Response of wheat canopy CO2 and water gas-exchange to soil water content under ambient and elevated CO2

The nature of the interaction between drought and elevated CO₂ partial pressure (pC_a) is critically important for the effects of global change on crops. Some crop models assume that the relative responses of transpiration and photosynthesis to soil water deficit are unaltered by elevated pC_a, while others predict decreased sensitivity to drought at elevated pC_a. These assumptions were tested by measuring canopy photosynthesis and transpiration in spring wheat (cv. Minaret) stands grown in boxes with 100 L rooting volume. Plants were grown under controlled environments with constant light (300 µmol m^?2 s^?1) at ambient (36 Pa) or elevated (68 Pa) pC_a and were well watered throughout growth or had a controlled decline in soil water starting at ear emergence. Drought decreased final aboveground biomass (?15%) and grain yield (?19%) while elevated pC_a increased biomass (+24%) and grain yield (+29%) and there was no significant interaction. Elevated pC_a increased canopy photosynthesis by 15% on average for both water regimes and increased dark respiration per unit ground area in well‐watered plants, but not drought‐grown ones. Canopy transpiration and photosynthesis were decreased in drought‐grown plants relative to well‐watered plants after about 20–25 days from the start of the drought. Elevated pC_a decreased transpiration only slightly during drought, but canopy photosynthesis continued to be stimulated so that net growth per unit water transpired increased by 21%. The effect of drought on canopy photosynthesis was not the consequence of a loss of photosynthetic capacity initially, as photosynthesis continued to be stimulated proportionately by a fixed increase in irradiance. Drought began to decrease canopy transpiration below a relative plant‐available soil water content of 0.6 and canopy photosynthesis and growth below 0.4. The shape of these responses were unaffected by pC_a, supporting the simple assumption used in some models that they are independent of pC_a.     

Partitioning net ecosystem carbon exchange with isotopic fluxes of CO2

Because biological and physical processes alter the stable isotopic composition of atmospheric CO₂, variations in isotopic content can be used to investigate those processes. Isotopic flux measurements of ¹³CO₂ above terrestrial ecosystems can potentially be used to separate net ecosystem CO₂ exchange (NEE) into its component fluxes, net photosynthetic assimilation (F_A) and ecosystem respiration (F_R). In this paper theory is developed to partition measured NEE into F_A and F_R, using measurements of fluxes of CO₂ and ¹³CO₂, and isotopic composition of respired CO₂ and forest air. The theory is then applied to fluxes measured (or estimated, for ¹³CO₂) in a temperate deciduous forest in eastern Tennessee (Walker Branch Watershed). It appears that there is indeed enough additional information in ¹³CO₂ fluxes to partition NEE into its photosynthetic and respiratory components. Diurnal patterns in F_A and F_R were obtained, which are consistent in magnitude and shape with patterns obtained from NEE measurements and an exponential regression between night‐time NEE and temperature (a standard technique which provides alternate estimates of F_R and F_A). The light response curve for photosynthesis (F_A vs. PAR) was weakly nonlinear, indicating potential for saturation at high light intensities. Assimilation‐weighted discrimination against ¹³CO₂ for this forest during July 1999 was 16.8–17.1‰, depending on canopy conductance. The greatest uncertainties in this approach lie in the evaluation of canopy conductance and its effect on whole‐canopy photosynthetic discrimination, and thus the indirect methods used to estimate isotopic fluxes. Direct eddy covariance measurements of ¹³CO₂ flux are needed to assess the validity of the assumptions used and provide defensible isotope‐based estimates of the component fluxes of net ecosystem exchange.     

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.     

Wood CO2 efflux in a primary tropical rain forest

The balance between photosynthesis and plant respiration in tropical forests may substantially affect the global carbon cycle. Woody tissue CO₂ efflux is a major component of total plant respiration, but estimates of ecosystem‐scale rates are uncertain because of poor sampling in the upper canopy and across landscapes. To overcome these problems, we used a portable scaffolding tower to measure woody tissue CO₂ efflux from ground level to the canopy top across a range of sites of varying slope and soil phosphorus content in a primary tropical rain forest in Costa Rica. The objectives of this study were to: (1) determine whether to use surface area, volume, or biomass for modeling and extrapolating wood CO₂ efflux, (2) determine if wood CO₂ efflux varied seasonally, (3) identify if wood CO₂ efflux varied by functional group, height in canopy, soil fertility, or slope, and (4) extrapolate wood CO₂ efflux to the forest. CO₂ efflux from small diameter woody tissue (<10 cm) was related to surface area, while CO₂ efflux from stems >10 cm was related to both surface area and volume. Wood CO₂ efflux showed no evidence of seasonality over 2 years. CO₂ efflux per unit wood surface area at 25° (F_A) was highest for the N‐fixing dominant tree species Pentaclethra macroloba, followed by other tree species, lianas, then palms. Small diameter F_A increased steeply with increasing height, and large diameter F_A increased with diameter. Soil phosphorus and slope had slight, but complex effects on F_A. Wood CO₂ efflux per unit ground area was 1.34±0.36 μmol m^?2 s^?1, or 508±135 g C m^?2 yr^?1. Small diameter wood, only 15% of total woody biomass, accounted for 70% of total woody tissue CO₂ efflux from the forest; while lianas, only 3% of total woody biomass, contributed one‐fourth of the total wood CO₂ efflux.     

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.     

Growth and phenology of mature temperate forest trees in elevated CO2

Are mature forest trees carbon limited at current CO₂ concentrations? Will ‘mid‐life’, 35 m tall deciduous trees grow faster in a CO₂‐enriched atmosphere? To answer these questions we exposed ca. 100‐year‐old temperate forest trees at the Swiss Canopy Crane site near Basel, Switzerland to a ca. 540 ppm CO₂ atmosphere using web‐FACE technology. Here, we report growth responses to elevated CO₂ for 11 tall trees (compared with 32 controls) of five species during the initial four treatment years. Tested across all trees, there was no CO₂ effect on stem basal area (BA) increment (neither when tested per year nor cumulatively for 4 years). In fact, the 4th year means were almost identical for the two groups. Stem growth data were standardized by pretreatment growth (5 years) in order to account for a priori individual differences in vigor. Although this experiment was not designed to test species specific effects, one species, the common European beech, Fagus sylvatica, showed a significant growth enhancement in the first year, which reoccurred during a centennial drought in the third year. None of the other dominant species (Quercus petraea, Carpinus betulus) showed a growth response to CO₂ in any of the 4 years or for all years together. The inclusion or exclusion of single individuals of Prunus avium and Tilia platyphyllos did not change the picture. In elevated CO₂, lateral branching in terminal shoots was higher in Fagus in 2002, when shoots developed from buds that were formed during the first season of CO₂ enrichment (2001), but there was no effect in later years and no change in lateral branching in any of the other species. In Quercus, there was a steady stimulation of leading shoot length in high‐CO₂ trees. Phenological variables (bud break, leaf fall, leaf duration) were highly species specific and were not affected by elevated CO₂ in any consistent way. Our 4‐year data set reflects a very dynamic and species‐specific response of tree growth to a step change in CO₂ supply. Stem growth after 4 years of exposure does not support the notion that mature forest trees will accrete wood biomass at faster rates in a future CO₂‐enriched atmosphere.     

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.     

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.