Control of the rate of photosynthetic carbon dioxide fixation

A simplified model of the reductive pentose phosphate pathway of photosynthesis is analysed in order to quantify the degree to which each of the constituent reactions controls the rate of CO₂ fixation (given by the control coefficient). The analysis focuses on the four largely irreversible reactions of the cycle together with the first irreversible reaction in the sucrose and starch synthetic pathways. The model assumes that the other reactions are at equilibrium. The photorespiratory and electron transport systems are not included in the model. The analysis demonstrates that: (1) an analytical approach can be used to investigate the distribution of flux control in autocatalytic and moiety-conserved cycles; (2) measurements of enzyme kinetic parameters and certain fluxes and substrate concentrations can be used to solve the equations defining the enzyme control coefficients; (3) the conservation of total stromal phosphate and the intricate regulatory mechanisms of the photosynthetic system result in a relationship between the control coefficients that is complex and may defy any intuitive assessment of ‘rate limitation’; (4) ribulose-1,5-bisphosphate carboxylase / oxygenase may, under certain conditions, be a major controller of the rate of CO₂ fixation and, by regulating the concentration of ribulose 1,5-bisphosphate, may be important in governing the ratio of organic to inorganic phosphate in the stroma; (5) the other enzymes may also serve an important role in determining the distribution of phosphate between organic and inorganic species because they catalyze reactions at the branch points between starch and sucrose synthesis and ribulose 1,5-bisphosphate regeneration; (6) these enzymes that catalyze ‘branch-pint’ reactions may have negative control coefficients because of their ability to reduce the total concentration of cycle intermediates; (7) an approach combining the use of the equations presented in this paper and flux and substrate concentration measurements may be adequate for determining the control coefficients of several enzymes of the reductive pentose phosphate pathway.     

Maximum carbon uptake rate dominates the interannual variability of global net ecosystem exchange

Terrestrial ecosystems contribute most of the interannual variability (IAV) in atmospheric carbon dioxide (CO₂) concentrations, but processes driving the IAV of net ecosystem CO₂ exchange (NEE) remain elusive. For a predictive understanding of the global C cycle, it is imperative to identify indicators associated with ecological processes that determine the IAV of NEE. Here, we decompose the annual NEE of global terrestrial ecosystems into their phenological and physiological components, namely maximum carbon uptake (MCU) and release (MCR), the carbon uptake period (CUP), and two parameters, α and β, that describe the ratio between actual versus hypothetical maximum C sink and source, respectively. Using long‐term observed NEE from 66 eddy covariance sites and global products derived from FLUXNET observations, we found that the IAV of NEE is determined predominately by MCU at the global scale, which explains 48% of the IAV of NEE on average while α, CUP, β, and MCR explain 14%, 25%, 2%, and 8%, respectively. These patterns differ in water‐limited ecosystems versus temperature‐ and radiation‐limited ecosystems; 31% of the IAV of NEE is determined by the IAV of CUP in water‐limited ecosystems, and 60% of the IAV of NEE is determined by the IAV of MCU in temperature‐ and radiation‐limited ecosystems. The Lund‐Potsdam‐Jena (LPJ) model and the Multi‐scale Synthesis and Terrestrial Model Inter‐comparison Project (MsTMIP) models underestimate the contribution of MCU to the IAV of NEE by about 18% on average, and overestimate the contribution of CUP by about 25%. This study provides a new perspective on the proximate causes of the IAV of NEE, which suggest that capturing the variability of MCU is critical for modeling the IAV of NEE across most of the global land surface.     

Calibration of simultaneous measurements of photosynthetic carbon dioxide uptake and oxygen evolution in leaves

The stoichiometric ratio of O2 evolution to CO2 uptake during photosynthesis reveals information about reductive metabolism, including the reduction of alternative electron acceptors, such as nitrite and oxaloacetate. Recently we reported that in simultaneous measurements of CO2 uptake and O2 evolution in a sunflower leaf, O2 evolution changed by 7% more than CO2 uptake when light intensity was varied. Since the O2/CO2 exchange ratio is approximately 1, small differences are important. Thus, these gas exchange measurements need precise calibration. In this work, we describe a new calibration procedure for such simultaneous measurements, based on the changes of O2 concentration caused by the addition of pure CO2 or O2 into a flow of dry air (20.95% O2) through one and the same capillary. The relative decrease in O2 concentration during the addition of CO2 and the relative increase in O2 concentration during the addition of O2 allowed us to calibrate the CO2 and O2 scales of the measurement system with an error (relative standard deviation, RSD) of <1%. Measurements on a sunflower leaf resulted in an O2/CO2 ratio between 1.0 and 1.03 under different CO2 concentrations and light intensities, in the presence of an ambient O2 concentration of 20-50 micromol mol(-1). This shows that the percentage use of reductive power from photochemistry in synthesis of inorganic or organic matter other than CO2 assimilation in the C3 cycle is very low in mature leaves and, correspondingly, the reduction of alternative acceptors is a weak source of coupled ATP synthesis.     

Productivity, carbon dioxide uptake and net energy return of microalgal bubble column photobioreactors

This work examined the energy return of Chlorella vulgaris and Dunaliella tertiolecta cultivated in a gas-sparged photobioreactor design where the power input for sparging was manipulated (10, 20, and 50 W m⁻³). Dry weight, organic carbon and heating values of the biomass were measured, plus a suite of variables including Fv/Fm and dissolved oxygen. A model for predicting the higher heating value of microalgal biomass was developed and used to measure the energetic performance of batch cultivations. High power inputs enhanced maximum biomass yields, but did not improve the energy return. Cultivation in 10 W m⁻³ showed up to a 39% higher cumulative net energy return than 50 W m⁻³, and increased the cumulative net energy ratio up to fourfold. The highest net energy ratio for power input was 19.3 (D. tertiolecta, 12% CO₂, 10 W m⁻³). These systems may be a sustainable method of biomass production, but their effectiveness is sensitive to operational parameters.     

Concomitant determination of folar nitrogen loss, net carbon dioxide uptake, and transpiration

A closed system was designed for concomitant determination of net CO₂ uptake, transpiration, and foliar nitrogen (N) loss in soybeans (Glycine max [L.] Merr.). The CO₂ uptake was monitored by infrared analysis of system air removed periodically in aliquots. Leaf vapors were trapped in calibrated tubes at Dry Ice temperature, and transpiration rate was determined from the quantity of condensate. Subsequent pyrochemiluminescent analysis of this condensate revealed reduced N forms, although a [ill] percentage (4 to 15%) of oxidized forms was found.     

The uptake of carbon dioxide by plant roots

Summary The uptake of C¹⁴O₂ by the roots of intact tomato plants from solution containing Na₂C¹⁴O₃ was studied at different light intensities as well as in darkness.Where plants had previously been starved for CO₂ for 12 hours, a higher rate of C¹⁴ uptake was observed than with plants which had been transferred directly from the soil to the radioactive solution.In general, the C¹⁴ content of the roots was slightly higher than that of the shoots. At light intensities under the compensation point and in darkness the C¹⁴ content of the shoots relative to the roots decreased. This was accompanied by release of C¹⁴O₂ during respiration, indicating that the absorbed C¹⁴ was readily translocated upwards and released as C¹⁴O₂ under these conditions. At light intensities above the compensation point no C¹⁴O₂ was released.     

The use of eddy covariance to infer the net carbon dioxide uptake of Brazilian rain forest

• 1 Eddy covariance measurements of CO₂ flux, based on four and six week campaigns in Rondôdnia, Brazil, have been used in conjunction with a model to scale up data to a whole year, and thus estimate the carbon balance of the tropical forest ecosystem, and the changes in carbon balance expected from small interannual variations in climatological conditions.
• 2 One possible source of error in this estimation arises from the difficulty in measuring fluxes under stably stratified meteorological conditions, such as occur frequently at night. Flux may be ‘lost’ because of low velocity advection, caused by nocturnal radiative cooling at sites on raised ground. Such effects may be detected by plotting the net ecosystem flux of CO₂, F_eco is a function of wind speed. If flux is ‘lost’ then F_eco is expected to decline with wind speed. In the present data set, this did not occur, and F_eco was similar to the nocturnal flux estimated independently from chamber measurements.
• 3 The model suggests that in 1992/3, the Gross Primary Productivity (GPP) was 203.3 mol C m^?2 y^?1 and ecosystem respiration was 194.8 mol C m^?2 y^?1, giving an ecosystem carbon balance of 8.5 mol C m^?2 y^?1, equivalent to a sink of 1.0 ton C ha^?1 y^?1. However, the sign and magnitude of this figure is very sensitive to temperature, because of the strong influence of temperature on respiration.
• 4 The model also suggests that the effect of temperature on the net carbon balance is strongly dependent on the partial pressure of CO₂.

     

Role of pyruvate kinase in non-specific changes of carbon photosynthetic metabolism, caused by the photophosphorylation inhibition]

The decrease in the level of NADP reduction in chloroplasts under injections of cofactors of pseudocyclic FMN photophosphorylation (vitamin K3 and methylviologen) into pea, tomato and cucumber leaves resulted in the decrease of 14CO2 autotrophic accumulation rate and in the change of distribution of assimilated carbon among the assimilation products. The inhibition of synthesis of labelled sugars and the increase of the content of 3-phosphoglyceric and glyceric acids in the labelled low molecular fraction were observed in all the experiments. Similar changes in the distribution of synthesized in Kalvin cycle labelled products, associated with the inhibition of its reduction unit, were observed under the effect of photophosphorylation uncoupling agents. However, the decrease of ATP/NADP ratio in chloroplasts resulted in the simultaneous increase of fixed 14CO2 incorporation into alanine. The role of pyruvate kinase in "alanine" effect, characteristic of non-specific changes of carbon photosynthetic metabolism, is discussed on the basis of the authors' previous data on the effect of phosphorylation on glycolysis reactions and on the basis of results of ADP introduction into leaf cuts.     

Radiative forcing of methane fluxes offsets net carbon dioxide uptake for a tropical flooded forest

Wetlands are important sources of methane (CH₄) and sinks of carbon dioxide (CO₂). However, little is known about CH₄ and CO₂ fluxes and dynamics of seasonally flooded tropical forests of South America in relation to local carbon (C) balances and atmospheric exchange. We measured net ecosystem fluxes of CH₄ and CO₂ in the Pantanal over 2014–2017 using tower‐based eddy covariance along with C measurements in soil, biomass and water. Our data indicate that seasonally flooded tropical forests are potentially large sinks for CO₂ but strong sources of CH₄, particularly during inundation when reducing conditions in soils increase CH₄ production and limit CO₂ release. During inundation when soils were anaerobic, the flooded forest emitted 0.11 ± 0.002 g CH₄‐C m^?2 d^?1 and absorbed 1.6 ± 0.2 g CO₂‐C m^?2 d^?1 (mean ± 95% confidence interval for the entire study period). Following the recession of floodwaters, soils rapidly became aerobic and CH₄ emissions decreased significantly (0.002 ± 0.001 g CH₄‐C m^?2 d^?1) but remained a net source, while the net CO₂ flux flipped from being a net sink during anaerobic periods to acting as a source during aerobic periods. CH₄ fluxes were 50 times higher in the wet season; DOC was a minor component in the net ecosystem carbon balance. Daily fluxes of CO₂ and CH₄ were similar in all years for each season, but annual net fluxes varied primarily in relation to flood duration. While the ecosystem was a net C sink on an annual basis (absorbing 218 g C m^?2 (as CH₄‐C + CO₂‐C) in anaerobic phases and emitting 76 g C m^?2in aerobic phases), high CH₄ effluxes during the anaerobic flooded phase and modest CH₄ effluxes during the aerobic phase indicate that seasonally flooded tropical forests can be a net source of radiative forcings on an annual basis, thus acting as an amplifying feedback on global warming.     

Diffusion-based process for carbon dioxide uptake and isoprene emission in gaseous/aqueous two-phase photobioreactors by photosynthetic microorganisms

Photosynthesis for the generation of fuels and chemicals from cyanobacteria and microalgae offers the promise of a single host organism acting both as photocatalyst and processor, performing sunlight absorption and utilization, as well as CO(2) assimilation and conversion into product. However, there is a need to develop methods for generating, sequestering, and trapping such bio-products in an efficient and cost-effective manner that is suitable for industrial scale-up and exploitation. A sealed gaseous/aqueous two-phase photobioreactor was designed and applied for the photosynthetic generation of volatile isoprene (C(5)H(8)) hydrocarbons, which operates on the principle of spontaneous diffusion of CO(2) from the gaseous headspace into the microalgal or cyanobacterial-containing aqueous phase, followed by photosynthetic CO(2) assimilation and isoprene production by the transgenic microorganisms. Volatile isoprene hydrocarbons were emitted from the aqueous phase and were sequestered into the gaseous headspace. Periodic replacement (flushing) of the isoprene (C(5)H(8)) and oxygen (O(2)) content of the gaseous headspace with CO(2) allowed for the simultaneous harvesting of the photoproducts and replenishment of the CO(2) supply in the gaseous headspace. Reduction in practice of the gaseous/aqueous two-phase photobioreactor is offered in this work with a fed-batch and a semi-continuous culturing system using Synechocystis sp. PCC 6803 heterologously expressing the Pueraria montana (kudzu) isoprene synthase (IspS) gene. Constitutive isoprene production was observed over 192 h of experimentation, coupled with cyanobacterial biomass accumulation. The diffusion-based process in gaseous/aqueous two-phase photobioreactors has the potential to be applied to other high-value photosynthetically derived volatile molecules, emanating from a variety of photosynthetic microorganisms.     

Mechanism of aluminum inhibition of net ca uptake by amaranthus protoplasts

Calcium ions serve as a second messenger in signal transduction and metabolic regulation. Effects of Al on calcium homeostasis remain to be elucidated. Short-term net ⁴⁵Ca²⁺ uptake by Amaranthus tricolor protoplasts was monitored from uptake media prepared to test the influence of pH, Al, and various inhibitors. Accumulation of ⁴⁵Ca²⁺ increased during the first 3 to 6 minutes and then leveled off or declined. Al and Ca²⁺ channel blockers (verapamil and bepridil) decreased net ⁴⁵Ca²⁺ uptake. This decrease was more pronounced when Al and bepridil were both present in uptake media, but Al did not aggravate verapamil-induced reduction of net ⁴⁵Ca²⁺ uptake. Erythrosin B and calmidazolium each increased net ⁴⁵Ca²⁺ uptake, probably by interfering with Ca²⁺ efflux. This effect was undetectable in the presence of Al. Mycophenolic acid decreased net ⁴⁵Ca²⁺ uptake; guanosine alleviated this effect. Al-induced reduction of net ⁴⁵Ca²⁺ uptake was not aggravated by mycophenolic acid. Net ⁴⁵Ca²⁺ uptake was generally less at pH 4.5 than at 5.5 for all treatments. It is concluded that Al ions affect net ⁴⁵Ca²⁺ uptake by binding to the verapamil-specific channel site that is different from the bepridil-specific one, as well as by interfering with the action of guanosine 5′-triphosphate-binding proteins.     

Relations between K+ uptake and photosynthetic uptake of inorganic carbon by aquatic plants

The uptake of K⁺ by the leafy shoots of four submersed higher aquatic plants (Elodea canadensis, Ranunculus aquatilis, R. trichophyllus, and Callitriche hamulata) with different HCO₃ ^- affinity was measured in successive 2-h periods under the conditions of high or low photosynthetic rates (i.e. at pH 7.5 or 10). At pH 7.5 the uptake of K⁺ by species with the higher HCO₃ ^- affinity (E. canadensis, R. trichophyllus) was significantly faster than that by species with a lower HCO₃ ^- affinity (R. aquatilis, C. hamulata). In the former group of species, the K⁺ uptake rate at pH 7.5 was 1.7 - 3.5 times higher than at pH 10. At pH 10, the soft-water species, R. aquatilis, had the lowest net photosynthetic rate (PN) of the three HCO₃ ^- users but, in contrast to the relative hard-water species, R. trichophyllus, showed a small K⁺ efflux (47 nmol kg^-1 s^-1). Thus, K⁺ uptake by shoots was not strictly correlated with PN. A significant K⁺ efflux (73 - 86 nmol kg^-1 s^-1) occurred from all HCO₃ ^- users in darkness. The relatively low K⁺ uptake by the strict CO₂ user, C. hamulata, was quite independent of PN and light or darkness. It may be suggested that uptake of K⁺ by shoots of submersed plants depends on their HCO₃ ^- affinity. This revised version was published online in July 2006 with corrections to the Cover Date.     

Remote estimation of carbon dioxide uptake by a Mediterranean forest

The estimation of the carbon balance in ecosystems, regions, and the biosphere is currently one of the main concerns in the study of the ecology of global change. Current remote sensing methodologies for estimating gross primary productivity are not satisfactory because they rely too heavily on (i) the availability of climatic data, (ii) the definition of land‐use cover, and (iii) the assumptions of the effects of these two factors on the radiation‐use efficiency of vegetation (RUE). A new methodology is urgently needed that will actually assess RUE and overcome the problems associated with the capture of fluctuations in carbon absorption in space and over time. Remote sensing techniques such as the widely used reflectance vegetation indices (e.g. NDVI, EVI) allow green plant biomass and therefore plant photosynthetic capacity to be assessed. However, there are vegetation types, such as the Mediterranean forests, with a very low seasonality of these vegetation indices and a high seasonality of carbon uptake. In these cases it is important to detect how much of this capacity is actually realized, which is a much more challenging goal. The photochemical reflectance index (PRI) derived from freely available satellite information (MODIS sensor) presented for a 5‐year analysis for a Mediterranean forest a positive relationship with the RUE. Thus, we show that it is possible to estimate RUE and GPP in real time and therefore actual carbon uptake of Mediterranean forests at ecosystem level using the PRI. This conceptual and technological advancement would avoid the need to rely on the sometimes unreliable maximum RUE.     

Pyrophosphate inhibition of carbon dioxide fixation in isolated pea chloroplasts by uptake in exchange for endogenous adenine nucleotides

Carbon dioxide-dependent O(2) evolution by isolated pea (Pisum sativum) chloroplasts was inhibited by inorganic pyrophosphate (PPi). Oxygen evolution was also inhibited by high concentrations of orthophosphate (Pi) and the inhibition was relieved by 3-phosphoglycerate. In contrast, the inhibition by PPi was not relieved by 3-phosphoglycerate, indicating that hydrolysis of PPi and accumulation of inhibitory concentrations of Pi were not occurring. In agreement with this suggestion, the percentage of (14)C-labeled products diffusing out of the chloroplasts was increased by Pi but not by PPi. The inhibition of O(2) evolution by PPi was reversed by ATP. The concentration of PPi required for 50% inhibition was 1.2 to 1.4 mm and the subsequent stimulation by ATP was half-maximal at 16 to 25 mum. Carbon dioxide-dependent O(2) evolution by spinach chloroplasts, or chloroplasts isolated from older pea plants, was not significantly inhibited by PPi.Chloroplasts were preloaded with (14)C-ATP and release of the labeled nucleotides was measured to assess the activity of adenine nucleotide transport across the inner chloroplast envelope membrane. A rapid exchange was promoted by the addition of exogenous ATP. Addition of PPi also resulted in a release of endogenous nucleotides. We suggest that PPi inhibits CO(2) fixation by entering the chloroplast in exchange for endogenous adenine nucleotides via the transporter on the inner envelope membrane. The subsequent depletion of the internal adenine nucleotide pool would result in decreased CO(2) fixation due to insufficient ATP. Addition of ATP to PPi-inhibited chloroplasts apparently results in uptake of catalytic amounts of ATP and restoration of the internal adenine nucleotide pool thus relieving the inhibition of CO(2) fixation.     

Spring photosynthetic onset and net CO2 uptake in Alaska triggered by landscape thawing

The springtime transition to regional‐scale onset of photosynthesis and net ecosystem carbon uptake in boreal and tundra ecosystems are linked to the soil freeze–thaw state. We present evidence from diagnostic and inversion models constrained by satellite fluorescence and airborne CO₂ from 2012 to 2014 indicating the timing and magnitude of spring carbon uptake in Alaska correlates with landscape thaw and ecoregion. Landscape thaw in boreal forests typically occurs in late April (DOY 111 ± 7) with a 29 ± 6 day lag until photosynthetic onset. North Slope tundra thaws 3 weeks later (DOY 133 ± 5) but experiences only a 20 ± 5 day lag until photosynthetic onset. These time lag differences reflect efficient cold season adaptation in tundra shrub and the longer dehardening period for boreal evergreens. Despite the short transition from thaw to photosynthetic onset in tundra, synchrony of tundra respiration with snow melt and landscape thaw delays the transition from net carbon loss (at photosynthetic onset) to net uptake by 13 ± 7 days, thus reducing the tundra net carbon uptake period. Two global CO₂ inversions using a CASA‐GFED model prior estimate earlier northern high latitude net carbon uptake compared to our regional inversion, which we attribute to (i) early photosynthetic‐onset model prior bias, (ii) inverse method (scaling factor + optimization window), and (iii) sparsity of available Alaskan CO₂ observations. Another global inversion with zero prior estimates the same timing for net carbon uptake as the regional model but smaller seasonal amplitude. The analysis of Alaskan eddy covariance observations confirms regional scale findings for tundra, but indicates that photosynthesis and net carbon uptake occur up to 1 month earlier in evergreens than captured by models or CO₂ inversions, with better correlation to above‐freezing air temperature than date of primary thaw. Further collection and analysis of boreal evergreen species over multiple years and at additional subarctic flux towers are critically needed.