RELATIONSHIP BETWEEN PHOTOSYNTHETIC OXYGEN CYCLING AND CARBON ASSIMILATION IN SYNECHOCOCCUS WH7803 (CYANOPHYTA)1

Mass spectrometric analysis of oxygen uptake and evolution in the light by marine Synechococcus WH7803 indicated that the respiration rate was near zero at low irradiance levels but increased significantly at high irradiances. The light intensity (I_r) at which oxygen uptake began to increase with increasing light intensity depended on the growth irradiance of the culture. In each case, I_r coincided with the minimum light intensity for saturation of carbon assimilation (I_k). At irradiances >I_r, net oxygen evolution rates paralleled carbon assimilation rates. Oxygen uptake at high light intensities was inhibited by DCMU, indicating that oxygen uptake was due to Mehler reaction activity. The onset of Mehler activity at I_k supports the idea that oxygen becomes an alternative sink for electrons from photosystem I when NADPH turnover is limited by the capacity of the dark reactions to utilize reductant.     

Effect of growth temperature on the electron flow for photorespiration in leaves of tobacco grown in the field

Photorespiration has been indicated as an important mechanism for maintaining CO₂ assimilation and alleviating photodamage under conditions of high light and low CO₂. We tested the hypothesis that plants grown under a high temperature had greater electron flow for photorespiration compared with those grown under a relative low temperature. Responses of photosynthetic electron flow and CO₂ assimilation to incident light intensity and intercellular CO₂ concentration were examined in leaves of tobacco cultivar ‘k326’. Plants were cultivated at three sites with different ambient temperatures (Zhengzhou, Zunyi and Jiangchuan). Under high light, plants grown in Zhengzhou (with the highest growth temperature in the three sites) showed higher effective quantum yield of photosystem II and total electron flow through photosystem II than that in Zunyi and Jiangchuan. However, regardless of light intensity and intercellular CO₂ status, there were no significant differences among sites in the photosynthetic CO₂ assimilation rate or electron flow devoted to the carboxylation of ribulose‐1,5‐bisphosphate (RuBP). As a result, plants grown at high temperature showed higher electron flow devoted to oxygenation of RuBP than plants grown at low temperature. These results suggested that enhancement of electron flow for photorespiration is an important strategy in tobacco for acclimating to high growth temperature.     

Effects of growth and measurement light intensities on temperature dependence of CO2 assimilation rate in tobacco leaves

Effects of growth light intensity on the temperature dependence of CO₂ assimilation rate were studied in tobacco (Nicotiana tabacum) because growth light intensity alters nitrogen allocation between photosynthetic components. Leaf nitrogen, ribulose 1·5‐bisphosphate carboxylase/oxygenase (Rubisco) and cytochrome f (cyt f) contents increased with increasing growth light intensity, but the cyt f/Rubisco ratio was unaltered. Mesophyll conductance to CO₂ diffusion (g_m) measured with carbon isotope discrimination increased with growth light intensity but not with measuring light intensity. The responses of CO₂ assimilation rate to chloroplast CO₂ concentration (C_c) at different light intensities and temperatures were used to estimate the maximum carboxylation rate of Rubisco (V_cmax) and the chloroplast electron transport rate (J). Maximum electron transport rates were linearly related to cyt f content at any given temperature (e.g. 115 and 179 µmol electrons mol^?1 cyt f s^?1 at 25 and 40 °C, respectively). The chloroplast CO₂ concentration (C_trans) at which the transition from RuBP carboxylation to RuBP regeneration limitation occurred increased with leaf temperature and was independent of growth light intensity, consistent with the constant ratio of cyt f/Rubisco. In tobacco, CO₂ assimilation rate at 380 µmol mol^?1 CO₂ concentration and high light was limited by RuBP carboxylation above 32 °C and by RuBP regeneration below 32 °C.     

Interaction between Light Intensity and NaCl Salinity and Their Effects on Growth, CO(2) Assimilation, and Photosynthate Conversion in Young Broad Beans

Seedings of Vicia faba were grown for four weeks at two different light intensities (55 and 105 watts per square meter) in a saline (50 millimolar NaCl) and nonsaline nutrient solution. NaCl salinity depressed growth and restricted protein formation, CO₂ assimilation, and especially the incorporation of photosynthates into the lipid fraction. Conversion of photosynthates in leaves was much more affected by salinity than was photosynthate turnover in roots. The detrimental effect of NaCl salinity on growth, protein formation, and CO₂ assimilation was greater under low than under high light conditions. Plants of the high light intensity treatment were more capable of excluding Na⁺ and Cl⁻ and accumulating nutrient cation species (Ca²⁺, K⁺, Mg²⁺) than plants grown under low light intensity. It is suggested that the improved ionic status provided better conditions for protein synthesis, CO₂ assimilation, and especially for the conversion of photosynthates into lipids.     

Increased CO2 and light intensity regulate growth and leaf gas exchange in tomato

Carbon dioxide concentration (CO₂) and light intensity are known to play important roles in plant growth and carbon assimilation. Nevertheless, the underlying physiological mechanisms have not yet been fully explored. Tomato seedlings (Solanum lycopersicum Mill. cv. Jingpeng No. 1) were exposed to two levels of CO₂ and three levels of light intensity and the effects on growth, leaf gas exchange and water use efficiency were investigated. Elevated CO₂ and increased light intensity promoted growth, dry matter accumulation and pigment concentration and together the seedling health index. Elevated CO₂ had no significant effect on leaf nitrogen content but did significantly upregulate Calvin cycle enzyme activity. Increased CO₂ and light intensity promoted photosynthesis, both on a leaf-area basis and on a chlorophyll basis. Increased CO₂ also increased light-saturated maximum photosynthetic rate, apparent quantum efficiency and carboxylation efficiency and, together with increased light intensity, it raised photosynthetic capacity. However, increased CO₂ reduced transpiration and water consumption across different levels of light intensity, thus significantly increasing both leaf-level and plant-level water use efficiency. Among the range of treatments imposed, the combination of increased CO₂ (800 µmol CO₂ mol⁻¹) and high light intensity (400 µmol m⁻² s⁻¹) resulted in optimal growth and carbon assimilation. We conclude that the combination of increased CO₂ and increased light intensity worked synergistically to promote growth, photosynthetic capacity and water use efficiency by upregulation of pigment concentration, Calvin cycle enzyme activity, light energy use and CO₂ fixation. Increased CO₂ also lowered transpiration and hence water usage.     

Photosynthetic Assimilation of NO(3) by Intact Cells of the Cyanobacterium Anacystis nidulans: Influence of NO(3) and NH(4) Assimilation on CO(2) Fixation

Illuminated suspensions of Anacystis nidulans, supplied with saturating concentrations of CO₂ evolved O₂ at a greater rate when nitrate was simultaneously present. The extent of the stimulation of noncyclic electron flow induced by nitrate was dependent on light intensity, being maximal under light saturating conditions. Accordingly, nitrate depressed the rate of CO₂ fixation at limiting but not at saturating light, this depression reflecting the competition between both processes for assimilatory power. In contrast, ammonium stimulated CO₂ fixation at any light intensity assayed, the stimulation being dependent on the incorporation of ammonium to carbon skeletons. The positive effect of ammonium on CO₂ fixation also appeared to occur when nitrate was the nitrogen source, since with either nitrogen source an increase in the incorporation of newly fixed carbon into acid-soluble metabolites took place. From these results, the in vivo partitioning of assimilatory power between photosynthetic nitrogen and carbon assimilation and the quantitative and qualitative effects of inorganic nitrogen assimilation on CO₂ fixation are discussed.     

Enhancement of photosynthetic CO2 assimilation in the absence of oxygen,as dependent upon species and temperature

Summary The assimilation of CO₂ was enhanced in seven plant species by 44% at 30° in the absence of oxygen, and in three of the species by 85% at 40°. Net assimilation of CO₂ was not significantly greater in the absence of O₂ than it was in air with leaves of three tropical grasses and of one dicotyledonous species, Amaranthus palmeri. For two species in 30°, the enhanced CO₂ assimilation values were similar to those of the tropical grasses and of A. palmeri. The absence of O₂ did not enhance net CO₂ assimilation in maize even in light of low intensity.     

Responses to shading of naturalized and non-naturalized exotic woody species

Background and Aims

Recent studies have suggested that responses to shading gradients may play an important role in establishment success of exotic plants, but hitherto few studies have tested this. Therefore, a common-garden experiment was conducted using multiple Asian woody plant species that were introduced to Europe >100 years ago in order to test whether naturalized and non-naturalized species differ in their responses to shading. Specifically, a test was carried out to determine whether naturalized exotic woody species maintained better growth under shaded conditions, and whether they expressed greater (morphological and physiological) adaptive plasticity in response to shading, relative to non-naturalized species.

Methods

Nineteen naturalized and 19 non-naturalized exotic woody species were grown under five light levels ranging from 100 to 7 % of ambient light. For all plants, growth performance (i.e. biomass), morphological and CO₂ assimilation characteristics were measured. For the CO₂ assimilation characteristics, CO₂ assimilation rate was measured at 1200 μmol m^–2 s^–1 (i.e. saturated light intensity, A₁₂₀₀), 50 μmol m^–2 s^–1 (i.e. low light intensity, A₅₀) and 0 μmol m^–2 s^–1 (A₀, i.e. dark respiration).

Key Results

Overall, the naturalized and non-naturalized species did not differ greatly in biomass production and measured morphological and CO₂ assimilation characteristics across the light gradient. However, it was found that naturalized species grew taller and reduced total leaf area more than non-naturalized species in response to shading. It was also found that naturalized species were more capable of maintaining a high CO₂ assimilation rate at low light intensity (A₅₀) when grown under shading.

Conclusions

The results indicate that there is no clear evidence that the naturalized species possess a superior response to shading over non-naturalized species, at least not at the early stage of their growth. However, the higher CO₂ assimilation capacity of the naturalized species under low-light conditions might facilitate early growth and survival, and thereby ultimately favour their initial population establishment over the non-naturalized species.     

The Effect of Reduced Light Intensity and Sub-optimal Potassium Supply on N2 Fixation and N Turnover in Rhizobium Infected Lucerne

The effect of low light intensity and suboptimal potassium supply on the fixation of molecular nitrogen by root nodules and the nitrogen turnover in the host plant was studied in Medicago sativa using ¹⁵N labelled molecular nitrogen. For the application of ¹⁵N₂ labelled gas a special box was used. Both low light intensity and a low potassium supply resulted in a substantial growth depression. In particular the protein content of tops, roots and nodules was lower in the plants of the low light intensity treatment as compared with the control plants. Decreasing potassium supply had a similar but less-pronounced effect on protein content. The low protein content was not a consequence of a lack of soluble amino nitrogen or NH₃, since these fractions were influenced to a lesser degree by the reduced light intensity and by the low potassium supply. This observation is supported by the data obtained with ¹⁵N. N₂ fixation and NH₃ assimilation were affected by both low light intensity and low K application to the same degree as the overall metabolism, thus showing no particular response to the treatments applied.     

Distinctive Light and CO(2)-Fixation Requirements of Nitrate and Ammonium Utilization by the Cyanobacterium Anacystis nidulans

The effect of light intensity on the rates of ammonium and nitrate uptake and of CO₂ fixation has been determined in intact Anacystis nidulans cells. Ammonium uptake became saturated at photon flux values of about 60 microeinsteins per square meter per second, whereas both nitrate uptake and CO₂ fixation reached saturation at about 250 microeinsteins per square meter per second, the rates of the two latter processes being tightly correlated at any light intensity assayed. Inhibition of ammonium assimilation resulted in the loss of correlation between CO₂ fixation and nitrate uptake, the latter process exhibiting then a reduced light requirement. The results establish a clear distinction between ammonium utilization and nitrate utilization with regard to their light requirement and to the nature of their dependence upon CO₂ fixation.     

Changes in Levels of Intermediates of the C(4) Cycle and Reductive Pentose Phosphate Pathway under Various Light Intensities in Maize Leaves

The rate of CO₂ assimilation and levels of metabolites of the C₄ cycle and reductive pentose phosphate pathway in attached leaves of maize (Zea mays L.) were measured over a range of light intensity from 0 to 1,900 microEinsteins per square meter per second under a saturated CO₂ concentration of 350 microliters per liter and a limiting CO₂ concentration of 133 microliters per liter. The level of ribulose 1,5-bisphosphate (RuBP) stayed almost constant (around 60 nanomoles per milligram chlorophyll [Chl]) from low to high light intensities under 350 microliters per liter. Levels of 3-phosphoglycerate (PGA) increased from 100 to 650 nanomoles per milligram Chl under 350 microliters per liter CO₂ with increasing light intensity. The calculated RuBP concentration of 6 millimolar (corresponded to 60 nanomoles per milligram Chl) was about two times above the estimated RuBP binding-site concentration on ribulose bisphosphate carboxylase-oxygenase (Rubisco) of ~2.6 millimolar in maize bundle sheath chloroplasts in the light. The ratio of RuBP/PGA increased with decreasing light intensity under 350 microliters per liter CO₂. These results suggest that RuBP carboxylation is under control of light intensity possibly due to a limited supply of CO₂ to Rubisco through the C₄ cycle whose activity is highly dependent on light intensity. Pyruvate level increased with increasing light intensity as long as photosynthesis rate increased. A positive relationship between levels of PGA and those of pyruvate during steady-state photosynthesis under various conditions suggests that an elevated concentration of PGA increases the carbon input into the C₄ cycle through the conversion of PGA to PEP and consequently the level of total intermediates of the C₄ cycle can be raised to mediate higher photosynthesis rate.     

Contribution of Metabolites of Photosynthesis to Postillumination CO(2) Assimilation in Response to Lightflects

In the shade plant Alocasia macrorrhiza grown in low light, photosynthetic CO₂ assimilation during a 5 second lightfleck plus postillumination CO₂ assimilation can allow up to 60% more photosynthesis than that which occurs during 5 seconds of steady state light of the same intensity (RL Chazdon, RW Pearcy 1986 Oecologia. 69: 524-531). Metabolites of photosynthesis were measured to determine if the pool of ribulose 1,5-bisphosphate (RuBP) could account for all of the postillumination CO₂ assimilation following a lightfleck in Alocasia. It was found that the pool of triose-P was much larger than that of RuBP and could account for five times more postillumination CO₂ assimilation than could RuBP. The same trend was seen in the sun plant Phaseolus vulgaris when it was grown in the shade. In contrast, sun-grown Alocasia and Phasiolus did not have a large pool of triose-P relative to RuBP following a lightfleck. In sun plants, carbon may rapidly be converted to RuBP in the light whereas in shade plants there may be a restriction in the path between the triose-P and RuBP pools. It is hypothesized that in shade plants the buildup of triose-P rather than RuBP during the lightfleck prevents inhibition of electron transport which may otherwise occur because of competition for ATP between the two kinases of the photosynthetic carbon reduction cycle. Utilization of the triose-P for postillumination CO₂ fixation would require the capacity for significant postillumination ATP synthesis. The extensive grana stacking and large intrathylakoid space which accompanies the high level of chlorophyll in low-light-grown Alocasia could be an important contributing factor to postillumination ATP formation.     

Photoinhibition and photoprotection in senescent leaves of field-grown wheat plants

Photosynthesis, photosystem II (PSII) photochemistry, photoinhibition and the xanthophyll cycle in the senescent flag leaves of wheat (Triticum aestivum L.) plants grown in the field were investigated. Compared to the non-senescent leaves, photosynthetic capacity was significantly reduced in senescent flag leaves. The light intensity at which photosynthesis was saturated also declined significantly. The light response curves of PSII photochemistry indicate that a down-regulation of PSII photochemistry occurred in senescent leaves in particular at high light. The maximal efficiency of PSII photochemistry in senescent flag leaves decreased slightly when measured at predawn but substantially at midday, suggesting that PSII function was largely maintained and photoinhibition occurred in senescent leaves when exposed to high light. At midday, PSII efficiency, photochemical quenching and the efficiency of excitation capture by open PSII centers decreased considerably, while non-photochemical quenching increased significantly. Moreover, compared with the values at early morning, a greater decrease in CO₂ assimilation rate was observed at midday in senescent leaves than in control leaves. The levels of antheraxanthin and zeaxanthin via the de-epoxidation of violaxanthin increased in senescent flag leaves from predawn to midday. An increase in the xanthophyll cycle pigments relative to chlorophyll was observed in senescent flag leaves. The results suggest that the xanthophyll cycle was activated in senescent leaves due to the decrease in CO₂ assimilation capacity and the light intensity for saturation of photosynthesis and that the enhanced formation of antheraxanthin and zeaxanthin at high light may play an important role in the dissipation of excess light energy and help to protect photosynthetic apparatus from photodamage. Our results suggest that the well-known function of the xanthophyll cycle to safely dissipate excess excitation energy is also important for maintaining photosynthetic function during leaf senescence.     

Effect of Light Intensity during Growth on Photoinhibition of Intact Attached Bean Leaflets

In the study reported here, two different photoinhibitory phenomena were compared within a single plant species. Bean plants were grown in three different light intensities to simulate sun and shade environments. The effects of photoinhibitory treatments on in vivo CO₂ assimilation rates and in vitro chloroplast electron transport reactions were investigated and the extent to which carbon metabolism served to prevent photoinhibition was characterized. It was shown that the photoinhibition which follows exposure of intact leaflets of low light-grown bean plants to high light intensity in normal air is essentially similar to that which occurs when leaflets of plants grown in full sunlight are illuminated in the absence of CO₂ at low O₂ partial pressures.     

Mechanism of Photosynthesis Decrease by Verticillium dahliae in Potato

Young, visually symptomless leaves from potato (Solanum tuberosum) plants infected with Verticillium dahliae exhibited reduced carbon assimilation rate, stomatal conductance, and intercellular CO₂, but no increase in dark respiration, no change in the relationship between carbon assimilation rate versus intercellular CO₂, and no change in light use efficiency when intercellular CO₂ was held constant. Therefore, the initial decrease in photosynthesis caused by V. dahliae was caused by stomatal closure. Errors in the intercellular CO₂ calculation caused by uneven distribution of carbon assimilation rate across the leaf were tested by ¹⁴CO₂ autoradiography. Patchiness was found at a low frequency. Low stomatal conductance was correlated with low leaf water potentials. Infection did not affect leaf osmotic potentials.     

Analysis of the photosynthetic response induced by variation potential in geranium

Electrical signals (action and variation potentials) caused by environmental stimuli induce a number of physiological responses in plants including changes in photosynthesis; however, mechanisms of these changes remain unclear. We investigated the influence of the variation potential on photosynthesis in geranium (Pelargonium zonale) under different conditions (control, low external CO₂ concentration, and actinic light absence). The variation potential caused by lamina burning induced a reduction in photosynthesis (decreases in effective quantum yields of photosystem I and II, CO₂ assimilation rate, and stomatal conductance) in unstimulated leaves under control conditions. Changes in the majority of light-stage parameters (photosystem I and II quantum yields, coefficients of photochemical and non-photochemical quenching, quantum yield of non-photochemical energy dissipation in photosystem I due to donor-side limitation) were correlated with a decrease in CO₂ assimilation rate. The changes were similar to those caused by lowering [CO₂]; their magnitudes decreased both under low external CO₂ concentration and without actinic light. These results support the hypothesis that Calvin cycle inactivation plays a key role in photosynthetic response induced by electrical signals. However, a decrease in electron transport through the PSI acceptor side also induced by variation potential was not correlated with a decrease in the CO₂ assimilation rate and did not depend on the external CO₂ concentration or actinic light intensity. Thus, we suggest that there are two different mechanisms of light-stage inactivation induced by the variation potential in geranium: one strongly dependent on dark-stage inactivation and one weakly dependent on dark-stage inactivation.     

Physiological performances of the blue-green alga Anacystis nidulans in continuous turbidostat fermenter culture

A sterile continuous turbidostat culture in a 2-1 fermenter was used to systematically measure the gas exchange rates of Anacystis nidulans in a highly turbulent system under strictly controlled environmental conditions. An extensive physiological characterization of Anacystis is given in terms of photosynthesis rates (CO₂ uptake and O₂ evolution) and dark respiration rates as function of different parameters such as stirrer speed, temperature, CO₂ and O₂ concentration, light intensity, culture density and pH. Steady state ATP levels and apparent photophosphorylation rates complete the performance data. The dependence of the photosynthetic quotient from the parameters enables a physiological characterization of the light dependent nitrate assimilation.     

Responses to different external light rhythms by the circadian rhythm of Crassulacean acid metabolism in Kalanchoe daigremontiana

Time series of net CO₂ exchange ( J CO₂) and leaf conductivity for water vapour (gH₂O) were measured and subsequently analysed mathematically in the Crassulacean acid metabolism (CAM) plant Kalanchoe daigremontiana (Hamet et Perrier de la Bâthie) under constant environmental conditions and under imposed external rhythms of lower and higher light intensity. The time series were analysed by Fourier methods and a correlation analysis considering the first time derivatives of J CO₂, gH₂Oand photosynthetically active photon flux density (PPFD). The ratio of internal to external CO₂ (c_i/c_a) was also considered in the analysis, leading to a discussion of the interaction of stomata and carbon assimilation under periodic stimulation. It is suggested that for stimulation with frequencies close to the endogeneous circadian period, stomatal conductance and carbon assimilation oscillate synchronously, guard‐cell movements trailing behind changes in internal CO₂ with a delay of 10–15 min. For stimulation frequencies far shorter than the endogeneous period, this synchrony can be disturbed due to independent responses of stomata and assimilation to light pulses, leading to an arrhythmic gas exchange pattern. These results are discussed in the context of understanding circadian oscillations as the output of a multioscillator, multisignalling pathway system on the organismic and metabolic level.     

Rapid and Specific Modulation of Stomatal Conductance by Blue Light in Ivy (Hedera helix) : An Approach to Assess the Stomatal Limitation of Carbon Assimilation

Low intensity (0.015 millimole per square meter per second) blue light applied to leaves of Hedera helix under a high intensity red light background (0.50 millimole per square meter per second red light) induced a specific stomatal opening response, with rapid kinetics comparable to those previously reported for stomata with `grass type' morphology. The response of stomatal conductance to blue light showed a transient `overshoot' behavior at high vapor pressure difference (2.25 ± 0.15 kiloPascals), but not at low vapor pressure difference (VPD) (0.90 ± 0.10 kilo-Pascal). The blue light-induced conductance increase was accompanied by an increase in net photosynthetic carbon assimilation, mediated by an increase in the intercellular concentration of carbon dioxide. Values of assimilation once the blue light-stimulated conductance increase reached steady state were less than those at the peak of the overshoot, but the ratios of assimilation to transpiration (A/E) and blue light-stimulated ΔA/ΔE were greater during the steady-state response than during the overshoot. These results indicate that significant stomatal limitation of assimilation can occur, but that this limitation may improve water use efficiency under high VPD conditions. Under high intensity red light, the decline in A/E associated with an increase in VPD was minimized when conductance was stimulated by additional low intensity blue light. This effect indicates that the blue light response of stomata may be important in H. helix for the optimization of water use efficiency under natural conditions of high irradiance and VPD.     

A method for measuring whole plant photosynthesis in Arabidopsis thaliana

Measurement of photosynthesis of intact leaves of Arabidopsis thaliana has been prohibitive due to the small leaf size and prostrate growth habit. Because of the widespread use of Arabidopsis for plant science research it is important to have a procedure for accurate, nondestructive measurement of its photosynthesis. We developed and tested a method for analysis of photosynthesis in whole plants of Arabidopsis. Net carbon assimilation and stomatal conductance were measured with an open gas exchange system and photosynthetic oxygen evolution was determined from chlorophyll fluorescence parameters. Individual plants were grown in 50 cubic centimeter tubes that were attached with an air tight seal to an enclosed gas exchange chamber for measurement of carbon dioxide and water exchange by the whole plant. Chlorophyll fluorescence from intact leaves was simultaneously measured with a pulse modulated fluorometer. Photosynthetic CO₂ assimilation and stomatal conductance rates were calculated with established gas exchange procedures and O₂ evolution was determined from chlorophyll fluorescence measurement of Photosystem II yield. Carbon assimilation and oxygen evolution in response to light intensity and ambient CO₂ concentration was measured and is presented here to demonstrate the potential use of this method for investigation of photosynthesis of Arabidopsis plants in controlled environment conditions.