Showing papers on "Photosynthesis published in 1992"

Coupled Photosynthesis-Stomatal Conductance Model for Leaves of C4 Plants

Abstract: Leaf based models of net photosynthesis (An) and stomatal conductance (g) are often components of whole plant, canopy and regional models of net primary productivity and surface energy balance. Since C4 metabolism shows unique responses to environmental conditions and C4 species are important agriculturally and ecologically, a realistic and accurate leaf model specific to C4 plants is needed. In this paper we develop a simple model for predicting An and g from leaves of C4 plants that is easily parameterised and that predicts many of the important environmental responses. We derive the leaf model from a simple biochemical-intercellular transport model of C4 photosynthesis that includes inorganic carbon fixation by PEP carboxylase, light dependent generation of PEP and RuBP, rubisco reaction kinetics, and the diffusion of inorganic carbon and O2 between the bundle sheath and mesophyll. We argue that under most conditions these processes can be described simply as three potentially limiting steps. The leaf photosynthesis model treats An as first order with respect to either light, CO2 or the amount of rubisco present and produces a continuous transition between limitations. The independent variables of the leaf photosynthesis model are leaf temperature (TI), intercellular CO2 levels and the absorbed quantum flux. A simple linear model of g in terms of An and leaf surface CO2 level (ps) and relative humidity (hs) is combined with the photosynthesis model to give leaf photosynthesis as a function of absorbed quantum flux, T1 and ps and hs levels. Gas exchange measurements from corn leaves exposed to varied light, CO2 and temperature levels are used to parameterise and test the models. Model parameters are determined by fitting the models to a set of 21 measurements. The behaviour of the models is compared with an independent set of 71 measurements, and the predictions are shown to be highly correlated with the data. Under most conditions the leaf model can be parameterised simply by determining the level of rubisco in the leaves. The effects of light environment, nutritional status and water stress levels on An and g can be accounted for by appropriate adjustment of the capacity for rubisco to fix CO2. We estimate rubsico capacity from CO2 and light saturated photosynthesis although leaf nitrogen content or rubisco assays from leaf extracts could also be used for this purpose.

Modelling photosynthesis of cotton grown in elevated CO2

Abstract: Cotton plants were grown in CO2-controlled growth chambers in atmospheres of either 35 or 65 Pa CO2. A widely accepted model of C3 leaf photosynthesis was parameterized for leaves from both CO2 treatments using non-linear least squares regression techniques, but in order to achieve reasonable fits, it was necessary to include a phosphate limitation resulting from inadequate triose phosphate utilization. Despite the accumulation of large amounts of starch (>50 g m−2) in the high CO2 plants, the photosynthetic characteristics of leaves in both treatments were similar, although the maximum rate of Rubisco activity (Vcmax), estimated from A versus Ci response curves measured at 29°C, was ∼10% lower in leaves from plants grown in high CO2. The relationship between key model parameters and total leaf N was linear, the only difference between CO2 treatments being a slight reduction in the slope of the line relating Vcmax to leaf N in plants grown at high CO2. Stomatal conductance of leaves of plants grown and measured at 65 Pa CO2 was approximately 32% lower than that of plants grown and measured at 35 Pa. Because photosynthetic capacity of leaves grown in high CO2 was only slightly less than that of leaves grown in 35 Pa CO2, net photosynthesis measured at the growth CO2, light and temperature conditions was approximately 25% greater in leaves of plants grown in high CO2, despite the reduction in leaf conductance. Greater assimilation rate was one factor allowing plants grown in high CO2 to incorporate 30% more biomass during the first 36 d of growth.

Relationship between photosystem II activity and CO2 fixation in leaves

Abstract: There is now potential to estimate photosystem II (PSII) activity in vivo from chlorophyll fluorescence measurements and thus gauge PSII activity per CO2 fixed. A measure of the quantum yield of photosystem II, ΦII (electron/photon absorbed by PSII), can be obtained in leaves under steady-state conditions in the light using a modulated fluorescence system. The rate of electron transport from PSII equals ΦII times incident light intensity times the fraction of incident light absorbed by PSII. In C4 plants, there is a linear relationship between PSII activity and CO2 fixation, since there are no other major sinks for electrons; thus measurements of quantum yield of PSII may be used to estimate rates of photosynthesis in C4 species. In C3 plants, both CO2 fixation and photorespiration are major sinks for electrons from PSII (a minimum of 4 electrons are required per CO2, or per O2 reacting with RuBP). The rates of PSII activity associated with photosynthesis in C3 plants, based on estimates of the rates of carboxylation (vo) and oxygenation (vo) at various levels of CO2 and O2, largely account for the PSII activity determined from fluorescence measurements. Thus, in C3 plants, the partitioning of electron flow between photosynthesis and photorespiration can be evaluated from analysis of fluorescence and CO2 fixation.

Photosynthesis and Production in a Changing Environment: A Field And Laboratory Manual

Abstract: Introduction: Photosynthesis and the changing environment. Measurement of plant biomass and net primary production. remote sensing of biomass and productivity. Plant growth analysis. Plant Microclimate. Controlled-environment studies. Canopy structure and light interception. Functional leaf anatomy. Water relations. Measurement of CO2 assimilation by plants in the field and the laboratory. Polarographic measurement of oxygen. Carbon isotope discrimination. Chlorophyll fluorescence as a tool in photosynthesis research. Modelling - leaf to canopy. Models of canopy carbon and water balance: tools for data synthesis. Carbon partitioning. Carbon metabolism. Chloroplasts and protoplasts. Thylakoid components and processes. Nitrogen fixation and nitrate reduction. Ammonia assimilation, photorespiration and amino acid biosynthesis. Micro-algae: laboratory growth techniques and biotechnology of biomass production. References. Index.

Regulation of Photosynthesis by End-Product Accumulation in Leaves of Plants Storing Starch, Sucrose, and Hexose Sugars

Abstract: In the present study, leaves of different plant species were girdled by the hot wax collar method to prevent export of assimilates. Photosynthetic activity of girdled and control leaves was evaluated 3 to 7 days later by two methods: (a) carbon exchange rate (CER) of attached leaves was determined under ambient CO2 concentrations using a closed gas system, and (b) maximum photosynthetic capacity (Amax) was determined under 3% CO2 with a leaf disc O2 electrode. Starch, hexoses, and sucrose were determined enzymically. Typical starch storers like soybean (Glycine max L.) (up to 87.5 milligrams of starch per square decimeter in girdled leaves), cotton (Gossypium hirsutum L.), and cucumber (Cucumis sativus L.) responded to 7 days of girdling by increased (80-100%) stomatal resistance (rs) and decreased Amax (>50%). On the other hand, spinach (Spinacia oleracea L.), a typical sucrose storer (up to 160 milligrams of sucrose per square decimeter in girdled leaves), showed only a slight reduction in CER and almost no change in Amax. Intermediate plants like tomato (Lycopersicon esculentum Mill.), sunflower (Helianthus annuus L.), broad bean (Vicia faba L.), bean (Phaseolus vulgaris L.), and pea (Pisum sativum L.), which upon girdling store both starch and sucrose, responded to the girdle by a considerable reduction in CER but only moderate inhibition of Amax, indicating that the observed reduction in CER was primarily a stomatal response. Both the wild-type tobacco (Nicotiana sylvestris) (which upon girdling stored starch and hexoses) and the starchless mutant (which stored only hexoses, up to 90 milligrams per square decimeter) showed 90 to 100% inhibition of CER and approximately 50% inhibition of Amax. In general, excised leaves (6 days) behaved like girdled leaves of the respective species, showing 50% reduction of Amax in wild-type and starchless N. sylvestris but only slight decline of Amax in spinach. The results of the present study demonstrate the possibility of the occurrence of end-product inhibition of photosynthesis in a large number of crop plants. The long-term inhibition of photosynthesis in girdled leaves is not confined to stomatal responses since the Amax declined up to 50%. The inhibition of Amax by girdling was strongest in starch storers, but starch itself cannot be directly responsible, because the starchless mutant of N. sylvestris was also strongly inhibited. Similarly, the inhibition cannot be attributed to hexose sugars either, because soybean, cotton, and cucumber are among the plants most strongly inhibited although they do not maintain a large hexose pool. Spinach, a sucrose storer, showed the least inhibition in both girdled and excised leaf systems, which indicates that sucrose is probably not directly responsible for the end-product inhibition of photosynthesis. The occurrence of strong end-product inhibition appears to be correlated with high acid-invertase activity in fully expanded leaves. The inhibition may be related to the nature of soluble sugar metabolism in the extrachloroplastic compartment and may be caused by a metabolite that has different rates of accumulation and turnover in sucrose storers and other plants.

Photoinhibition of photosynthesis represents a mechanism for the long-term regulation of photosystem II.

Abstract: The obligate shade plant, Tradescantia albiflora Kunth grown at 50 μmol photons · m(-2) s(-1) and Pisum sativum L. acclimated to two photon fluence rates, 50 and 300 μmol · m(-2) · s(-1), were exposed to photoinhibitory light conditions of 1700 μmol · m(-2) · s(-1) for 4 h at 22° C. Photosynthesis was assayed by measurement of CO2-saturated O2 evolution, and photosystem II (PSII) was assayed using modulated chlorophyll fluorescence and flash-yield determinations of functional reaction centres. Tradescantia was most sensitive to photoinhibition, while pea grown at 300 μmol · m(-2) · s(-1) was most resistant, with pea grown at 50 μmol · m(-2) · s(-1) showing an intermediate sensitivity. A very good correlation was found between the decrease of functional PSII reaction centres and both the inhibition of photosynthesis and PSII photochemistry. Photoinhibition caused a decline in the maximum quantum yield for PSII electron transport as determined by the product of photochemical quenching (qp) and the yield of open PSII reaction centres as given by the steady-state fluorescence ratio, F'vF'm, according to Genty et al. (1989, Biochim. Biophys. Acta 990, 81-92). The decrease in the quantum yield for PSII electron transport was fully accounted for by a decrease in F'vF'm, since qp at a given photon fluence rate was similar for photoinhibited and noninhibited plants. Under lightsaturating conditions, the quantum yield of PSII electron transport was similar in photoinhibited and noninhibited plants. The data give support for the view that photoinhibition of the reaction centres of PSII represents a stable, long-term, down-regulation of photochemistry, which occurs in plants under sustained high-light conditions, and replaces part of the regulation usually exerted by the transthylakoid ΔpH gradient. Furthermore, by investigating the susceptibility of differently lightacclimated sun and shade species to photoinhibition in relation to qp, i.e. the fraction of open-to-closed PSII reaction centres, we also show that irrespective of light acclimation, plants become susceptible to photoinhibition when the majority of their PSII reaction centres are still open (i.e. primary quinone acceptor oxidized). Photoinhibition appears to be an unavoidable consequence of PSII function when light causes sustained closure of more than 40% of PSII reaction centres.

Stress Tolerance of Photosystem II in Vivo Antagonistic Effects of Water, Heat, and Photoinhibition Stresses

Abstract: The in vivo photochemical activity of photosystem II was inferred from modulated chlorophyll fluorescence and photoacoustic measurements in intact leaves of several plant species (Lycopersicon esculentum Mill., Solanum tuberosum L., Solanum nigrum L.) exposed to various environmental stresses (drought, heat, strong light) applied separately or in combination. Photosystem II was shown to be highly drought-resistant: even a drastic desiccation in air of detached leaf samples only marginally affected the quantum yield for photochemistry in photosystem II. However, water stress markedly modified the responses of photosystem II to superimposed constraints. The stability of photosystem II to heat was observed to increase strongly in leaves exposed to water stress conditions: heat treatments (e.g. 42°C in the dark), which caused a complete and irreversible inhibition of photosystem II in well-watered (tomato) leaves, resulted in a small and fully reversible reduction of the photochemical efficiency of photosystem II in drought-stressed leaves. In vivo photoacoustic data indicated that photosystem I was highly resistant to both heat and water stresses. When leaves were illuminated with intense white light at 25°C, photoinhibition damage of photosystem II was more pronounced in water-stressed leaves than in undesiccated controls. However, in nondehydrated leaves, photoinhibition of photosystem II was strongly temperature dependent, being drastically stimulated at high temperatures above 38 to 40°C. As a consequence, when exposed to strong light at high temperature, photosystem II photochemistry was significantly less inhibited in dehydrated leaves than in control well-hydrated leaves. Our results demonstrate the existence of a marked antagonism between physicochemical stresses, with water stress enhancing the resistance of photosystem II to constraints (heat, strong light at high temperature) that are usually associated with drought in the field.

The effect of water stress on photosynthetic carbon metabolism in four species grown under field conditions

Abstract: . The effect of gradually-developing water-stress has been studied in Lupinus albus L., Helianthus annuus L., Vitis vinifera cv. Rosaki and Eucalyptus globulus Labill. Water was withheld and diurnal rhythms were investigated 4–8d later, when the predawn water deficit was more negative than in watered plants, and the stomata closed almost completely early during the photoperiod. The contribution of ‘stomatal’ and ‘non-stomatal’ components to the decrease of photosynthetic rate was investigated by (1) comparing the changes of the rate of photosynthesis in air with the changes of stomatal conductance and (2) measuring photosynthetic capacity in saturating irradiance and 15% CO2. Three species (lupin, eucalyptus and sunflower) showed larger changes of stomatal conductance than photosynthesis in air, and showed little or no decrease of photosynthetic capacity in saturating CO2. Photosynthesis in air also recovered fully overnight after watering the plants in the evening. In grapevines, stomatal conductance and photosynthesis in air changed in parallel, there was a marked decrease of photosynthetic capacity, and photosynthesis and stomatal conductance did not recover overnight after watering water-stressed plants. Relative water content remained above 90% in grapevine. We conclude that non-stomatal components do not play a significant role in lupins, sunflower or eucalyptus, but could in grapevine. The effect of water-stress on partitioning of photosynthate was investigated by measuring the amounts of sucrose and starch in leaves during a diurnal rhythm, and by measuring the partitioning of 14C-carbon dioxide between sucrose and starch. In all four species, starch was depleted in water-stressed leaves but sucrose was maintained at amounts similar to, or higher than, those in watered plants. Partitioning into sucrose was increased in lupins and eucalyptus, and remained unchanged in grapevine and sunflower. It is concluded that water-stressed leaves in all four species maintain high levels of soluble sugars in their leaves, despite having lower rates of field photosynthesis, decreased rates of export, and low amounts of starch in their leaves.

Variations in the natural abundance of oxygen-18 and deuterium in plant carbohydrates

Abstract: Variations in the natural abundance of 18O and 2H in plant cellulose are influenced by the isotopic composition of the water directly involved in metabolism—the metabolic water fraction. The isotopic distinction between the metabolic source water and total tissue water must reflect the formation of isotopic gradients within the tissue that are influenced by the rate of water turnover, by properties of the water conducting system and by environmental conditions. It seems that the 18O abundance in the metabolic water is conserved in cellulose with a relatively constant isotope effect. The relationship of the 2H abundance between metabolic water and cellulose is more complex. Hydrogen incorporated into photosynthetic products during primary reduction steps is highly depleted in 2H. However, a large proportion of these hydrogens are subsequently replaced by exchange with water, leading to 2H enrichment during heterotrophic metabolism. Deciphering the oxygen isotope ratio of cellulose could help in providing insights into the carbon and oxygen fluxes exchanged between plants and the atmosphere. This is because the 18O abundance in cellulose records the 18O abundance in the metabolic water, which in turn, controls the oxygen isotopic signatures of the CO2 and O2 released by plants into the atmosphere. The hydrogen isotope effects associated with carbohydrate metabolism provide insights into the autotrophic state of a plant tissue. This is because the hydrogen isotope ratio of carbohydrates must reflect the net effects of the two opposing isotope effects associated with photosynthesis and heterotrophic metabolism.

Growth of cotton under continuous salinity stress: influence on allocation pattern, stomatal and non-stomatal components of photosynthesis and dissipation of excess light energy.

Abstract: Cotton (Gossypium hirsutum L.) plants were grown in flowing-culture solutions containing 0%, 26% and 55% natural seawater under controlled and otherwise identical conditions. Leaf Na(+) content rose to 360 mM in 55% seawater, yet the K(+) content was maintained above 100 mM. The K(+)/Na(+) selectivity ratio was much greater in the saline plants than in the control plants. All plants were healthy and able to complete the life cycle but relative growth rate fell by 46% in 26% seawater and by 83% in 55% seawater. Much of this reduction in growth was caused by a decreased allocation of carbon to leaf growth versus root growth. The ratio of leaf area/plant dry weight fell by 32% in 26% seawater and by 50% in 55 % seawater while the rate of carbon gain per unit leaf area fell by only 20% in 26% seawater and by as much as 66% in 55% seawater. Partial stomatal closure accounted for nearly all of the fall in the photosynthesis rate in 26% seawater but in 55% seawater much of the fall also can be attributed to non-stomatal factors. As a result of the greater effect of salinity on stomatal conductance than on CO2-uptake rate, photosynthetic water-use efficiency was markedly improved by salinity. This was also confirmed by stablecarbon-isotope analyses of leaf sugar and of leaf cellulose and starch. - Although non-stomatal photosynthetic capacity at the growth light was reduced by as much as 42% in 55% seawater, no effects were detected on the intrinsic photon yield of photosynthesis nor on the efficiency of photosystem II photochemistry, chlorophyll a/b ratio, carotenoid composition or the operation of the xanthophyll cycle. Whereas salinity caused in increase in mesophyll thickness and content of chloroplast pigments it caused a decrease in total leaf nitrogen content. The results indicate that the salinity-induced reduction in non-stomatal photosynthetic capacity was not caused by any detrimental effect on the photosynthetic apparatus but reflects a decreased allocation to enzymes of carbon fixation. - Rates of energy dissipation via CO2 fixation and photorespiration, calculated from gas-exchange measurements, were insufficient to balance the rate of light-energy absorption at the growth light. Salinity therefore would have been expected to cause the excess excitation energy to rise, leading to an increased nonradiative dissipation in the pigment bed and resulting increases in non-photochemical fluorescence quenching and zeaxanthin formation. However, no such changes could be detected, implying that salinity may have increased energy dissipation via a yet unidentified energy-consuming process. This lack of a response to salinity stress is in contrast to the responses elicited by short-term water stress which caused strong non-photochemical quenching and massive zeaxanthin formation.

An extremely low‐light adapted phototrophic sulfur bacterium from the Black Sea

Abstract: Five strains of a brown phototrophic sulfur bacterium (Chlorobium phaeobacteroides) were isolated from the chemocline of the Black Sea (80-m depth). All contain bacteriochlorophyll e as the main photosynthetic pigment. The strains revealed extreme low-light adaptation of growth compared to 12 other green and purple sulfur bacterial strains. At very low light intensities (<4 µEinst m‒2 s‒1), the Black Sea strain MN 1 oxidized sulfide faster than the type strain 2430; the latter reached three times higher oxidation rates at light saturation. Low-light adaptation is achieved by an increase of light-harvesting pigments (175% compared to the type strain) and a very low maintenance energy requirement. The efficiency of energy transfer (59%) within light-harvesting structures (chlorosomes) is comparable in other green sulfur bacteria and, therefore, appears to be limited by the molecular organization of the chlorosomes. From data in the literature, a light transmission of 0.0005% of surface irradiance was calculated for the chemocline of the Black Sea. Extrapolation of our laboratory data revealed that anoxygenic photosynthesis could account for 4% of total sulfide oxidation under average light conditions in situ and for 13% at maximal surface irradiance in summer.

Changes in Activities of Enzymes of Carbon Metabolism in Leaves during Exposure of Plants to Low Temperature.

Abstract: The aim of this study was to determine the response of photosynthetic carbon metabolism in spinach and bean to low temperature. (a) Exposure of warm-grown spinach and bean plants to 10°C for 10 days resulted in increases in the total activities of a number of enzymes, including ribulose 1,5-bisphosphate carboxylase (Rubisco), stromal fructose 1,6 bisphosphatase (Fru 1,6-P2ase), sedoheptulose 1,7-bisphosphatase (Sed 1,7-P2ase), and the cytosolic Fru 1,6-P2ase. In spinach, but not bean, there was an increase in the total activity of sucrose-phosphate synthase. (b) The CO2-saturated rates of photosynthesis for the cold-acclimated spinach plants were 68% greater at 10°C than those for warm-acclimated plants, whereas in bean, rates of photosynthesis at 10°C were very low after exposure to low temperature. (c) When spinach leaf discs were transferred from 27 to 10°C, the stromal Fru 1,6-P2ase and NADP-malate dehydrogenase were almost fully activated within 8 minutes, and Rubisco reached 90% of full activation within 15 minutes of transfer. An initial restriction of Calvin cycle fluxes was evident as an increase in the amounts of ribulose 1,5-bisphosphate, glycerate-3-phosphate, Fru 1,6-P2, and Sed 1,7-P2. In bean, activation of stromal Fru 1,6-P2ase was weak, whereas the activation state of Rubisco decreased during the first few minutes after transfer to low temperature. However, NADP-malate dehydrogenase became almost fully activated, showing that no loss of the capacity for reductive activation occurred. (d) Temperature compensation in spinach evidently involves increases in the capacities of a range of enzymes, achieved in the short term by an increase in activation state, whereas long-term acclimation is achieved by an increase in the maximum activities of enzymes. The inability of bean to activate fully certain Calvin cycle enzymes and sucrose-phosphate synthase, or to increase nonphotochemical quenching of chlorophyll fluorescence at 10°C, may be factors contributing to its poor performance at low temperature.

Iron-induced changes in light harvesting and photochemical energy conversion processes in eukaryotic marine algae.

Abstract: The role of iron in regulating light harvesting and photochemical energy conversion processes was examined in the marine unicellular chlorophyte Dunaliella tertiolecta and the marine diatom Phaeodactylum tricornutum. In both species, iron limitation led to a reduction in cellular chlorophyll concentrations, but an increase in the in vivo, chlorophyll-specific, optical absorption cross-sections. Moreover, the absorption cross-section of photosystem II, a measure of the photon target area of the traps, was higher in iron-limited cells and decreased rapidly following iron addition. Iron-limited cells exhibited reduced variable/maximum fluorescence ratios and a reduced fluorescence per unit absorption at all wave-lengths between 400 and 575 nm. Following iron addition, variable/maximum fluorescence ratios increased rapidly, reaching 90% of the maximum within 18 to 25 h. Thus, although more light was absorbed per unit of chlorophyll, iron limitation reduced the transfer efficiency of excitation energy in photosystem II. The half-time for the oxidation of primary electron acceptor of photosystem II, calculated from the kinetics of decay of variable maximum fluorescence, increased 2-fold under iron limitation. Quantitative analysis of western blots revealed that cytochrome f and subunit IV (the plastoquinone-docking protein) of the cytochrome b6/f complex were also significantly reduced by lack of iron; recovery from iron limitation was completely inhibited by either cycloheximide or chloramphenicol. The recovery of maximum photosynthetic energy conversion efficiency occurs in three stages: (a) a rapid (3-5 h) increase in electron transfer rates on the acceptor side of photosystem II correlated with de novo synthesis of the cytochrome b6/f complex; (b) an increase (10-15 h) in the quantum efficiency correlated with an increase in D1 accumulation; and (c) a slow (>18 h) increase in chlorophyll levels accompanied by an increase in the efficiency of energy transfer from the light-harvesting chlorophyll proteins to the reaction centers.

Decreased ribulose-1,5-bisphosphate carboxylase-oxygenase in transgenic tobacco transformed with ‘antisense’ rbcS

Abstract: The effect of nitrogen supply during growth on the contribution of ribulose-1,5-bisphosphate carboxylase-oxygenase (Rubisco; EC 4.1.1.39) to the control of photosynthesis was examined in tobacco (Nicotiana tabacum L.). Transgenic plants transformed with antisense rbcS to produce a series of plants with a progressive decrease in the amount of Rubisco were used to allow the calculation of the flux-control coefficient of Rubisco for photosynthesis (CR). Several points emerged from the data: (i) The strength of Rubisco control of photosynthesis, as measured by CR, was altered by changes in the short-term environmental conditions. Generally, CR was increased in conditions of increased irradiance or decreased CO2. (ii) The amount of Rubisco in wild-type plants was reduced as the nitrogen supply during growth was reduced and this was associated with an increase in CR. This implied that there was a specific reduction in the amount of Rubisco compared with other components of the photosynthetic machinery. (iii) Plants grown with low nitrogen and which had genetically reduced levels of Rubisco had a higher chlorophyll content and a lower chlorophyll a/b ratio than wild-type plants. This indicated that the nitrogen made available by genetically reducing the amount of Rubisco had been re-allocated to other cellular components including light-harvesting and electron-transport proteins. It is argued that there is a “luxury” additional investment of nitrogen into Rubisco in tobacco plants grown in high nitrogen, and that Rubisco can also be considered a nitrogen-store, all be it one where the opportunity cost of the nitrogen storage is higher than in a non-functional storage protein (i.e. it allows for a slightly higher water-use efficiency and for photosynthesis to respond to temporarily high irradiance).

Reduction of Ribulose-1,5-Bisphosphate Carboxylase/Oxygenase Content by Antisense RNA Reduces Photosynthesis in Transgenic Tobacco Plants

Abstract: A complementary DNA for the small subunit of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) was cloned from tobacco (Nicotiana tabacum) and fused in the antisense orientation to the cauliflower mosaic virus 35S promoter. This antisense gene was introduced into the tobacco genome, and the resulting transgenic plants were analyzed to assess the effect of the antisense RNA on Rubisco activity and photosynthesis. The mean content of extractable Rubisco activity from the leaves of 10 antisense plants was 18% of the mean level of activity of control plants. The soluble protein content of the leaves of anti-small subunit plants was reduced by the amount equivalent to the reduction in Rubisco. There was little change in phosphoribulokinase activity, electron transport, and chlorophyll content, indicating that the loss of Rubisco did not affect these other components of photosynthesis. However, there was a significant reduction in carbonic anhydrase activity. The rate of CO2 assimilation measured at 1000 micromoles quanta per square meter per second, 350 microbars CO2, and 25°C was reduced by 63% (mean value) in the antisense plants and was limited by Rubisco activity over a wide range of intercellular CO2 partial pressures (pi). In control leaves, Rubisco activity only limited the rate of CO2 assimilation below a pi of 400 microbars. Despite the decrease in photosynthesis, there was no reduction in stomatal conductance in the antisense plants, and the stomata still responded to changes in pi. The unchanged conductance and lower CO2 assimilation resulted in a higher pi, which was reflected in greater carbon isotope discrimination in the leaves of the antisense plants. These results suggest that stomatal function is independent of total leaf Rubisco activity.

The CO2 concentrating mechanism in cyanobacteria and microalgae

Abstract: Over the past 10 years it has become clear that cyanobacteria and microalgae possess mechanisms for actively acquiring inorganic carbon from the external medium and are able to use this to elevate the CO 2 concentration around the active site of the primary photosynthetic carboxylating enzyme, ribulose bisphosphate carboxylase-oxygenase (Rubisco)

Physical and chemical basis of carbon isotope fractionation in plants

Abstract: Naturally-occurring variations in the abundances of the stable isotopes of carbon and other elements can be used to understand the dynamics of natural processes in chemistry, biochemistry, biology, medicine, ecology and other fields. The use of carbon-13 isotopic abundances as an indicator of photosynthetic function in plants has become common. The purpose of this article is to describe the physical and chemical processes that contribute to the abundances of carbon-13 in plant materials, and to provide a framework for understanding how those processes control the isotopic contents of natural materials.

Distinctive Responses of Ribulose-1,5-Bisphosphate Carboxylase and Carbonic Anhydrase in Wheat Leaves to Nitrogen Nutrition and their Possible Relationships to CO2-Transfer Resistance

Abstract: The amounts of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), total chlorophyll (Chl), and total leaf nitrogen were measured in fully expanded, young leaves of wheat (Triticum aestivum L.), rice (Oryza sativa L.), spinach (Spinacia oleracea L.), bean (Phaseolus vulgaris L.), and pea (Pisum sativum L.). In addition, the activities of whole-chain electron transport and carbonic anhydrase were measured. All plants were grown hydroponically at different nitrogen concentrations. Although a greater than proportional increase in Rubisco content relative to leaf nitrogen content and Chl was found with increasing nitrogen supply for rice, spinach, bean, and pea, the ratio of Rubisco to total leaf nitrogen or Chl in wheat was essentially independent of nitrogen treatment. In addition, the ratio of Rubisco to electron transport activities remained constant only in wheat. Nevertheless, gas-exchange analysis showed that the in vivo balance between the capacities of Rubisco and electron transport in wheat, rice, and spinach remained almost constant, irrespective of nitrogen treatment. The in vitro carbonic anhydrase activity in wheat was very low and strongly responsive to increasing nitrogen content. Such a response was not found for the other C3 plants examined, which had 10- to 30-fold higher carbonic anhydrase activity than wheat at any leaf-nitrogen content. These distinctive responses of carbonic anhydrase activity in wheat were discussed in relation to CO2-transfer resistance and the in vivo balance between the capacities of Rubisco and electron transport.

Photosynthesis of oak trees [Quercus petraea (Matt.) Liebl.] during drought under field conditions: diurnal course of net CO2 assimilation and photochemical efficiency of photosystem II

Abstract: Adult trees of Quercus petraea were submitted to controlled water shortage in a natural stand near Nancy, France. Diurnal course of net CO2 assimilation rate (A) was measured in situ together with chlorophyll a fluorescence determined on dark adapted leaves. In 1990, trees experienced a strong water stress, with predawn and midday leaf water potentials below –2·0 and –3·0 MPa, respectively. Diurnal course of A of well-watered trees exhibited sometimes important midday decreases in A related to high temperature and vapour pressure deficit. Decreases in initial (Fo) and maximal (Fm) fluorescence and sometimes in photochemical efficiency of photosystem II (Fv/Fm) were observed and probably revealed the onset of mechanisms for thermal de-excitation. These mechanisms were shown to be sensitive to dithiothreitol. All these effects were reversible and vanished almost completely overnight. Therefore, they may be considered as protective mechanisms adjusting activity of photosystem II to the electron requirement for photosynthesis. Water stress amplified these reactions: A was strongly decreased, showing important midday depression; diurnal reductions in Fm and Fv/Fm were enhanced. The same trends were observed during summer 1991, despite a less marked drought. These protective mechanisms seemed very effective, as no photoinhibitory damage to PS II could be detected in either water stressed or control trees.

Photoinactivation of Catalase Occurs under Both High- and Low-Temperature Stress Conditions and Accompanies Photoinhibition of Photosystem II

Abstract: Severe photoinactivation of catalase (EC 1.11.1.6) and a decline of variable fluorescence (Fv), indicating photoinhibition of photosynthesis, were observed as rapid and specific symptoms in leaves exposed to a high heat-shock temperature of 40°C as well as in leaves exposed to low chilling temperatures in white light of only moderately high photosynthetic photon flux density of 520 μE m−2 s−1. Other parameters, such as peroxidase (EC 1.11.1.7), glycolate oxidase (EC 1.1.3.1), glutathione reductase (EC 1.6.4.2), or the chlorophyll content, were hardly affected under these conditions. At a compatible temperature of 22°C, the applied light intensity did not induce severe photoinactivations. In darkness, exposures to high or low temperatures did not affect catalase levels. Also, decline of Fv in light was not related to temperature sensitivity in darkness. The effective low-temperature ranges inducing photoinactivation of catalase differed significantly for chilling-tolerant and chilling-sensitive plants. In leaves of rye (Secale cereale L.) and pea (Pisum sativum L.), photoinactivation occurred only below 15°C, whereas inactivation occurred at 15°C in cucumber (Cucumis sativus L.) and maize (Zea mays L.). The behavior of Fv was similar, but the difference between chilling-sensitive and chilling-tolerant plants was less striking. Whereas the catalase polypeptide, although photoinactivated, was not cleaved at 0 to 4°C, the D1 protein of photosystem II was greatly degraded during the low-temperature treatment of rye leaves in light. Rye leaves did not exhibit symptoms of any major general photodamage, even when they were totally depleted of catalase after photoinactivation at 0 to 4°C, and catalase recovered rapidly at normal temperature. In cucumber leaves, the decline of catalase after exposures to bright light at 0 to 4°C was accompanied by bleaching of chlorophyll, and the recovery observed at 25°C was slow and required several days. Similar to the D1 protein of photosystem II, catalase differs greatly from other proteins by its inactivation and high turnover in light. Inasmuch as catalase and D1 protein levels depend on continuous repair synthesis, preferential and rapid declines are generally to be expected in light whenever translation is suppressed by stress actions, such as heat or chilling, and recovery will reflect the repair capacity of the plants.

The path of carbon in photosynthesis: improved crop yields with methanol.

Abstract: Foliar sprays of aqueous 10-50% methanol increased growth and development of C3 crop plants in arid environments. The effects of low levels (< 1 ml per plant) of methanol were observed for weeks after the brief time necessary for its rapid metabolism. Within several hours, foliar treatment with methanol resulted in increased turgidity. Plants treated with nutrient-supplemented methanol showed up to 100% increases in yields when maintained under direct sunlight in desert agriculture. In the shade and when winter crops were treated with methanol, plants showed no improvement of growth. When repeatedly treated with nutrient-supplemented methanol, shaded plants showed symptoms of toxicity. Repeated methanol treatments with glycine caused increased turgidity and stimulated plant growth without injury under indirect sunlight, but indoors with artificial illumination, foliar damage developed after 48 hr. Addition of glycerophosphate to glycine/methanol solutions allowed treatment of artificially illuminated plants indoors without injury. Plants with C4 metabolism showed no increase in productivity by methanol treatment. Plants given many applications of aqueous methanol showed symptoms of nutrient deficiency. Supplementation with a source of nitrogen sustained growth, eliminating symptoms of deficiency. Adjustment of carbon/nitrogen ratios was undertaken in the field by decreasing the source of nitrogen in the final application, resulting in early maturation; concomitantly, irrigation requirements were reduced.

Regulation of Photosynthetic Rate of Two Sunflower Hybrids under Water Stress

Abstract: The effect of short-term water stress on photosynthesis of two sunflower hybrids (Helianthus annuus L. cv Sungro-380 and cv SH-3622), differing in productivity under field conditions, was measured. The rate of CO2 assimilation of young, mature leaves of SH-3622 under well-watered conditions was approximately 30% greater than that of Sungro-380 in bright light and elevated CO2; the carboxylation efficiency was also larger. Growth at large photon flux increased assimilation rates of both hybrids. The changes in leaf composition, including cell numbers and sizes, chlorophyll content, and amounts of total soluble and ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) protein, and in Rubisco activity and amount of ribulose-1,5-bisphosphate (RuBP) were determined to assess the factors regulating the differences in assimilation of the hybrids at high and low water potentials. The amounts of chlorophyll, soluble protein, Rubisco protein and the initial activity of Rubisco and its activation state did not differ significantly between hybrids. However, unstressed leaves of SH-3622 had more, smaller cells per unit area and 60% more RuBP per unit leaf area than that of Sungro-380. Water stress developing over 4 days decreased the assimilation of both hybrids similarly. Changes in the amounts of chlorophyll, soluble and Rubisco protein, and Rubisco activity and activation state were small and were not sufficient to explain the decrease in photosynthesis; neither was decreased stomatal conductance (or stomatal “patchiness”). Reduction of photosynthesis per unit leaf area from 25 to 5 micromoles CO2 per square meter per second in both hybrids was caused by a decrease in the amount of RuBP from approximately 130 to 40 micromoles per square meter in SH-3622 and from 80 to 40 micromoles per square meter in Sungro. Differences between hybrids and their response to water stress is discussed in relation to control of RuBP regeneration.

Timing of ozone stress and resulting status of ribulose bisphosphate carboxylase/oxygenase and associated net photosynthesis

Abstract: summary Experiments were conducted to determine how the timing of an exposure to ozone influenced the impact of the gas on ribulose- 1,5-bisphosphate carboxylase/oxygenase (Rubisco) throughout the life span of a designated leaf. Saplings of Populus maximowizii×trichocarpa NE 388 received 5-d exposures to O3 in growth chambers during and at the termination of presumed synthesis of Rubisco in a designated leaf. Ozone had no detectable impact on Rubisco activity or quantity when the exposure occurred during the time of increasing concentration of the protein in the leaf. When the concentration of Rubisco was near its peak, O3 induced a reduction in quantity and activity of Rubisco, but after cessation of the O3 stress, levels converged with those of the untreated tissue. When O3exposure occurred after full leaf expansion, minimal effects of the gas could be detected. When plants of hybrid popular or Raphanus sativus L. cv. Cherry Belle received chronic O3 treatment throughout the lifespan of the leaf, Rubisco activity and quantity declined more rapidly and never converged with values of untreated tissue. Studies of gas exchange revealed that changes in Rubisco were associated with a decline in net photosynthesis (A) and that these effects preceded the observed reduction in foliar conductance. CO2 response curves were measured periodically, and the initial slope (linear portion) of the curve, reflecting carboxylation capacity, declined more rapidly with leaf age in O3-treated than in untreated poplars. There was no effect of O3 on stomatal limitation to CO2, assimilation except for a slight increase during the last 2 wk of the 9-wk experiment. This supported the hypothesis that O3 effects on A were associated with CO2-fixing capability of the leaf.

The CO2concentrating mechanism in cyanobactiria and microalgae

Abstract: Over the past 10 years it has become clear that cyanobacteria and microalgae possess mechanisms for actively acquiring inorganic carbon from the external medium and are able to use this to elevate the CO2 concentration around the active site of the primary photosynthetic carboxylating enzyme, ribulose bisphosphate carboxylase-oxygenase (Rubisco). This results in a vastly enhanced photosynthetic affinity for inorganic carbon (Ci) and improved photosynthetic efficiency. The CO2 concentrating mechanism is dependent on the existence of membrane bound Ci transport systems, and a microenvironment within the cell where the accumulated Ci can be used to elevate CO2 at the site of Rubisco. Evidence presented in this review suggests that in cyanobacteria this is achieved by the packaging of Rubisco and carbonic anhydrase (CA) into discrete structures, which are termed carboxysomes. Analogous structures in microalgae, termed pyrenoids, may perform a similar function. The recovery and analysis of high-CO2-requiring mutants has greatly advanced our understanding of the mechanisms and genes underlying these systems, especially in cyanobacteria, and this review places particular emphasis on the contribution made by molecular genetic approaches.

Diversity, Ecology, and Taxonomy of the Cyanobacteria

Abstract: The cyanobacteria are photosynthetic prokaryotes possessing the ability to synthesize chlorophyll a and at least one phycobilin pigment; typically water acts as the electron donor during photosynthesis, leading to the release of oxygen. They are by far the largest group of photosynthetic prokaryotes, as judged by their widespread occurrence, frequent abundance, and morphological diversity. Not only are they represented at the present day in most types of illuminated environment, except for those at lower pH values, but they have one of the longest geological records (Schopf and Walter, 1982). Much of the earth’s original atmospheric oxygen was probably formed by organisms quite similar to modern cyanobacteria (Knoll, 1985) and they are still responsible for a considerable proportion of photosynthetic oxygen evolution in the oceans.