Regulation of photosynthesis: Analysis of a model for sensitivity of delayed luminescence oscillation and the CO<Subscript>2</Subscript> fixation rate to variation of the model parameters

The oscillation behavior of delayed luminescence was addressed using the earlier proposed mathematical model [Karavaev and Kukushkin, Biofizika 38, 958 (1993)]. The oscillation frequency and damping factor were calculated by Lyapunov analysis. In calculations, each of the model parameters was varied in a broad range. The results suggest that, besides the oscillation mode observed experimentally, there may be other oscillations that are more rapidly damped. Analysis of how variation in the model parameters affects the CO₂ fixation rate has shown that CO₂ assimilation differently depends on the light absorbed by photosystems I and II.     

Relationships between quantum yield for CO2 assimilation, activity of key enzymes and CO2 leakiness in Amaranthus cruentus, a C4 dicot, grown in high or low light

In C(4) photosynthesis, a part of CO(2) fixed by phosphoenolpyruvate carboxylase (PEPC) leaks from the bundle-sheath cells. Because the CO(2) leak wastes ATP consumed in the C(4) cycle, the leak may decrease the efficiency of CO(2) assimilation. To examine this possibility, we studied the light dependence of CO(2) leakiness (phi), estimated by the concurrent measurements of gas exchange and carbon isotope discrimination, initial activities of ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) and pyruvate, orthophosphate dikinase (PPDK), the phosphorylation state of PEPC and the CO(2) assimilation rate using leaves of Amaranthus cruentus (NAD-malic enzyme subtype, dicot) plants grown in high light (HL) and low light (LL). phi was constant at photon flux densities (PFDs) >200 micromol m(-2) s(-1) and was around 0.3. At PFDs <150 micromol m(-2) s(-1), phi increased markedly as PFD decreased. At 40 micromol m(-2) s(-1), phi was 0.76 in HL and 0.55 in LL leaves, indicating that the efficiency of CO(2) assimilation at low PFD was greater in LL leaves. The activities of Rubisco and PPDK, and the phosphorylated state of PEPC all decreased as PFD decreased. Theoretical calculations with a mathematical model clearly showed that the increase in phi with decreasing PFD contributed to the decrease in the CO(2) assimilation rate. It was also shown that the 'conventional' quantum yield of photosynthesis obtained by fitting the straight line to the light response curve of the CO(2) assimilation rate at the low PFD region is seriously overestimated. Ecological implications of the increase in phi in LL are discussed.     

Oscillatory response of photosynthesis in leaves to environmental perturbations: a mathematical model

Oscillations in the yield of chlorophyll fluorescence, in oxygen evolution, and in CO2 uptake observed with leaves upon perturbation of steady-state conditions are suggested to be due to the interdependence of turnover of adenylates and Calvin cycle intermediates. This suggestion is quantified in a mathematical model; the behavior of the model system in the neighborhood of the singular point of the system is analyzed. The linearized system is solved analytically, a condition for the occurrence of oscillations is given, and explicit expressions for the oscillation period and the damping constant are derived. The model is shown to be capable of exhibiting oscillations with the period observed with algae or leaves, whereas calculated values of the damping constant are higher than those measured for leaves or algae.     

Optimizing the statistical estimation of the parameters of the Farquhar-von Caemmerer-Berry model of photosynthesis

The model of Farquhar, von Caemmerer and Berry is the standard in relating photosynthetic carbon assimilation and concentration of intercellular CO(2). The techniques used in collecting the data from which its parameters are estimated have been the object of extensive optimization, but the statistical aspects of estimation have not received the same attention. The model segments assimilation into three regions, each modeled by a distinct function. Three parameters of the model, namely the maximum rate of Rubisco carboxylation (V(c max)), the rate of electron transport (J), and nonphotorespiratory CO(2) evolution (R(d)), are customarily estimated from gas exchange data through separate fitting of the component functions corresponding to the first two segments. This disjunct approach is problematic in requiring preliminary arbitrary subsetting of data into sets believed to correspond to each region. It is shown how multiple segments can be estimated simultaneously, using the entire data set, without predetermination of transitions by the investigator. Investigation of the number of parameters that can be estimated in the two-segment model suggests that, under some conditions, it is possible to estimate four or even five parameters, but that only V(c max), J, and R(d), have good statistical properties. Practical difficulties and their solutions are reviewed, and software programs are provided.     

The mechanism of acetate assimilation in purple nonsulfur bacteria lacking the glyoxylate pathway: enzymes of the citramalate cycle in Rhodobacter sphaeroides

The mechanism of acetate assimilation by the purple nonsulfur bacterium Rhodobacter sphaeroides, which lacks the glyoxylate shortcut, has been studied. In a previous work, proceeding from data on acetate assimilation by Rba. sphaeroides cell suspensions, a suggestion was made regarding the operation, in this bacterium, of the citramalate cycle. This cycle was earlier found in Rhodospirillum rubrum in the form of an anaplerotic reaction sequence that operates during growth on acetate instead of the glyoxylate shortcut, which is not present in the latter bacterium. The present work considers the enzymes responsible for acetate assimilation in Rba. sphaeroides. It is shown that this bacterium possesses the key enzymes of the citramalate cycle: citramalate synthase, which catalyzes condensation of acetyl-CoA and pyruvate and, as a result, forms citramalate, and 3-methylmalyl-CoA lyase, which catalyzes the cleavage of 3-methylmalyl-CoA to glyoxylate and propionyl-CoA. The regeneration of pyruvate, which is the acetyl-CoA acceptor in the citramalate cycle, involves propionyl-CoA and occurs via the following reaction sequence: propionyl-CoA (+ CO2) --> methylmalonyl-CoA --> succinyl-CoA --> succinate --> fumarate --> malate --> oxalacetate (- CO2) --> phosphoenolpyruvate --> pyruvate. The independence of the cell growth and the acetate assimilation of CO2 is due to the accumulation of CO2/HCO3- (released during acetate assimilation) in cells to a level sufficient for the effective operation of propionyl-CoA carboxylase.     

Transpiration and assimilation of early Devonian land plants with axially symmetric telomes-simulations on the tissue level

Early terrestrial ancestors of the land flora are characterized by a simple, axially symmetric habit and evolved in an atmosphere with much higher CO(2)concentrations than today. In order to gain information about the ecophysiological interrelationships of these plants, a model dealing with their gaseous exchange, which is basic to transpiration and photosynthesis, is introduced. The model is based on gas diffusion inside a porous medium and on a well-established photosynthesis model and allows for the simulation of the local gas fluxes through the various tissue layers of a plant axis. Necessary parameters consist of kinetical properties of the assimilation process and other physiological parameters (which have to be taken from extant plants), as well as physical constants and anatomical parameters which can be obtained from well-preserved fossil specimens. The model system is applied to an Early Devonian land plant, Aglaophyton major. The results demonstrate that, under an Early Devonian CO(2)concentration, A. major shows an extremely low transpiration rate and a low, but probably sufficiently high assimilation rate. Variation of the atmospheric CO(2)concentration shows that the assimilation is fully saturated even if the CO(2)content is decreased to about one-third of the initial value. This result indicates that A. major was probably able to exist under a wide range of atmospheric CO(2)concentrations. Further applications of this model system to ecophysiological studies of early land plant evolution are discussed.     

Use of heterotrophic CO2 assimilation as a measure of metabolic activity in planktonic and sessile bacteria

We have examined whether assimilation of CO2 can be used as a measure of metabolic activity in planktonic and sessile heterotrophic bacteria. CO2 assimilation by environmental samples and pure cultures of heterotrophic bacteria was studied using 14CO2 and 13CO2 as tracers. Heterotrophic growth on complex organic substrates resulted in assimilation of CO2 into cell biomass by activated sludge, drinking water biofilm, and pure cultures of Escherichia coli ATCC 25922, Es. coli ATCC 13706, Rhodococcus ruber, Burkholderia sp., Bacillus circulans, Pseudomonas putida, Pseudomonas stutzeri, and Pseudomonas aeruginosa. Analysis of 13C-labelled phospholipid fatty acids (PLFAs) confirmed that heterotrophic bacteria may assimilate 13CO2 into cell macromolecules such as membrane lipids. All major PLFAs extracted from activated sludge and drinking water biofilm samples were enriched in 13C after incubation with CO2. Between 1.4% and 6.5% of the biomass produced by cultures of P. putida and a drinking water biofilm during growth in complex media was apparently derived from assimilation of CO2. Resting cells assimilated less CO2 compared to actively growing cells, and CO2 assimilation activity correlated with the amount of biomass produced during heterotrophic growth. The 14CO2 assimilation assay was evaluated as a tool to examine inhibitory effects of biocides on planktonic and sessile heterotrophs (biofilms). On the basis of 14CO2 assimilation activity, the minimum inhibitory concentration (MIC) of benzalkonium chloride was estimated to 21.1 and 127.2 mg l(-1) for planktonic and biofilm samples, respectively. The results indicate that assimilation of isotopically labelled CO2 can be used as a relatively simple measure of metabolic activity in heterotrophic bacteria. CO2 assimilation assays may be used to study the effects of antimicrobial agents on growth and survival of planktonic and sessile heterotrophic organisms.     

典型城区与郊区环境大叶黄杨气体交换及叶绿素荧光特性比较

采用Li-6400便携式光合作用测定系统对夏秋季典型城区与郊区环境下大叶黄杨的气体交换和叶绿素荧光特性进行了现场实验比较研究.研究显示,叶片净光合速率的大小由总光合速率(光合能力)和呼吸速率共同决定,城区环境温度较高、相对湿度较低、大气CO2浓度较高, 不同月份城区和郊区样点大叶黄杨的净光合速率差异显著性存在不同.城区环境下大叶黄杨的胞间CO2浓度、叶面水气压亏缺、蒸腾速率高于郊区环境.城区环境中温度、大气CO2浓度等的变化会影响叶片呼吸作用,造成呼吸速率升高或是降低,城区环境中污染物浓度变化也会损伤叶片光合结构从而导致总光合能力降低,这两者都会引起净光合速率的变化.通过大叶黄杨叶片叶绿素荧光指标的进一步对比分析发现,城区大叶黄杨叶片叶绿素总量、叶绿素a/b、Fv/Fm、Fv/Fo、qP、ΦPSⅡ、ETR降低,但qN升高.表明叶片叶绿体PSⅡ的功能受到负面影响.城区大叶黄杨叶片荧光参数的变化,从微观机制上表明城区环境中污染物浓度的上升导致叶绿素及叶绿体光合结构受损的确是叶片光合能力下降的主要原因之一.     

Inorganic Carbon-Stimulated O2 Photoreduction Is Suppressed by NO2- Assimilation in Air-Grown Cells of Synechococcus UTEX 625

The effect of NO2- assimilation on O2 exchange and CO2 fixation of the cyanobacterium, Synechococcus UTEX 625, was studied mass spectrometrically. Upon addition of 1 mM inorganic carbon to the medium, inorganic carbon pools developed and accelerated O2 photoreduction 5-fold when CO2 fixation was inhibited. During steady-state photosynthesis at saturating light, O2 uptake represented 32% of O2 evolution and balanced that portion of O2 evolution that could not be accounted for by CO2 fixation. Under these conditions, NO2- assimilation reduced O2 uptake by 59% but had no influence on CO2 fixation. NO2- assimilation decreased both CO2 fixation and O2 photoreduction at low light and and increased net O2 evolution at all light intensities. The increase in net O2 evolution observed during simultaneous assimilation of carbon and nitrogen over carbon alone was due to a suppression of O2 photoreduction by NO2- assimilation. When CO2 fixation was precluded, NO2- assimilation inhibited O2 photoreduction and stimulated O2 evolution. When the electron supply was limiting (low light), competition among O2, CO2, and NO2- for electrons could be observed, but when the electron supply was not limiting (saturating light), O2 photoreduction and/or NO2- reduction caused electron transport that was additive to that for maximum CO2 fixation.     

Effects of irradiance and leaf water deficit on net carbon dioxide assimilation and mesophyll and transport resistances

Rate of net CO₂ assimilation by soil-grown soybean plants were studied over a range of relative leaf water contents at each of four levels of irradiance. There was a large interaction between light level and leaf water deficit on the rate of CO₂ assimilation. The effect of leaf water deficit on assimilation became larger as irradiance increased. Both stomatal resistance to CO₂ transport and mesophyll resistance to CO₂ assimilation increased as leaf-water deficit increased. The increase in both resistance with changing leaf-water content was largest at high irradiance and became smaller as irradiance decreased. Relief of soil-moisture stress by watering induced large oscillations of CO₂ assimilation, stomatal resistance, and mesophyll resistance. The oscillation of the mesophyll resistance occurred in the absence of changes in relative water content and appeared to be related to oscillations in leaf temperature. The observed increase in mesophyll resistance with decreasing leaf-water content under nonoscillative conditions may be caused by changes in leaf temperature rather than leaf water content.     

Method for measuring substrate preferences by individual members of microbial consortia proposed for bioaugmentation

In this study we used the assimilation of isotope labeled CO(2) to measure the substrate preferences by two different bioaugmentation mixtures proposed for bioremediation of diesel oil contamination. All active microorganisms assimilate CO(2) in various carboxylation processes involved in growth. The CO(2) assimilation by the two mixtures was measured upon addition of glucose, diesel oil or specific compounds present in diesel oil (naphthalene, toluene, hexadecane, and octane). It was shown that within short term incubations with diesel oil (<5 h), one bioaugmentation mixture was superior to the other regarding the assimilation of CO(2). This observation was confirmed in a labor-intensive long term microcosm study (60 days). The applied method open various possibilities for fast pre-testing of substrate-preferences by microbial-bioaugmentation mixtures without microcosm experiments, onsite tests, and complicated chemical analysis. This study also demonstrates the possibility to obtain further information on the substrate preferences at a single cell level of phylogenetically defined microbial subgroups in bioaugmentation mixtures, based on combined analyses of microautoradiography and fluorescence in situ hybridization.     

胡杨叶片气孔导度特征及其对环境因子的响应

依据2005年对极端干旱区荒漠河岸林胡杨的观测资料,对胡杨气孔运动进行了分析研究以揭示胡杨的水分利用特征与抗旱机理。结果表明:(1)胡杨叶片气孔导度日变化呈现为周期波动曲线,其波动周期为2 h,傍晚(20:00)波动消失;净光合速率和蒸腾速率与气孔导度的波动相对应而呈现同步周期波动。(2)胡杨的阳生叶气孔导度高于阴生叶,且不同季节气孔导度值不同,阳生叶气孔导度的季节变幅大于阴生叶。(3)胡杨气孔导度与气温、相对湿度和叶水势有显著相关关系,当CO2浓度较小时,胡杨气孔导度随CO2浓度的增加而增加,当CO2浓度达到一定值后气孔导度不再增加,反而随CO2浓度的增加大幅度降低。(4)胡杨适应极端干旱区生境的气孔调节机制为反馈式反应,即由于叶水势降低导致气孔导度减小,从而减少蒸腾耗水,达到节约用水、适应干旱的目的,表明胡杨的水分利用效率随气孔限制值的增大而减小,二者呈显著负相关。     

Formation of carbon monoxide from CO2 and H2 by Methanobacterium thermoautotrophicum

Cell suspensions of Methanobacterium thermoautotrophicum were found to reduce CO2 with H2 to CO at a maximal rate of 100 nmol X min-1 X mg protein-1. Half-maximal rates were obtained at a H2 and a CO2 concentration in the gas phase of 10% and 30%, respectively. The CO concentration in the gas phase surpassed the equilibrium concentration by a factor of more than 15 which indicates that CO2 reduction with H2 to CO was energy-driven. This was substantiated by the observation that the cells only formed CO when they also generated methane and that CO formation was completely inhibited by uncouplers. CO formation by cell suspensions and by growing cells was inhibited by cyanide. Neither methane formation nor the electrochemical proton potential were affected by this inhibitor. Cyanide was shown to inactivate specifically the carbon monoxide dehydrogenase present in M. thermoautotrophicum. It is therefore concluded that reduction of CO2 to CO is catalyzed by this enzyme. CO production by growing cells was 5-10-times slower than by resting cells. This is explained by effective CO assimilation in growing cells; when CO assimilation was inhibited by propyl iodide the rate of CO production immediately increased more than tenfold.     

Modeling the synchronization of yeast respiratory oscillations

The budding yeast Saccharomyces cerevisiae exhibits autonomous oscillations when grown aerobically in continuous culture with ethanol as the primary carbon source. A single cell model that includes the sulfate assimilation and ethanol degradation pathways recently has been developed to study these respiratory oscillations. We utilize an extended version of this single cell model to construct large cell ensembles for investigation of a proposed synchronization mechanism involving hydrogen sulfide. Ensembles with as many as 10,000 cells are used to simulate population synchronization and to compute transient number distributions from asynchronous initial cell states. Random perturbations in intracellular kinetic parameters are introduced to study the synchronization of single cells with small variations in their unsynchronized oscillation periods. The cell population model is shown to be consistent with available experimental data and to provide insights into the regulatory mechanisms responsible for the synchronization of yeast metabolic oscillations.     

Nitrate reduction and assimilation in Chlorella

1. Nitrate reduction and assimilation have been studied in Chlorella pyrenoidosa under growth conditions by observing effects on the CO(2)/O(2) gas exchange quotient. 2. During assimilation of glucose in the dark, nitrate reduction is noted as an increase in the R.Q. to about 1.6 caused by an increased rate of carbon dioxide production. 3. During photosynthesis at low light intensity nitrate reduction is evidenced by a reduction in the CO(2)O(2) quotient to about 0.7 caused by a decreased rate of carbon dioxide uptake. 4. Chlorella will assimilate nitrogen from either nitrate or ammonia. When both sources are supplied, only ammonia is utilized and no nitrate reduction occurs. It is inferred that under the usual conditions of growth nitrate is reduced only at a rate required for subsequent cellular syntheses. The effect of nitrate reduction on the CO(2)O(2) quotient therefore provides a measure of the relative rate of nitrogen assimilation. 5. Over-all photosynthetic metabolism may be described from elementary analysis of the cells since excretory products are negligible. The gas exchange predicted in this way is in good agreement with the observed CO(2)/O(2) quotients.     

Effect of the pacing interval and extrasystole on amplitude and damping of diastolic oscillations in the rabbit papillary muscle

Diastolic mechanical oscillations of right ventricular rabbit papillary muscles investigated at 15 degrees C in the perfusion chamber were analysed using a relaxation model with 6 parameters. From this analysis follows that the first diastolic oscillation amplitude plotted against the driving interval ("interval amplitude curve" of the first diastolic oscillation) shows tow maxima thus differing from the shape of the interval strength curve of the preceding driven contraction. It is concluded that the amplitude of diastolic oscillations is less determined by the Ca-amount released from the sarcoplasmic reticulum during the action potential than by the frequency of processes which effect the Ca release and the Ca-sequestration. The damping ratio varies within the range of tested driving intervals (0.36 to 10 s). After an extrasystole damping ratio and period of diastolic oscillation are diminished compared with the values after regularly driven contractions.     

Cellular glutathione redox homeostasis plays an important role in the brassinosteroid-induced increase in CO2 assimilation in Cucumis sativus

Brassinosteroids (BRs) play a vital role in plant growth, stress tolerance and productivity. Here, the involvement of BRs in the regulation of CO(2) assimilation and cellular redox homeostasis was studied. The effects of BRs on CO(2) assimilation were studied in cucumber (Cucumis sativus) through the analysis of the accumulation of H(2)O(2) and glutathione and photosynthesis-related enzyme activities using histochemical and cytochemical detection or a spectrophotometric assay, and Rubisco activase (RCA) using western blot analysis and immunogold labeling. Exogenous BR increased apoplastic H(2)O(2) accumulation, the ratio of reduced to oxidized glutathione (GSH:GSSG) and CO(2) assimilation, whereas a BR biosynthetic inhibitor had the opposite effects. BR-induced CO(2) assimilation was decreased by a H(2)O(2) scavenger or inhibition of H(2)O(2) generation, GSH biosynthesis and the NADPH-generating pentose phosphate pathway. BR-, H(2)O(2) - or GSH-induced CO(2) assimilation was associated with increased activity of enzymes in the Benson-Calvin cycle. Immunogold labeling and western blotting showed that BR increased the content of RCA and this effect was blocked by inhibitors of redox homeostasis. These results strongly suggest that BR-induced photosynthesis involves an H(2)O(2) -mediated increase in the GSH:GSSG ratio, which may positively regulate the synthesis and activation of redox-sensitive enzymes in carbon fixation.     

Computer simulation of damped oscillations during peroxidase-catalyzed oxidation of indole-3-acetic acid

Oscillation patterns in horseradish peroxidase (HRP)-catalyzed oxidation of indole-3-acetic acid (IAA) at neutral pH were studied using computer simulation. Under certain conditions, such as the presence of a reaction promoter and continuous intake of oxygen from the gaseous phase, the simulated system exhibits damped oscillations of the concentrations of oxygen in the aqueous phase, [O(2)](aq), and of all the reaction intermediates. The critical concentration of oxygen in aqueous phase, [O(2)](cr)(aq), was used to describe the nature of the oscillations. The critical concentration is the concentration at which the system abruptly changes its properties. If [O(2)](aq) is higher than [O(2)](cr)(aq) then the reaction develops as an avalanche, otherwise, the reaction stops. The nature of oscillations is accounted for by the interaction of two processes: the consumption/accumulation of oxygen and the accumulation/consumption of reaction intermediates. Oscillations are always damped. Neither HRP or umbelliferone (Umb) deactivation nor IAA consumption can account for the damping. The nature of the damping is determined by the termination reactions of free radical intermediates and ROOH. The three major parameters of oscillations: period of oscillations, initial amplitude of oscillations and the rate of damping were studied as functions of: (i) oxygen concentration in the gaseous phase, (ii) initial oxygen concentration in aqueous phase, (iii) the concentration of IAA and (iv) the initial concentration of HRP.     

Interrelationship of the mass transfer coefficient with regard to O2 and CO2 in the process of antibiotic biosynthesis]

A complex of studies on the effect of technological parameters on the mass transfer coeficients with respect to O2 and CO2 was carried out. It was shown that the ratio between the mass transfer coefficients with respect to O2 and CO2 was constant and equal to 20 for the fermentation broths of the antibiotic-producing organisms studied.     

Quantum nature of the damping of Langmuir oscillations and the boson peak in plasma

It is shown that the damping of Langmuir plasma oscillations is quantum in nature and that the damping rate, which is proportional to the fourth power of the electron charge, is caused by thermal electron fluctuations and depends nonanalytically on the Plank constant ℏ at ℏ → 0. At frequencies of ∼T/ℏ, the damping rate has a maximum, which can be identified with a boson peak.