CO2 and intracellular pH

Abstract The experimental determination of cytoplasmic and vacuolar pH values is discussed. Despite variation in these values evidence indicates that intracellular pH values are normally regulated within narrow limits. The regulatory mechanisms proposed involve the metabolic consumption of OH^& and the active efflux of H ⁺. The evidence for intracellular pH modification in response to CO₂ hydration and the production of HCO^?₃ and H⁺ is examined. Theoretical calculations and experimental data indicate that CO₂ concentrations as high as 5% will lower intracellular pH. Conversely, variation in CO₂ levels around atmospheric concentrations is unlikely to perturb intracellular pH. High CO₂ levels are found in bulky tissues, and flooded root systems. Evidence is presented that the slow diffusion of dissolved CO₂ compared to gaseous CO₂ results in its accumulation. It is proposed that the accumulation of respiratory CO₂ may reduce intracellular pH values when plant tissues, cells or protoplasts are maintained in a liquid culture medium. Finally, the possible role of dark CO₂ fixation and organic acid synthesis in the regulation of intracellular pH is examined.     

Oscillatory Transpiration and CO2 Exchange of Citrus Leaves at the CO2 Compensation Concentration

CO₂ and H₂O vapor exchange were measured by enclosing citrus (Citrus sinensis cv. Sour Orange) leaves in a temperature controlled transparent leaf chamber. Introduction of dry air into the closed circuit gas flow caused cyclic oscillation in CO₂ and H₂O vapor exchange. It is suggested that oscillation in the CO₂ exchange at the CO₂ compensation concentration is due to oscillation in non-stomatal resistance to CO₂. Three types of oscillation were observed: 3–6 min (peak to peak) in young leaves, 30 min in mature leaves, and 160 min in old leaves.     

Seed production and seed quality in a calcareous grassland in elevated CO2

In diverse plant communities the relative contribution of species to community biomass may change considerably in response to elevated CO₂. Along with species‐specific biomass responses, reproduction is likely to change as well with increasing CO₂ and might further accelerate shifts in species composition. Here, we ask if, after 5 years of CO₂ exposure, seed production and seed quality in natural nutrient‐poor calcareous grassland are affected by elevated CO₂ (650 μ L L^?1 vs 360 μ L L^?1) and how this might affect long‐term community dynamics. The effect of elevated CO₂ on the number of flowering shoots (+ 24%, P < 0.01) and seeds (+ 29%, P = 0.06) at the community level was similar to above ground biomass responses in this year, suggesting that the overall allocation to sexual reproduction remained unchanged. Compared among functional groups of species we found a 42% increase in seed number (P < 0.01) of graminoids, a 33% increase (P = 0.07) in forbs, and no significant change in legumes (? 38%, n.s.) under elevated CO₂. Large responses particularly of two graminoid species and smaller responses of many forb species summed up to the significant or marginally significant increase in seed number of graminoids and forbs, respectively. In several species the increase in seed number resulted both from an increase in flowering shoots and an increase in inflorescence size. In most species, seeds tended to be heavier (+ 12%, P < 0.01), and N‐concentration of seeds was significantly reduced in eight out of 13 species. The fraction of germinating seeds did not differ between seeds produced in ambient and elevated CO₂, but time to germination was significantly shortened in two species and prolonged in one species when seeds had been produced in elevated CO₂. Results suggest that species specific increases in seed number and changes in seed quality will exert substantial cumulative effects on community composition in the long run.     

Oceanic sinks for atmospheric CO2

There is approximately 50 times more inorganic carbon in the global ocean than in the atmosphere. On time scales of decades to millions of years, the interaction between these two geophysical fluids determines atmospheric CO₂ levels. During glacial periods, for example, the ocean serves as the major sink for atmospheric CO₂, while during glacial–interglacial transitions, it is a source of CO₂ to the atmosphere. The mechanisms responsible for determining the sign of the net exchange of CO₂ between the ocean and the atmosphere remain unresolved. There is evidence that during glacial periods, phytoplankton primary productivity increased, leading to an enhanced sedimentation of particulate organic carbon into the ocean interior. The stimulation of primary production in glacial episodes can be correlated with increased inputs of nutrients limiting productivity, especially aeolian iron. Iron directly enhances primary production in high nutrient (nitrate and phosphate) regions of the ocean, of which the Southern Ocean is the most important. This trace element can also enhance nitrogen fixation, and thereby indirectly stimulate primary production throughout the low nutrient regions of the central ocean basins. While the export flux of organic carbon to the ocean interior was enhanced during glacial periods, this process does not fully account for the sequestration of atmospheric CO₂. Heterotrophic oxidation of the newly formed organic carbon, forming weak acids, would have hydrolyzed CaCO₃ in the sediments, increasing thereby oceanic alkalinity which, in turn, would have promoted the drawdown of atmospheric CO₂. This latter mechanism is consistent with the stable carbon isotope pattern derived from air trapped in ice cores. The oceans have also played a major role as a sink for up to 30% of the anthropogenic CO₂ produced during the industrial revolution. In large part this is due to CO₂ solution in the surface ocean; however, some, poorly quantified fraction is a result of increased new production due to anthropogenic inputs of combined N, P and Fe. Based on ‘circulation as usual’, models predict that future anthropogenic CO₂ inputs to the atmosphere will, in part, continue to be sequestered in the ocean. Human intervention (large-scale Fe fertilization; direct CO₂ burial in the deep ocean) could increase carbon sequestration in the oceans, but could also result in unpredicted environmental perturbations. Changes in the oceanic thermohaline circulation as a result of global climate change would greatly alter the predictions of C sequestration that are possible on a ‘circulation as usual’ basis.     

The respiratory source of CO2

Abstract Approximately half of the carbon plants fix in photosynthesis is lost in dark respiration. The major pathways for dark respiration and their control are briefly discussed in the context of a growing plant. It is suggested that whole-plant respiration may be largely ADP-limited and that fine control of the respiratory network serves to select the respiratory substrate and to partition carbon between the numerous possible fates within the network. The striking stoichiometry between whole-plant growth and respiration is reviewed, and the relationships between substrate-limited growth and ADP-limited respiration are discussed.     

Germination of cocklebur seed and growth of their axial and cotyledonary tissues in response to C2H4, CO2 and/or O2 under water stress

Abstract. Germination modes of lower seeds of cocklebur (Xanthium pennsylvanicum Wallr.) under different water stresses, prepared with mannitol solution, were examined in relation to gaseous factors. As the concentration of mannitol increased, germination was increasingly inhibited at a mode which was drawn by two straight lines having different slopes and meeting at an angle. One is a sharp line occurring at the lower concentrations of mannitol; the other is a gentle line occurring at higher concentrations of mannitol. The former reflected the growth response of axial tissues to mild water stress, whereas the latter reflected the growth response of cotyledonary tissues to severe water stress. The germination potential of cocklebur seeds increased with increasing temperature. Thus, the seeds were more resistant to water stress at higher than al lower temperatures. This increased germination potential under water stress resulted from the greater growth potential of axial tissues, but not cotyledonary tissues, at higher temperature. Increased O₂ levels improved both the reduced axial and cotyledonary growth under water stress. Carbon dioxide predominantly enhanced axial growth under water stress, whereas C₂H₄ exclusively enhanced cotyledonary growth. Thus, these gases were effective in potentiating germination under water stress. When combined with each other, these gases caused more pronounced growth of the axial and cotyledonary tissues, leading to germination under more severe water stresses. Maximal axial and cotyledonary growth under water stress occurred in the simultaneous presence of CO₂, C₂H₄ and O₂, which allowed the germination at higher mannitol concentrations above 0.6 kmol m^?3 From these results, it was suggested that cocklebur seeds would override water stress by depending upon both the Corresponding axial growth and the C₂H₄-responding cotyledonary growth.     

Soil CO2 efflux in a boreal pine forest under atmospheric CO2 enrichment and air warming

The response of forest soil CO₂ efflux to the elevation of two climatic factors, the atmospheric concentration of CO₂ (↑CO₂ of 700 μmol mol⁻¹) and air temperature (↑ T with average annual increase of 5°C), and their combination (↑CO₂+↑ T ) was investigated in a 4-year, full-factorial field experiment consisting of closed chambers built around 20-year-old Scots pines ( Pinus sylvestris L.) in the boreal zone of Finland. Mean soil CO₂ efflux in May–October increased with elevated CO₂ by 23–37%, with elevated temperature by 27–43%, and with the combined treatment by 35–59%. Temperature elevation was a significant factor in the combined 4-year efflux data, whereas the effect of elevated CO₂ was not as evident. Elevated temperature had the most pronounced impact early and late in the season, while the influence of elevated CO₂ alone was especially notable late in the season. Needle area was found to be a significant predictor of soil CO₂ efflux, particularly in August, a month of high root growth, thus supporting the assumption of a close link between whole-tree physiology and soil CO₂ emissions. The decrease in the temperature sensitivity of soil CO₂ efflux observed in the elevated temperature treatments in the second year nevertheless suggests the existence of soil response mechanisms that may be independent of the assimilating component of the forest ecosystem. In conclusion, elevated atmospheric CO₂ and air temperature consistently increased forest soil CO₂ efflux over the 4-year period, their combined effect being additive, with no apparent interaction.     

Effects of elevated atmospheric CO2 on net ecosystem CO2 exchange of a scrub–oak ecosystem

We report the results of a 2‐year study of effects of the elevated (current ambient plus 350 μmol CO₂ mol^?1) atmospheric CO₂ concentration (C_a) on net ecosystem CO₂ exchange (NEE) of a scrub–oak ecosystem. The measurements were made in open‐top chambers (OTCs) modified to function as open gas‐exchange systems. The OTCs enclosed samples of the ecosystem (ca. 10 m² surface area) that had regenerated after a fire, 5 years before, in either current ambient or elevated C_a. Throughout the study, elevated C_a increased maximum NEE (NEE_max) and the apparent quantum yield of the NEE (φ_NEE) during the photoperiod. The magnitude of the stimulation of NEE_max, expressed per unit ground area, was seasonal, rising from 50% in the winter to 180% in the summer. The key to this stimulation was effects of elevated C_a, and their interaction with the seasonal changes in the environment, on ecosystem leaf area index, photosynthesis and respiration. The separation of these factors was difficult. When expressed per unit leaf area the stimulation of the NEE_max ranged from 7% to 60%, with the increase being dependent on increasing soil water content (W_soil). At night, the CO₂ effluxes from the ecosystem (NEE_night) were on an average 39% higher in elevated C_a. However, the increase varied between 6% and 64%, and had no clear seasonality. The partitioning of NEE_night into its belowground (R_below) and aboveground (R_above) components was carried out in the winter only. A 35% and 27% stimulation of NEE_night in December 1999 and 2000, respectively, was largely due to a 26% and 28% stimulation of R_below in the respective periods, because R_below constituted ca. 87% of NEE_night. The 37% and 42% stimulation of R_above in December 1999 and 2000, respectively, was less than the 65% and 80% stimulation of the aboveground biomass by elevated C_a at these times. An increase in the relative amount of the aboveground biomass in woody tissue, combined with a decrease in the specific rate of stem respiration of the dominant species Quercus myrtifolia in elevated C_a, was responsible for this effect. Throughout this study, elevated C_a had a greater effect on carbon uptake than on carbon loss, in terms of both the absolute flux and relative stimulation. Consequently, for this scrub–oak ecosystem carbon sequestration was greater in the elevated C_a during this 2‐year study period.