Fossil plant evidence for Early and Middle Jurassic paleoenvironmental changes in Lanzhou area, Northwest China

Estimating the paleoclimate changes through CO₂ levels has become a promising area of geological research. This paper focuses on analysis of fossil Ginkgo in continuous sedimentary series in northwestern China using plant anatomy and organic geochemistry approaches. The CO₂ variation curve during Early and Middle Jurassic is reconstructed based on the stomatal ratio method, which is consistent with the estimated results of GEOCARB III. In comparison with the carbon isotopic composition measured from the fossil leaves of Ginkgo, we suggest that the stomata-based CO₂ concentrations for the Jurassic are generally consistent with the predictions of the geochemical model, ranging from 1000 to 1600 ppmv. Measurements of carbon isotope values demonstrate that the water use efficiency of the fossil Ginkgo in a “green house” world is higher than that of the living Ginkgo. Investigations of physiological responses of plants to the increasing CO₂ level at the present and in the future should help demonstrate this effect. The cause of the carbon isotopic shift at the boundary between Aalenian and Bajocian for the Yaojie Basin is unclear and therefore needs further investigation.     

Inorganic Carbon Uptake during Photosynthesis : II. Uptake by Isolated Asparagus Mesophyll Cells during Isotopic Disequilibrium

The species of inorganic carbon (CO₂ or HCO₃⁻) taken up a source of substrate for photosynthetic fixation by isolated Asparagus sprengeri mesophyll cells is investigated. Discrimination between CO₂ or HCO₃⁻ transport, during steady state photosynthesis, is achieved by monitoring the changes (by ¹⁴C fixation) which occur in the specific activity of the intracellular pool of inorganic carbon when the inorganic carbon present in the suspending medium is in a state of isotopic disequilibrium. Quantitative comparisons between theoretical (CO₂ or HCO₃⁻ transport) and experimental time-courses of ¹⁴C incorporation, over the pH range of 5.2 to 7.5, indicate that the specific activity of extracellular CO₂, rather than HCO₃⁻, is the appropriate predictor of the intracellular specific activity. It is concluded, therefore, that CO₂ is the major source of exogenous inorganic carbon taken up by Asparagus cells. However, at high pH (8.5), a component of net DIC uptake may be attributable to HCO₃⁻ transport, as the incorporation of ¹⁴C during isotopic disequilibrium exceeds the maximum possible incorporation predicted on the basis of CO₂ uptake alone. The contribution of HCO₃⁻ to net inorganic carbon uptake (pH 8.5) is variable, ranging from 5 to 16%, but is independent of the extracellular HCO₃⁻ concentration. The evidence for direct HCO₃⁻ transport is subject to alternative explanations and must, therefore, be regarded as equivocal. Nonlinear regression analysis of the rate of ¹⁴C incorporation as a function of time indicates the presence of a small extracellular resistance to the diffusion of CO₂, which is partially alleviated by a high extracellular concentration of HCO₃⁻.     

Effect of Varying CO(2) Partial Pressure on Photosynthesis and on Carbon Isotope Composition of Carbon-4 of Malate from the Crassulacean Acid Metabolism Plant Kalanchoë daigremontiana Hamet et Perr

Intact leaves of Kalanchoë daigremontiana were exposed to CO₂ partial pressures of 100, 300, and 1000 microbars. Malic acid was extracted, purified, and degraded in order to obtain isotopic composition of carbon-1 and carbon-4. From these data, it is possible to calculate the carbon isotope composition of newly fixed carbon in malate. In all three treatments, the isotopic composition of newly introduced carbon is the same as that of the CO₂ source and is independent of CO₂ partial pressures over the range tested. Comparison with numerical models described previously (O'Leary 1981 Phytochemistry 20: 553-567) indicates that we would expect carbon 4 of malate to be 4‰ more negative than source CO₂ if diffusion is totally limiting or 7‰ more positive than source CO₂ if carboxylation is totally limiting. Our results demonstrate that stomatal aperture adjusts to changing CO₂ partial pressures and maintains the ratio of diffusion resistance to carboxylation resistance approximately constant. In this study, carboxylation and diffusion resistances balance so that essentially no fractionation occurs during malate synthesis. Gas exchange studies of the same leaves from which malate was extracted show that the extent of malate synthesis over the whole night is nearly independent of CO₂ partial pressure, although there are small variations in CO₂ uptake rate. Both the gas exchange and the isotope studies indicate that the ratio of external to internal CO₂ partial pressure is the same in all three treatments. Inasmuch as a constant ratio will result in constant isotope fractionation, this observation may explain why plants in general have fairly invariable ¹³C contents, despite growing under a variety of environmental conditions.     

The Role of External Carbonic Anhydrase in Inorganic Carbon Acquisition by Chlamydomonas reinhardii at Alkaline pH

The role of external carbonic anhydrase in inorganic carbon acquisition and photosynthesis by Chlamydomonas reinhardii at alkaline pH (8.0) was studied. Acetazolamide (50 micromolar) completely inhibited external carbonic anhydrase (CA) activity as determined from isotopic disequilibrium experiments. Under these conditions, photosynthetic rates at low dissolved inorganic carbon (DIC) were far greater than could be maintained by CO₂ supplied from the spontaneous dehydration of HCO₃⁻ thereby showing that C. reinhardii has the ability to utilize exogenous HCO₃⁻. Acetazolamide increased the concentration of DIC required to half-saturate photosynthesis from 38 to 80 micromolar, while it did not affect the maximum photosynthetic rate. External CA activity was also removed from the cell-wall-less mutant (CW-15) by washing. This had no effect on the photosynthetic kinetics of the algae while the addition of acetazolamide to washed cells (CW-15) increased the K_½^DIC from 38 to 80 micromolar. Acetazolamide also caused a buildup of the inorganic carbon pool upon NaHCO₃ addition, indicating that this compound partially inhibited internal CA activity. The effects of acetazolamide on the photosynthetic kinetics of C. reinhardii are likely due to the inhibition of internal rather than a consequence of the inhibition of external CA. Further analysis of the isotopic disequilibrium experiments at saturating concentration of DIC provided evidence consistent with active CO₂ transport by C. reinhardii. The observation that C. reinhardii has the ability to take up both CO₂ and bicarbonate throws into question the role of external CA in the accumulation of DIC in this alga.     

Increased Atmospheric SO2 Detected from Changes in Leaf Physiognomy across the Triassic–Jurassic Boundary Interval of East Greenland

The Triassic–Jurassic boundary (Tr–J; ∼201 Ma) is marked by a doubling in the concentration of atmospheric CO₂, rising temperatures, and ecosystem instability. This appears to have been driven by a major perturbation in the global carbon cycle due to massive volcanism in the Central Atlantic Magmatic Province. It is hypothesized that this volcanism also likely delivered sulphur dioxide (SO₂) to the atmosphere. The role that SO₂ may have played in leading to ecosystem instability at the time has not received much attention. To date, little direct evidence has been presented from the fossil record capable of implicating SO₂ as a cause of plant extinctions at this time. In order to address this, we performed a physiognomic leaf analysis on well-preserved fossil leaves, including Ginkgoales, bennettites, and conifers from nine plant beds that span the Tr–J boundary at Astartekløft, East Greenland. The physiognomic responses of fossil taxa were compared to the leaf size and shape variations observed in nearest living equivalent taxa exposed to simulated palaeoatmospheric treatments in controlled environment chambers. The modern taxa showed a statistically significant increase in leaf roundness when fumigated with SO₂. A similar increase in leaf roundness was also observed in the Tr–J fossil taxa immediately prior to a sudden decrease in their relative abundances at Astartekløft. This research reveals that increases in atmospheric SO₂ can likely be traced in the fossil record by analyzing physiognomic changes in fossil leaves. A pattern of relative abundance decline following increased leaf roundness for all six fossil taxa investigated supports the hypothesis that SO₂ had a significant role in Tr–J plant extinctions. This finding highlights that the role of SO₂ in plant biodiversity declines across other major geological boundaries coinciding with global scale volcanism should be further explored using leaf physiognomy.     

Rates of Microbial Metabolism in Deep Coastal Plain Aquifers

Rates of microbial metabolism in deep anaerobic aquifers of the Atlantic coastal plain of South Carolina were investigated by both microbiological and geochemical techniques. Rates of [2-¹⁴C]acetate and [U-¹⁴C]glucose oxidation as well as geochemical evidence indicated that metabolic rates were faster in the sandy sediments composing the aquifers than in the clayey sediments of the confining layers. In the sandy aquifer sediments, estimates of the rates of CO₂ production (millimoles of CO₂ per liter per year) based on the oxidation of [2-¹⁴C] acetate were 9.4 × 10⁻³ to 2.4 × 10⁻¹ for the Black Creek aquifer, 1.1 × 10⁻² for the Middendorf aquifer, and <7 × 10⁻⁵ for the Cape Fear aquifer. These estimates were at least 2 orders of magnitude lower than previously published estimates that were based on the accumulation of CO₂ in laboratory incubations of similar deep subsurface sediments. In contrast, geochemical modeling of groundwater chemistry changes along aquifer flowpaths gave rate estimates that ranged from 10⁻⁴ to 10⁻⁶ mmol of CO₂ per liter per year. The age of these sediments (ca. 80 million years) and their organic carbon content suggest that average rates of CO₂ production could have been no more than 10⁻⁴ mmol per liter per year. Thus, laboratory incubations may greatly overestimate the in situ rates of microbial metabolism in deep subsurface environments. This has important implications for the use of laboratory incubations in attempts to estimate biorestoration capacities of deep aquifers. The rate estimates from geochemical modeling indicate that deep aquifers are among the most oligotrophic aquatic environments in which there is ongoing microbial metabolism.     

Net Carbon Emissions from Deforestation in Bolivia during 1990-2000 and 2000-2010: Results from a Carbon Bookkeeping Model

Accurate estimates of global carbon emissions are critical for understanding global warming. This paper estimates net carbon emissions from land use change in Bolivia during the periods 1990–2000 and 2000–2010 using a model that takes into account deforestation, forest degradation, forest regrowth, gradual carbon decomposition and accumulation, as well as heterogeneity in both above ground and below ground carbon contents at the 10 by 10 km grid level. The approach permits detailed maps of net emissions by region and type of land cover. We estimate that net CO₂ emissions from land use change in Bolivia increased from about 65 million tons per year during 1990–2000 to about 93 million tons per year during 2000–2010, while CO₂ emissions per capita and per unit of GDP have remained fairly stable over the sample period. If we allow for estimated biomass increases in mature forests, net CO₂ emissions drop to close to zero. Finally, we find these results are robust to alternative methods of calculating emissions.     

Long-term Effects of Free Air CO2 Enrichment (FACE) on Soil Respiration

Emissions of CO₂ from soils make up one of the largest fluxes in the global C cycle, thus small changes in soil respiration may have large impacts on global C cycling. Anthropogenic additions of CO₂ to the atmosphere are expected to alter soil carbon cycling, an important component of the global carbon budget. As part of the Duke Forest Free-Air CO₂ Enrichment (FACE) experiment, we examined how forest growth at elevated (+200 ppmv) atmospheric CO₂ concentration affects soil CO₂ dynamics over 7 years of continuous enrichment. Soil respiration, soil CO₂ concentrations, and the isotopic signature of soil CO₂ were measured monthly throughout the 7 years of treatment. Estimated annual rates of soil CO₂ efflux have been significantly higher in the elevated plots in every year of the study, but over the last 5 years the magnitude of the CO₂ enrichment effect on soil CO₂ efflux has declined. Gas well samples indicate that over 7 years fumigation has led to sustained increases in soil CO₂ concentrations and depletion in the δ¹³C of soil CO₂ at all but the shallowest soil depths.     

Long-term effects of elevated atmospheric CO<Subscript>2</Subscript> on species composition and productivity of a southern African C<Subscript>4</Subscript> dominated grassland in the vicinity of a CO<Subscript>2</Subscript> exhalation

We describe the long-term effects of a CO₂ exhalation, created more than 70 years ago, on a natural C₄ dominated sub-tropical grassland in terms of ecosystem structure and functioning. We tested whether long-term CO₂ enrichment changes the competitive balance between plants with C₃ and C₄ photosynthetic pathways and how CO₂ enrichment has affected species composition, plant growth responses, leaf properties and soil nutrient, carbon and water dynamics. Long-term effects of elevated CO₂ on plant community composition and system processes in this sub-tropical grassland indicate very subtle changes in ecosystem functioning and no changes in species composition and dominance which could be ascribed to elevated CO₂ alone. Species compositional data and soil δ¹³C isotopic evidence suggest no detectable effect of CO₂ enrichment on C₃:C₄ plant mixtures and individual species dominance. Contrary to many general predictions C₃ grasses did not become more abundant and C₃ shrubs and trees did not invade the site. No season length stimulation of plant growth was found even after 5 years of exposure to CO₂ concentrations averaging 610 μmol mol⁻¹. Leaf properties such as total N decreased in the C₃ but not C₄ grass under elevated CO₂ while total non-structural carbohydrate accumulation was not affected. Elevated CO₂ possibly lead to increased end-of-season soil water contents and this result agrees with earlier studies despite the topographic water gradient being a confounding problem at our research site. Long-term CO₂ enrichment also had little effect on soil carbon storage with no detectable changes in soil organic matter found. There were indications that potential soil respiration and N mineralization rates could be higher in soils close to the CO₂ source. The conservative response of this grassland suggests that many of the reported effects of elevated CO₂ on similar ecosystems could be short duration experimental artefacts that disappear under long-term elevated CO₂ conditions.     

Growth and Photosynthesis of the Cyanobacterium Synechococcus leopoliensis in HCO(3)-Limited Chemostats

Synechococcus leopoliensis was grown in HCO₃⁻-limited chemostats. Growth at 50% the maximum rate occurred when the inorganic carbon concentration was 10 to 15 micromolar (or 5.6 to 8.4 nanomolar CO₂). The O₂ to CO₂ ratios during growth were as high as 192,000 to 1. At growth rates below 80% the maximum rate, essentially all the supplied inorganic carbon was converted to organic carbon, and the cells were carbon limited. Carbon-limited cells used HCO₃⁻ rather than CO₂ for growth. They also exhibited a very high photosynthetic affinity for inorganic carbon in short-term experiments. Cells growing at greater than 80% maximum growth rate, in the presence of high dissolved inorganic carbon, were termed carbon sufficient. These cells had photosynthetic affinities that were about 1000-fold lower than HCO₃⁻-limited cells and also had a reduced capacity for HCO₃⁻ transport. HCO₃⁻-limited cells are reminiscent of the air-grown cells of batch culture studies while the carbon sufficient cells are reminiscent of high-CO₂ grown cells. However, the low affinity cells of the present study were growing at CO₂ concentrations less than air saturation. This suggests that supranormal levels of CO₂ not required to induce the physiological changes usually ascribed to high CO₂ cells.     

Mind the gap: non‐biological processes contributing to soil CO2 efflux

Widespread recognition of the importance of soil CO₂ efflux as a major source of CO₂ to the atmosphere has led to active research. A large soil respiration database and recent reviews have compiled data, methods, and current challenges. This study highlights some deficiencies for a proper understanding of soil CO₂ efflux focusing on processes of soil CO₂ production and transport that have not received enough attention in the current soil respiration literature. It has mostly been assumed that soil CO₂ efflux is the result of biological processes (i.e. soil respiration), but recent studies demonstrate that pedochemical and geological processes, such as geothermal and volcanic CO₂ degassing, are potentially important in some areas. Besides the microbial decomposition of litter, solar radiation is responsible for photodegradation or photochemical degradation of litter. Diffusion is considered to be the main mechanism of CO₂ transport in the soil, but changes in atmospheric pressure and thermal convection may also be important mechanisms driving soil CO₂ efflux greater than diffusion under certain conditions. Lateral fluxes of carbon as dissolved organic and inorganic carbon occur and may cause an underestimation of soil CO₂ efflux. Traditionally soil CO₂ efflux has been measured with accumulation chambers assuming that the main transport mechanism is diffusion. New techniques are available such as improved automated chambers, CO₂ concentration profiles and isotopic techniques that may help to elucidate the sources of carbon from soils. We need to develop specific and standardized methods for different CO₂ sources to quantify this flux on a global scale. Biogeochemical models should include biological and non‐biological CO₂ production processes before we can predict the response of soil CO₂ efflux to climate change. Improving our understanding of the processes involved in soil CO₂ efflux should be a research priority given the importance of this flux in the global carbon budget.     

Experimental determination of stable isotope variability in Globigerina bulloides: implications for paleoceanographic reconstructions

We have quantified the environmental and physiological parameters responsible for stable isotopic disequilibrium in the non-symbiotic planktic foraminifera, Globigerina bulloides, via controlled experiments with living specimens. Individual test chambers secreted in the laboratory were amputated, pooled with other chambers from defined positions in the shell whorl and analyzed for their carbon and oxygen isotopic composition. When temperature, δ¹⁸O_water and δ¹³C of ΣCO₂ are kept constant, the chamber δ¹³C and δ¹⁸O values increase 2.6 and 0.8%. respectively between the smallest chambers (chs. 1–9, shell size ≈180 μm) and final chamber (ch. 14, shell size ≈500 μm). Feeding experiments with prey of different δ¹³C values show that 8–15% of the chamber δ¹³C signal is due to the incorporation of metabolic CO₂. The observed ontogenetic trend is responsible for the stable isotope size-dependency in this species and may be due to a fractionation mechanism involving the incorporation of metabolic CO₂ during calcification. Temperature experiments show that shell δ¹⁸O varies as predicted by paleotemperature equations, but is offset from equilibrium. We present correction factors that should be applied to δ¹³C and δ¹⁸O data from well constrained size ranges to yield either oxygen isotope equilibrium or ambient δ¹³C of seawater ΣCO₂. Our results suggest that for paleoceanographic applications, shells in the 270–320 μm size range are optimal for paleoenvironmental reconstructions     

The curious consistency of carbon biosignatures over billions of years of Earth-life coevolution

The oldest and most wide-ranging signal of biological activity (biosignature) on our planet is the carbon isotope composition of organic materials preserved in rocks. These biosignatures preserve the long-term evolution of the microorganism-hosted metabolic machinery responsible for producing deviations in the isotopic compositions of inorganic and organic carbon. Despite billions of years of ecosystem turnover, evolutionary innovation, organismic complexification, and geological events, the organic carbon that is a residuum of the global marine biosphere in the rock record tells an essentially static story. The ~25‰ mean deviation between inorganic and organic ¹³C/¹²C values has remained remarkably unchanged over >3.5 billion years. The bulk of this record is conventionally attributed to early-evolved, RuBisCO-mediated CO₂ fixation that, in extant oxygenic phototrophs, produces comparable isotopic effects and dominates modern primary production. However, billions of years of environmental transition, for example, in the progressive oxygenation of the Earth’s atmosphere, would be expected to have accompanied shifts in the predominant RuBisCO forms as well as enzyme-level adaptive responses in RuBisCO CO₂-specificity. These factors would also be expected to result in preserved isotopic signatures deviating from those produced by extant RuBisCO in oxygenic phototrophs. Why does the bulk carbon isotope record not reflect these expected environmental transitions and evolutionary innovations? Here, we discuss this apparent discrepancy and highlight the need for greater quantitative understanding of carbon isotope fractionation behavior in extant metabolic pathways. We propose novel, laboratory-based approaches to reconstructing ancestral states of carbon metabolisms and associated enzymes that can constrain isotopic biosignature production in ancient biological systems. Together, these strategies are crucial for integrating the complementary toolsets of biological and geological sciences and for interpretation of the oldest record of life on Earth.Subject terms: Bacterial evolution, Applied microbiology, Biogeochemistry     

The contribution of C<Subscript>3</Subscript> and C<Subscript>4</Subscript> plants to the carbon cycle of a tallgrass prairie: an isotopic approach

The photosynthetic pathway composition (C₃:C₄ mixture) of an ecosystem is an important controller of carbon exchanges and surface energy flux partitioning, and therefore represents a fundamental ecophysiological distinction. To assess photosynthetic mixtures at a tallgrass prairie pasture in Oklahoma, we collected nighttime above-canopy air samples along concentration and isotopic gradients throughout the 1999 and 2000 growing seasons. We analyzed these samples for their CO₂ concentration and carbon isotopic composition and calculated C₃:C₄ proportions with a two-source mixing model. In 1999, the C₄ percentage increased from 38% in spring (late April) to 86% in early fall (mid-September). The C₄ percentages inferred from ecosystem respiration measurements in 2000 indicate a smaller shift, from 67% in spring (early May) to 77% in mid-summer (late July). We also sampled daytime CO₂ concentration and carbon isotope gradients above the canopy to determine ecosystem discrimination against ¹³CO₂ during net uptake. These discrimination values were always lower than corresponding nighttime ecosystem respiration isotopic signatures would suggest. After accounting for the isotopic disequilibria between respiration and photosynthesis resulting from seasonal variations in the C₃:C₄ mixture, we estimated canopy photosynthetic discrimination. The C₄ percentage calculated from this approach agrees with the percentage determined from nighttime respiration for sampling periods in both growing seasons. Isotopic imbalances between photosynthesis and respiration are likely to be common in mixed C₃:C₄ ecosystems and must be considered when using daytime isotopic measurements to constrain ecosystem physiology. Given the global extent of such ecosystems, isotopic imbalances likely contribute to global variations in the carbon isotopic composition of atmospheric CO₂.     

Warming Reduces Carbon Losses from Grassland Exposed to Elevated Atmospheric Carbon Dioxide

The flux of carbon dioxide (CO₂) between terrestrial ecosystems and the atmosphere may ameliorate or exacerbate climate change, depending on the relative responses of ecosystem photosynthesis and respiration to warming temperatures, rising atmospheric CO₂, and altered precipitation. The combined effect of these global change factors is especially uncertain because of their potential for interactions and indirectly mediated conditions such as soil moisture. Here, we present observations of CO₂ fluxes from a multi-factor experiment in semi-arid grassland that suggests a potentially strong climate – carbon cycle feedback under combined elevated [CO₂] and warming. Elevated [CO₂] alone, and in combination with warming, enhanced ecosystem respiration to a greater extent than photosynthesis, resulting in net C loss over four years. The effect of warming was to reduce respiration especially during years of below-average precipitation, by partially offsetting the effect of elevated [CO₂] on soil moisture and C cycling. Carbon losses were explained partly by stimulated decomposition of soil organic matter with elevated [CO₂]. The climate – carbon cycle feedback observed in this semiarid grassland was mediated by soil water content, which was reduced by warming and increased by elevated [CO₂]. Ecosystem models should incorporate direct and indirect effects of climate change on soil water content in order to accurately predict terrestrial feedbacks and long-term storage of C in soil.     

Photosynthesis and inorganic carbon transport in isolated asparagus mesophyll cells

The possibility of HCO₃⁻ transport into isolated leaf mesophyll cells of Asparagus sprengeri Regel has been investigated. Measurement of the inorganic carbon pool in these cells over an external pH range 6.2 to 8.0, using the silicone-fluid filtration technique, indicated that the pool was larger than predicted by passive ¹⁴CO₂ distribution, suggesting that HCO₃⁻ as well as CO₂ crosses the plasmalemma. Intracellular pH values, calculated from the distribution of ¹⁴CO₂ between the cells and the medium, were found to be higher (except at pH 8.0) than those previously determined by 5,5-dimethyl[2-¹⁴C]oxazolidine-2,4-dione distribution. It is suggested that the inorganic carbon accumulated above predicted concentrations may be bound to proteins and membranes and thus may not represent inorganic carbon actively accumulated by the cells, inasmuch as in a closed system at constant CO₂ concentration, the photosynthetic rates at pH 7.0 and 8.0 were 5 to 8 times lower than the maximum rate which could be supported by CO₂ arising from the spontaneous dehydration of HCO₃⁻. Furthermore, CO₂ compensation points of the cells in liquid media at 21% O₂ at pH 7.0 and 8.0, and the K_½ CO₂ (CO₂ concentration supporting the half maximal rate of O₂ evolution) at 2% O₂ at pH 7.0 and 8.0 are not consistent with HCO₃⁻ transport. These results indicate that the principal inorganic carbon species crossing the plasmalemma in these cells is CO₂.     

Forms of Dissolved Carbon Dioxide Required for Photosystem II Activity in Chloroplast Membranes

High concentrations of both bicarbonate and formate inhibit photosynthetic O₂ evolution at pH 8.0. At this pH, only 2.4% of the total dissolved carbon dioxide exists as CO₂. At pH 7.3, where 11% of the total dissolved carbon dioxide exists as CO₂, HCO₃⁻ no longer inhibits. While formate still inhibits O₂ evolution at pH 7.3, its effect can be partially overcome if CO₂ is also present. The rate of binding of added ¹⁴C-labeled inorganic carbon is nearly 10-fold more rapid when the internal pH of thylakoid membranes is at 6.0 than when it is at 7.8. These observations suggest that CO₂, not HCO₃⁻, is initially bound to the photosystem II reaction center and that the location of the binding site is on the inside thylakoid surface. However, additional data presented here suggest that, after binding, CO₂ is hydrated to HCO₃⁻ + H⁺ in a pH-dependent reaction. Two possible explanations of the “bicarbonate effect” are presented.     

Effects of six years atmospheric CO2 enrichment on plant,soil, and soil microbial C of a calcareous grassland

Stimulated plant production and often even larger stimulation of photosynthesis at elevated CO₂ raise the possibility of increased C storage in plants and soils. We analysed ecosystem C partitioning and soil C fluxes in calcareous grassland exposed to elevated CO₂ for 6 years. At elevated CO₂, C pools increased in plants (+23%) and surface litter (+24%), but were not altered in microbes and soil organic matter. Soils were fractionated into particle size and density separates. The amount of low-density macroorganic C, an indicator of particulate soil C inputs from root litter, was not affected by elevated CO₂. Incorporation of C fixed during the experiment (C_new) was tracked by C isotopic analysis of soil fractions which were labelled due to ¹³C depletion of the commercial CO₂ used for atmospheric enrichment. This data constrains estimates of C sequestration (absolute upper bound) and indicates where in soils potentially sequestered C is stored. C_new entered soils at an initial rate of 210±42 g C m^–2 year^–1, but only 554±39 g C_new m^–2 were recovered after 6 years due to the low mean residence time of 1.8 years. Previous process-oriented measurements did not indicate increased plant–soil C fluxes at elevated CO₂ in the same system (¹³C kinetics in soil microbes and fine roots after pulse labelling, and minirhizotron observations). Overall experimental evidence suggests that C storage under elevated CO₂ occurred only in rapidly turned-over fractions such as plants and detritus, and that potential extra soil C inputs were rapidly re-mineralised. We argue that this inference does not conflict with the observed increases in photosynthetic fixation at elevated CO₂, because these are not good predictors of plant growth and soil C fluxes for allometric reasons. C sequestration in this natural system may also be lower than suggested by plant biomass responses to elevated CO₂ because C storage may be limited by stabilisation of C_new in slowly turned-over soil fractions (a prerequisite for long-term storage) rather than by the magnitude of C inputs per se.     

Estimated Drainage of Carbon from the Tricarboxylic Acid Cycle for Protein Synthesis in Suspension Cultures of Paul's Scarlet Rose Cells

The amount of carbon (μmoles of carbon atoms) drained from the tricarboxylic acid cycle for protein synthesis was compared with μmoles of CO₂ released from the cycle at 2-day intervals during the growth of suspension cultures of Paul's Scarlet rose. We concluded that during the period of most rapid protein synthesis (day 0-4) one-sixth as much carbon was drained from the tricarboxylic acid cycle for protein synthesis as was released as CO₂. By day 8, one-thirtieth of the amount of carbon released as CO₂ was incorporated into protein. Net protein synthesis stopped on day 8, but the evolution of CO₂/culture continued at its maximum rate until day 10.     

Higher than expected CO2 fertilization inferred from leaf to global observations

Several lines of evidence point to an increase in the activity of the terrestrial biosphere over recent decades, impacting the global net land carbon sink (NLS) and its control on the growth of atmospheric carbon dioxide (c_a). Global terrestrial gross primary production (GPP)—the rate of carbon fixation by photosynthesis—is estimated to have risen by (31 ± 5)% since 1900, but the relative contributions of different putative drivers to this increase are not well known. Here we identify the rising atmospheric CO₂ concentration as the dominant driver. We reconcile leaf‐level and global atmospheric constraints on trends in modeled biospheric activity to reveal a global CO₂ fertilization effect on photosynthesis of 30% since 1900, or 47% for a doubling of c_a above the pre‐industrial level. Our historic value is nearly twice as high as current estimates (17 ± 4)% that do not use the full range of available constraints. Consequently, under a future low‐emission scenario, we project a land carbon sink (174 PgC, 2006–2099) that is 57 PgC larger than if a lower CO₂ fertilization effect comparable with current estimates is assumed. These findings suggest a larger beneficial role of the land carbon sink in modulating future excess anthropogenic CO₂ consistent with the target of the Paris Agreement to stay below 2°C warming, and underscore the importance of preserving terrestrial carbon sinks.