied entirely by CO₂ refixation.     

Diel, episodic and seasonal changes in pH and concentrations of inorganic carbon in a productive lake

• 1 Two pH electrodes and a thermistor were used to record conditions in the surface of Esthwaite Water every 15 min over a 12-month period. Combined with approximately weekly measurements of alkalinity they allowed inorganic carbon speciation to be calculated.
• 2 Large changes in pH from 7.1 to nearly 10.3, and hence in concentrations of inorganic carbon species, were measured over a year. Carbon speciation and pH varied on a diel, episodic and seasonal basis. Diel variation of up to pH 1.8 was recorded, although typical daily variation was between 0.03 and 1.06 (5 and 95 percentiles). Daily change in concentration of inorganic carbon varied between 4 and 63 mmol m^-3 (5 and 95 percentiles).
• 3 During lake stratification, episodes of high pH, typically of 1–2 weeks' duration were interspersed with episodes of lower pH. These changes appeared to relate to the weather: e.g. low wind velocity, high pressure, low rainfall and high sunshine hours correlated with periods of high pH.
• 4 Seasonal progression of carbon depletion generally followed stratification and the development of high phytoplankton biomass. When the lake was isothermal, the phytoplankton biomass caused relatively small amounts of carbon depletion.
• 5 During autumn, winter and spring, the lake had concentrations of CO₂* (free CO₂) up to 0.12 mol m^-3 which is nearly seven times the calculated atmospheric equilibrium concentration so the lake will accordingly be losing carbon to the atmosphere. In contrast, during periods of elevated pH the concentration of CO₂* was reduced close to zero and the lake will take up atmospheric CO₂. The rates of transfer between water and the atmosphere were estimated using a chemical equilibrium model with three boundary layer thicknesses. The calculations show that over a year the lake loses CO₂ to the atmosphere with the current mean atmospheric level of 360 μmol mol^-1, at between 0.28 and 2.80 mol m^-2 yr^-1. During elevated pH, rates of CO₂-influx increased up to nearly tenfold as a result of chemical-enhancement by parallel flux of HCO^-₃. Input of CO₂* to the lake from the catchment is suggested to be the main source of the carbon lost to the atmosphere.
• 6 The turnover time for CO₂ between the air and water was calculated to be 1 year for the gross influx and 3.3 years for the net flux. These values are less than the average water residence time of 0.25 years, which indicates that over a year inflow from streams is a more important source of inorganic carbon than the atmosphere.
• 7 Influx of CO₂ from the atmosphere was calculated to be roughly equivalent to between 1 and 4% of the rates of production in the water during mid-summer indicating that this source of inorganic carbon is not a major one in this lake.
• 8 Influx of CO₂ from the hypolimnion was estimated on one occasion to be 6.9 10^-9 mol m^-2 s^-1 using transfer values based on mass eddy-diffusion. These rates are equivalent to 23% of the rate of influx of CO₂ from the atmosphere on this occasion which suggests that the hypolimnion is probably a small source of inorganic carbon to the epilimnion. The exception appears to be during windy episodes when pH is depressed. Calculations based on depth-profiles of CO₂* and HCO^-₃ suggest that the measured changes in pH can be accounted for by entrainment of hypolimnetic water into the epilimnion.
• 9 The solubility product for calcite was exceeded by up to about sixfold which, although insufficient to allow homogeneous precipitation, may have allowed heterogeneous precipitation around algal particles.