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Showing papers on "Photosynthesis published in 1993"


Journal ArticleDOI
TL;DR: Various protective mechanisms and an efficient repair cycle of Photosystem II allow plants to survive light stress and probably allows for coordinated biodegradation and biosynthesis of the D1 protein.

2,223 citations


Journal ArticleDOI
TL;DR: During normally-encountered degrees of water deficit the capacity of the antioxidant systems and their ability to respond to increased active oxygen generation may be sufficient to prevent overt expression of damage.
Abstract: Water deficits cause a reduction in the rate of photosynthesis. Exposure to mild water deficits, when relative water content (RWC) remains above 70%, primarily causes limitation to carbon dioxide uptake because of stomatal closure. With greater water deficits, direct inhibition of photosynthesis occurs. In both cases limitation of carbon dioxide fixation results in exposure of chloroplasts to excess excitation energy. Much of this can be dissipated by various photoprotective mechanisms. These include dissipation as heat via carotenoids, photorespiration, CAM idling and, in some species, leaf movements and other morphological features which minimize light absorption. The active oxygen species superoxide and singlet oxygen are produced in chloroplasts by photoreduction of Oxygen and energy transfer from triplet excited chlorophyll to oxygen, respectively. Hydrogen peroxide and hydroxyl radicals can form as a result of the reactions of superoxide. All these species are reactive and potentially damaging, causing lipid peroxidation and inactivation of enzymes. They are normally scavenged by a range of antioxidants and enzymes which are present in the chloroplast and other subcellular compartments. When carbon dioxide fixation is limited by water deficit, the rate of active oxygen formation increases in chloroplasts as excess excitation energy, not dissipated fay the photoprotective mechanisms, is used to form superoxide and singlet oxygen. However, photorespiratory hydrogen peroxide production in peroxisomes decreases. Increased superoxide can be detected by EPR (electron paramagnetic resonance) in chloroplasts from droughted plants. Stiperoxide formation leads to changes suggestive of oxidative damage including lipid peroxidation and a decrease in ascorbate. These changes are not, however, apparent until severe water deficits develop, and they could also be interpreted as secondary effects of water deficit-induced senescence or wounding. Non-lethal water deficits often result in increased activity of superoxide dismutase, glutathione reductase and monodehydroascorbate reductase. Increased capacity of these protective enzymes may be part of a general antioxidative response in plants involving regulation of protein synthesis or gene expression. Since the capacity of these enzymes is also increased by other treatments which cause oxidative damage, and which alter the balance between excitation energy input and carbon dioxide fixation such as low temperature and high irradiance, it is suggested that water deficit has the same effect. Light levels that are not normally excessive do become excessive and photoprotective/antioxidative systems are activated. Some of the photoprotective mechanisms themselves could result in active oxygen formation. Photoinhibitory damage also includes a component of oxidative damage. During normally-encountered degrees of water deficit the capacity of the antioxidant systems and their ability to respond to increased active oxygen generation may be sufficient to prevent overt expression of damage. Desiccation-tolerant tissues such as bryophytes, lichens, spores, seeds, some algae and a few vascular plant leaves can survive desiccation to below 30-40% RWC, A component of desiccation damage in seeds and bacteria is oxygen-dependent. Desiccation causes oxidation of glutathione, a major antioxidant, and appearance of a free radical signal detected by EPR in a number of tissues suggesting that oxidative damage has occurred. In photosynthetic cells damage may arise from photooxidation. Disruption of membrane-bound electron tranport systems in partially hydrated tissue could lead to reduction of oxygen to superoxide. Oxidation of lipids and sulphydryl groups may also occur in dry tissue. Tolerant cells recover upon rehydration and arc able to reduce their glutathione pool. Non-tolerant species go on to show further oxidative damage including lipid peroxidation. It is difficult to attribute this subsequent damage to the cause or effect of death. Embryos in seeds lose desiccation tolerance soon after imbibition. This is associated with membrane damage that has been attributed to superoxide-mediated deesterification of phospholipids and loss of lipophilic antioxidants. These effects are discussed in relation to other mechanisms involved in desiccation tolerance. Contents Summary 27 I. Introduction 28 II. Generation of active oxygen and defence mechanisms in plant cells 29 III. The effect of water deficit on photosynthesis 31 IV. Mechanisms for active oxygen generation during water deficit 36 V. Evidence for oxidative damage during water deficit 39 VI. Desiccation 47 VII. Conclusions 52 Acknowledgements 53 References 53.

2,008 citations


Journal ArticleDOI
TL;DR: The results obtained demonstrated the existence of a general positive, linear relationship between plant decomposition rates and nitrogen and phosphorus concentrations, and reflect the coupling of phosphorus and nitrogen in the basic biochemical processes of both plants and their microbial decomposers.
Abstract: The strength and generality of the relationship between decomposition rates and detritus carbon, nitrogen, and phosphorus concentrations was assessed by comparing published reports of decomposition rates of detritus of photosynthetic organisms, from unicellular algae to trees. The results obtained demonstrated the existence of a general positive, linear relationship between plant decomposition rates and nitrogen and phosphorus concentrations. Differences in the carbon, nitrogen, and phosphorus concentrations of plant detritus accounted for 89% of the variance in plant decomposition rates of detritus orginating from photosynthetic organisms ranging from unicellular microalgae to trees. The results also demonstrate that moist plant material decomposes substantially faster than dry material with similar nutrient concentrations. Consideration of lignin, instead of carbon, concentrations did not improve the relationships obtained. These results reflect the coupling of phosphorus and nitrogen in the basic biochemical processes of both plants and their microbial decomposers, and stress the importance of this coupling for carbon and nutrient flow in ecosystems.

872 citations


Journal ArticleDOI
TL;DR: C4 and CAM photosynthesis are evolutionarily derived from C3 photosynthesis, with a tendency toward ecological adaptation of C4 plants into warm, monsoonal climates and CAM plants into water-limited habitats and in an anthropogenically altered CQ2 environment, C 4 plants may lose their competitive advantage over C3 plants.
Abstract: C4 and CAM photosynthesis are evolutionarily derived from C3 photosynthesis. The morphological and biochemical modifications necessary to achieve either C4 or CAM photosynthesis are thought to have independently arisen numerous times within different higher plant taxa. It is thought that C4 photosynthesis evolved in response to the low atmospheric CO2 concentrations that arose sometime after the end of the Cretaceous. Low CO2 concentrations result in significant increases in photorespiration of C3 plants, reducing productivity; both C3-C4 intermediate and C4 plants exhibit reduced photorespiration rates. In contrast, it may be argued that CAM arose either in response to selection of increased water-use efficiency or for increased carbon gain. Globally, all three pathways are widely distributed today, with a tendency toward ecological adaptation of C4 plants into warm, monsoonal climates and CAM plants into water-limited habitats. In an anthropogenically altered CQ2 environment, C4 plants may lose their competitive advantage over C3 plants. 411

758 citations


Journal ArticleDOI
28 Jan 1993-Nature
TL;DR: It is proposed that the global expansion of C4 biomass may be related to lower atmospheric carbon dioxide levels because C4 photosynthesis is favoured over C3 photosynthesis when there are low concentrations of carbon dioxide in the atmosphere.
Abstract: THE most common and the most primitive pathway of the three different photosynthetic pathways used by plants is the C3 pathway, or Calvin cycle, which is characterized by an initial CO2 carboxylation to form phosphoglyceric acid, a 3-carbon acid. The carbon isotope composition (δ13C) of C3 plants varies from about −23 to −35%l–3 and averages about −26%. Virtually all trees, most shrubs, herbs and forbs, and cool-season grasses and sedges use the C3 pathway. In the C4 pathway (Hatch–Slack cycle), CO2 initially combines with phosphoenol pyruvate to form the 4-carbon acids malate or aspartic acid, which are translocated to bundle sheath cells where CO2 is released and used in Calvin cycle reactions1–4. The carbon isotope composition of C4 plants ranges from about −10 to −14%, averaging about −13% for modern plants1–3. Warm-season grasses and sedges are the most abundant C4 plants, although C4 photosynthesis is found in about twenty families5. The third photosynthetic pathway, CAM, combines features of both C3 and C4 pathways. CAM plants, which include many succulents, have intermediate carbon isotope compositions and are also adapted to conditions of water and CO2 stress. The modern global ecosystem has a significant component of C4 plants, primarily in tropical savannas, temperate grasslands and semi-desert scrublands. Studies of palaeovegetation from palaeosols and palaeodiet from fossil tooth enamel indicate a rapid expansion of C4 biomass in both the Old World and the New World starting 7 to 5 million years ago. We propose that the global expansion of C4 biomass may be related to lower atmospheric carbon dioxide levels because C4 photosynthesis is favoured over C3 photosynthesis when there are low concentrations of carbon dioxide in the atmosphere.

654 citations


Journal ArticleDOI
TL;DR: In this paper, a pump-and-probe fluorescence technique was used to measure the change in the quantum yield of fluorescence induced by the strong pump flash, which was then used to derive the absolute absorption cross sections for photosystem 2 and the maximum rate of photosynthetic electron transport at light saturation.
Abstract: We describe the theory and practice of estimating photosynthetic rates from light-stimulated changes in the quantum yield of chlorophyll fluorescence. By means of a pump-and-probe fluorescence technique, where weak probe flashes are used to measure the change in the quantum yield of fluorescence induced by the strong pump flash, it is possible to derive the absolute absorption cross sections for photosystem 2, the quantum yield for photochemistry, and the maximum rate of photosynthetic electron transport at light saturation. In conjunction with a semiempirical biophysical model of photosynthesis, these parameters can bc used to calculate the instantaneous rate of gross photosynthesis in situ under ambient irradiance. A profiling pump-and-probe fluorometer was constructed and interfaced with a CTD, and vertical profiles of variable fluorescence were obtained on four cruises in the northwest Atlantic Ocean. The derived photosynthetic rates were compared with concurrent estimates of production based on radiocarbon uptake. The correlation coefficient between the two estimates of primary production, normalized to Chl a, was 0.86; linear regression analysis yielded a slope of 1.06. There is a 3-4-fold range in the maximum change in the quantum yields of photochemistry and absorption cross-sections in natural phytoplankton communities. Uncertainties in the pump-and-probe-derived estimates of photosynthesis arc primarily due to temporal mismatches between instantaneous and time-integrated measures of production and in biological variability in the ratio of the number of PS2 reaction centers to total Chl a. Almost all measurements of phytoplankton photosynthesis in situ are based on the timedependent incorporation of radiocarbon into particulate matter or on changes in concentration of dissolved oxygen in the bulk fluid.

611 citations


Journal ArticleDOI
TL;DR: The phycobilisome, the light-harvesting apparatus of cyanobacteria and red algae, is discussed, which allows these organisms to efficiently utilize available light energy to drive photosynthetic electron transport and CO2 fixation.

592 citations


Journal ArticleDOI
TL;DR: It is demonstrated that SOD is a critical component of the active-oxygen-scavenging system of plant chloroplasts and indicate that modification of SOD expression in transgenic plants can improve plant stress tolerance.
Abstract: Transgenic tobacco plants that express a chimeric gene that encodes chloroplast-localized Cu/Zn superoxide dismutase (SOD) from pea have been developed. To investigate whether increased expression of chloroplast-targeted SOD could alter the resistance of photosynthesis to environmental stress, these plants were subjected to chilling temperatures and moderate (500 mumol of quanta per m2 per s) or high (1500 mumol of quanta per m2 per s) light intensity. During exposure to moderate stress, transgenic SOD plants retained rates of photosynthesis approximately 20% higher than untransformed tobacco plants, implicating active oxygen species in the reduction of photosynthesis during chilling. Unlike untransformed plants, transgenic SOD plants were capable of maintaining nearly 90% of their photosynthetic capacity (determined by their photosynthetic rates at 25 degrees C) following exposure to chilling at high light intensity for 4 hr. These plants also showed reduced levels of light-mediated cellular damage from the superoxide-generating herbicide methyl viologen. These results demonstrate that SOD is a critical component of the active-oxygen-scavenging system of plant chloroplasts and indicate that modification of SOD expression in transgenic plants can improve plant stress tolerance.

555 citations


Journal ArticleDOI
TL;DR: It is concluded that the photosynthetic response to low temperatures and susceptibility to photoinhibition are consequences of the overwintering strategy of the plant species.
Abstract: Cold acclimation requires adjustment to a combination of light and low temperature, conditions which are potentially photoinhibitory. The photosynthetic response of plants to low temperature is dependent upon time of exposure and the developmental history of the leaves. Exposure of fully expanded leaves of winter cereals to short-term, low temperature shiftsinhibits whereas low temperature growthstimulates electron transport capacity and carbon assimilation. However, the photosynthetic response to low temperature is clearly species and cultivar dependent. Winter annuals and algae which actively grow and develop at low temperature and moderate irradiance acquire a resistance to irradiance 5- to 6-fold higher than their growth irradiance. Resistance to short-term photoinhibition (hours) in winter cereals is a reflection of the increased capacity to keep QA oxidized under high light conditions and low temperature. This is due to an increased capacity for photosynthesis. These characteristics reflect photosynthetic acclimation to low growth temperature and can be used to predict the freezing tolerance of cereals. It is proposed that the enhanced photosynthetic capacity reflects an increased flux of fixed carbon through to sucrose in source tissue as a consequence of the combined effects of increased storage of carbohydrate as fructans in the vacuole of leaf mesophyll cells and an enhanced export to the crown due to its increased sink activity. Long-term exposure (months) of cereals to low temperature photoinhibition indicates that this reduction of photochemical efficiency of PS II represents a stable, long-term down regulation of PS II to match the energy requirements for CO2 fixation. Thus, photoinhibition in vivo should be viewed as the capacity of plants to adjust photosynthetically to the prevailing environmental conditions rather than a process which necessarily results in damage or injury to plants. Not all cold tolerant, herbaceous annuals use the same mechanism to acquire resistance to photoinhibition. In contrast to annuals and algae, overwintering evergreens become dormant during the cold hardening period and generally remain susceptible to photoinhibition. It is concluded that the photosynthetic response to low temperatures and susceptibility to photoinhibition are consequences of the overwintering strategy of the plant species.

493 citations


Journal ArticleDOI
TL;DR: This overview of the current knowledge on the molecular aspects of the biology of cyanobacteria is presented, as well as on their mechanisms of resistance to metal ions and their responses to metabolic stress.
Abstract: Dating from the Pre-Cambrian era, cyanobacteria have a long history of adaptation to the Earth's environment. By evolving oxygen via photosynthetic reactions similar to those of plants and green algae, these prokaryotes were essential to the evolution of the present biosphere. They continue to make a large contribution to the equilibrium of the Earth's atmosphere by production oxygen and removing carbon dioxide. To survive in extreme or variable environments, cyanobacteria have developed specific regulatory systems, in addition to more general mechanisms equivalent to those of other prokaryotes or photosynthesis eukaryotes. Specific regulatory systems control the differentiation of specialized nitrogen-fixing cells and of cell types facilitating the dispersion of species. In the past decade, considerable progress has been made towards understanding the expression of the cyanobacterial genome in response to variations in the intensity and spectral quality of incident light and in response to nutritional conditions, especially carbon, nitrogen and sulphur sources. These studies have provided insights into the relationships between carbon and nitrogen intermediary metabolism, and a start towards understanding of the interconnected pathways which lead from the perception of environmental signals to the regulation of enzyme activities and gene expression. Cyanobacterial regulatory mechanisms share common features with those of other prokaryotes, but are unique since these essentially photo-autotrophic organisms must maintain a proper cellular C/N balance, in spite of dailty variations in incident light. Thus an appropriate coordination between photosynthesis and other metabolic processes must be achieved through control of the catalytic activity of key enzymes by reducing equivalents and ATP produced by photosynthetic or respiratory electron transport. Recently discovered kinases/phosphatases act by post-translational modification of specific proteins which probably act as signal transducers or modulators of gene expression in a manner similar to the well-known two-component regulatory systems described in other bacteria. In this overview, we present our current knowledge on the molecular aspects of the biology of cyanobacteria, as well as on their mechanisms of resistance to metal ions and their responses to metabolic stress.

434 citations


Journal ArticleDOI
TL;DR: Although cell chlorophyll a (chl a) content decreased in nutrient‐starved cells, the ratios of light‐harvesting accessory pigments ( chl c and fucoxanthin) to chl a were unaffected by nutrient starvation, indicating that chlorosis mirrored a general reduction in cell protein content.
Abstract: The effects of nitrate, phosphate, and iron starvation and resupply on photosynthetic pigments, selected photosynthetic proteins, and photosystem II (PSII) photochemistry were examined in the diatom Phaeodactylum tricornutum Bohlin (CCMP 1327). Although cell chlorophyll a (chl a) content decreased in nutrient-starved cells, the ratios of light-harvesting accessory pigments (chl c and fucoxanthin) to chl a were unaffected by nutrient starvation. The chl a-specific light absorpition coefficient (a*) and the functional absorption cross-section of PSII (σ) increased during nutrient starvation, consistent with reduction of intracellular self-shading (i.e. a reduction of the “package effect”) as cells became chlorotic. The light-harvesting complex proteins remained a constant proportion of total cell protein during nutrient starvation, indicating that chlorosis mirrored a general reduction in cell protein content. The ratio of the xanthophylls cycle pigments diatoxanthin and diadinoxanthin to chl a increased during nutrient starvation. These pigments are thought to play a photo-protective role by increasing dissipation of excitation energy in the pigment bed upstream from the reaction centers. Despite the increase in diatoxanthin and diadinoxanthin, the efficiency of PSII photochemistry, as measured by the ration of variable to maximum fluorescence (Fv/Fm) of dark-adapted cells, declined markedly under nitrate and iron starvation and moderately under phosphate starvation. Parallel to changes in Fv/Fm were decreases in abundance of the reaction center protein D1 consistent with damage of PSII reaction centers in nutrient-starved cells. The relative abundance of the carboxylating enzyme, ribulose bisphosphate carboxylase/oxygenase (RUBISCO), decreased in response to nitrate and iron starvation but not phosphate starvation. Most marked was the decline in the abundance of the small subunit of RUBISCO in nitrate-starved cells. The changes in pigment content and fluorescence characteristics were typically reversed within 24 h of resupply of the limiting nutrient.

Journal ArticleDOI
TL;DR: It is suggested that both carotenoids and flavonoids may be involved in plant UV-B photoprotection, but only carotanoids are directly linked to photoprotsection of photosynthetic function.
Abstract: The increase in ultraviolet-B (UV-B; 0.290–0.320 [mu]m) radiation received by plants due to stratospheric ozone depletion heightens the importance of understanding UV-B tolerance. Photosynthetic tissue is believed to be protected from UV-B radiation by UV-B-absorbing compounds (e.g. flavonoids). Although synthesis of flavonoids is induced by UV-B radiation, its protective role on photosynthetic pigments has not been clearly demonstrated. This results in part from the design of UV-B experiments in which experimental UV-A irradiance has not been carefully controlled, since blue/UV-A radiation is involved in the biosynthesis of the photosynthetic pigments. The relationship of flavonoids to photosynthetic performance, photosynthetic pigments, and growth measures was examined in an experiment where UV-A control groups were included at two biologically effective daily UV-B irradiances, 14.1 and 10.7 kJ m-2. Normal, chlorophyll-deficient, and flavonoid-deficient pigment isolines of two soybean (Glycine max) cultivars that produced different flavonol glycosides (Harosoy produced kaempferol, Clark produced quercetin and kaempferol) were examined. Plants with higher levels of total flavonoids, not specific flavonol glycosides, were more UV-B tolerant as determined by growth, pigment, and gas-exchange variables. Regression analyses indicated no direct relationship between photosynthesis and leaf levels of UV-B-absorbing compounds. UV-B radiation increased photosynthetic pigment content, along with UV-B-absorbing compounds, but only the former (especially carotenoids) was related to total biomass (r2 = 0.61, linear) and to photosynthetic efficiency (negative, exponential relationship, r2 = 0.82). A reduction in photosynthesis was associated primarily with a stomatal limitation rather than photosystem II damage. This study suggests that both carotenoids and flavonoids may be involved in plant UV-B photoprotection, but only carotenoids are directly linked to photoprotection of photosynthetic function. These results additionally show the importance of UV-A control in UV-B experiments conducted using artificial lamps and filters.

Journal ArticleDOI
01 Jan 1993-Nature
TL;DR: In this article, the authors present data indicating that the increase in CO2 has enhanced biospheric carbon fixation and altered species abundances by increasing the water-use efficiency of biomass production of C3 plants.
Abstract: ATMOSPHERIC CO2 concentration was 160 to 200 μmol mol−1 during the Last Glacial Maximum (LGM; about 18,000 years ago)1, rose to about 275 (μmol mol−1 10,000 years ago2,3, and has increased to about 350 μmol mol−1 since 1800 (ref. 4). Here we present data indicating that this increase in CO2 has enhanced biospheric carbon fixation and altered species abundances by increasing the water-use efficiency of biomass production of C3 plants, the bulk of the Earth's vegetation. We grew oats (Avena sativa), wild mustard (Brassica kaber) and wheat (Triticum aes-tivum cv. Seri M82 and Yaqui 54), all C3 annuals, and selected C4 grasses along daytime gradients of Glacial to present atmospheric CO2 concentrations in a 38-m-long chamber. We calculated parameters related to leaf photosynthesis and water-use efficiency from stable carbon isotope ratios (13C/12C) of whole leaves. Leaf water-use efficiency and above-ground biomass/plant of C3 species increased linearly and nearly proportionally with increasing CO2 concentrations. Direct effects of increasing CO2 on plants must be considered when modelling the global carbon cycle and effects of climate change on vegetation.

Journal ArticleDOI
TL;DR: The effects of long-term CO2 enhancement and varying nutrient availability on photosynthesis and ribulose-1,5-bisphosphate carboxylase/oxygenase (rubisco) were studied on loblolly pine seedlings and demonstrated acclimation of photosynthetic processes to elevated CO2 through reallocation of N.
Abstract: The effects of long-term CO2 enhancement and varying nutrient availability on photosynthesis and ribulose-1,5-bisphosphate carboxylase/oxygenase (rubisco) were studied on loblolly pine (Pinus taeda L.) seedlings grown in two atmospheric CO2 partial pressures (35 and 65 Pa) and three nutrient treatments (low N, low P, and high N and P). Measurements taken in late autumn (November) after 2 years of CO2 enrichment and nutrient addition showed that photosynthetic rates were higher for plants grown at elevated CO2 only when they received supplemental N. Total rubisco activity and rubisco content decreased at elevated CO2, but there was an increase in activation state. At elevated CO2, proportionately less N was found in rubisco and more N was found in the light reaction components. These results demonstrate acclimation of photosynthetic processes to elevated CO2 through reallocation of N. Loblolly pine grown in nutrient conditions similar to native soils (low N availability) had lower needle N and chlorophyll content, lower total rubisco activity and content, and lower photosynthetic rates than plants grown at high N and P. This suggests that the magnitude of the photosynthetic response to a future, high-CO2 environment will be dependent on soil fertility in the system.


Book ChapterDOI
01 Jan 1993
TL;DR: This chapter explains the variability of carbon isotope fractionation during photosynthesis, and presents an example of this bimodal distribution for 351 species of the grass family.
Abstract: Publisher Summary This chapter explains the variability of carbon isotope fractionation during photosynthesis. Early surveys showed that the carbon isotope ratios of C3 and C4 plants fall into two nonoverlapping categories. The chapter presents an example of this bimodal distribution for 351 species of the grass family. While the isotope ratios of the two groups of plants are related to function and to structure, the variation within each category is because of the influence of environmental factors on the kinetics of photosynthesis. Isotope ratio measurements can thus be used for interpreting the relative magnitudes of competing processes that take place during metabolism. As far as the carbon isotope composition is concerned, the succulents that have the ability to utilize the crassulacean acid metabolism constitute a separate category. These plants do not specifically differ from C3 plants in their structure, but they have the ability to fix CO2 as malate along the C4 pathway at night.

Journal ArticleDOI
TL;DR: Analysis of the characteristics of chlorophyll fluorescence induction in sub- and super-saturating light revealed two targets of mild heat stress: an irreversible inhibition of electron donation to PSII and a reversible reduction of excitation energy trapping by the PSII reaction centers, with the former effect being identified as the major determinant of the loss of photosynthesis.

Journal ArticleDOI
TL;DR: Six day old rice seedlings were grown in a nutrient solution with either Cd or Ni and the effect of Cd on carbohydrate distribution and content was similar to that of Ni, and the possible mechanisms involved in the abnormal carbohydrate accumulation and distribution are discussed.
Abstract: Six day old rice seedlings (Oryza sativa L. cv. Bahia) were grown for 5 or 10 days in a nutrient solution with either Cd (0.01, 0.1 mmol/l) or Ni (0.1, 0.5 mmol/l). Both Cd and Ni reduced the length of shoots and roots depending on the concentration and type of ion tested. On the other hand, the dry weight to fresh weight ratio was increased by heavy metal treatments, especially in the aerial part of 0.5 mmol/l Ni treated plants. The application of 0.1 mmol/l Cd and 0.5 mmol/l Ni to the seedlings produced an inhibition of the transport of carbohydrate reserves from the seeds from which plants were developing, to the rest of the plant. Net photosynthesis was also inhibited in treated plants. However, the total carbohydrate content in the shoots of these plants was higher than in controls. Thus, the starch, soluble sugars and sucrose content in the shoots of 0.5 mmol/l Ni treated plants was respectively up to 2.6, 2.8 and 4 times greater compared to controls. The distribution of assimilates between organs was also affected by the treatment: the carbohydrate content increased in the stem and second leaf but it was not affected or decreased in the root and third leaf. Although less evident, the effect of Cd on carbohydrate distribution and content was similar to that of Ni. The possible mechanisms involved in the abnormal carbohydrate accumulation and distribution are discussed.

Journal ArticleDOI
TL;DR: In this article, the functioning of isolated spinach (Spinacia oleracea L.) leaf mitochondria has been studied in the presence of metabolite concentrations similar to those that occur in the cytosol in vivo.
Abstract: The functioning of isolated spinach (Spinacia oleracea L.) leaf mitochondria has been studied in the presence of metabolite concentrations similar to those that occur in the cytosol in vivo. From measurements of the concentration dependence of the oxidation of the main substrates, glycine and malate, we have concluded that the state 3 oxidation rate of these substrates in vivo is less than half of the maximal rates due to substrate limitation. Analogously, we conclude that under steady-state conditions of photosynthesis, the oxidation of cytosolic NADH by the mitochondria does not contribute to mitochondrial respiration. Measurements of mitochondrial respiration with glycine and malate as substrates and in the presence of a defined malate:oxaloacetate ratio indicated that about 25% of the NADH formed in vivo during the oxidation of these metabolites inside the mitochondria is oxidized by a malate-oxaloacetate shuttle to serve extramitochondrial processes, e.g. reduction of nitrate in the cytosol or of hydroxypyruvate in the peroxisomes. The analysis of the products of the oxidation of malate indicates that in the steady state of photosynthesis the activity of the tricarboxylic acid cycle is very low. Therefore, we have concluded that the mitochondrial oxidation of malate in illuminated leaves produces mainly citrate, which is converted via cytosolic aconitase and NADP-isocitrate dehydrogenase to yield 2-oxoglutarate as the precursor for the formation of glutamate and glutamine, which are the main products of photosynthetic nitrate assimilation.

Journal ArticleDOI
TL;DR: It is postulate that, during short term exposure of plants to cadmium in the early stages of growth, the Calvin cycle reactions are more likely than photosystem II to be the primary target of the toxic influence of cadMium.
Abstract: Bean plants (Phaseolus vulgaris L. cv. Scarlett), germinated in darkness for I week, were transferred to light (200 μmol m-2 s-1 ) and cultivated for I week in a complete nutrient solution. After this period, cadmium ions in the form of CdSO4 were added at the concentrations of 0.10.20 and 50 μM. The effects of this metal on the properties of photosystem II photochemistry were studied by means of modulated fluorescence analysis. Steady state photochemical quenching. non-photochemical quenching and terminal fluorescence were determined in control and cadmiumtreated plants. We postulate that, during short term exposure of plants to cadmium in the early stages of growth, the Calvin cycle reactions are more likely than photosystem II to be the primary target of the toxic influence of cadmium. The reduced demand for ATP and NADPH upon Calvin cycle inhibition causes a down-regulation of photosystem II photochemistry and of the yield of linear electron transport.

Journal ArticleDOI
TL;DR: Although acclimation to low irradiance reduced the photosynthetic capacity per unit nitrogen by 12%, the considerable reorganisation of proteins within the thylakoids increased potential daily photosynthesis by 20% over that which would have been gained by a 'sun' leaf.
Abstract: Nitrogen redistribution between and within leaves was examined in a plot of lucerne (Medicago sativa L. cv. Aurora) in relation to potential canopy photosynthesis. The canopy was sampled during regrowth after cutting and just prior to flowering. As leaves were progressively shaded by the newly produced leaves, nitrogen content fell and photosynthetic acclimation occurred. The rate of acclimation in the canopy was the same as occurred following a step change to 23 or 6% sunlight. The profile of leaf nitrogen content was stable with respect to leaf area index and independent of time of sampling. Optimal profiles of nitrogen distribution between leaves, photosynthetic rate per unit chlorophyll and nitrogen partitioning within leaves were calculated from the relationships between photosynthesis and nitrogen in conjunction with the light environment of the preceding 3 days. The optimal nitrogen content of the leaves should vary in proportion to the relative daily irradiance at each leaf. The observed distribution achieved 88% of the potential daily photosynthesis, while a uniform nitrogen distribution yielded only 80%. Photosynthetic acclimation and nitrogen partitioning within each leaf both responded to daily irradiance similarly to the calculated optimum except at the two extremes. At the top of the canopy, photosynthetic rate per unit of chlorophyll did not increase as much as the calculated optimum, while at the base of the canopy, nitrogen partitioning failed to fall as much as the calculated optimum. This may reflect the constraints on the flexibility of the photosynthetic system. Nitrogen redistribution between leaves made a dramatic contribution to increasing the potential photosynthesis by the canopy. Although acclimation to low irradiance reduced the photosynthetic capacity per unit nitrogen by 12%, the considerable reorganisation of proteins within the thylakoids increased potential daily photosynthesis by 20% over that which would have been gained by a 'sun' leaf. However, in terms of canopy photosynthesis, which is dominated by leaves intercepting most of the light, acclimation contributed only a few per cent to the potential daily canopy photosynthesis.

Journal ArticleDOI
TL;DR: Intertidal benthic algae were the least sensitive to UV-B irradiation and this may be related to adaptation, through the accumulation ofUV-B screening compounds, to high light/high UV- B levels.
Abstract: Several species of marine benthic algae, four species of phytoplankton and two species of seagrass have been subjected to ultraviolet B irradiation for varying lengths of time and the effects on respiration, photosynthesis and fluorescence rise kinetics studied. No effect on respiration was found. Photosynthesis was inhibited to a variable degree in all groups of plants after irradiation over periods of up to 1 h and variable fluorescence was also inhibited in a similar way. The most sensitive plants were phytoplankton and deep-water benthic algae. Intertidal benthic algae were the least sensitive to UV-B irradiation and this may be related to adaptation, through the accumulation of UV-B screening compounds, to high light/high UV-B levels. Inhibition of variable fluorescence (Fv) of the fluorescence rise curve was a fast and sensitive indicator of UV-B damage. Two plants studied, a brown alga and a seagrass, showed very poor recovery of Fv over a period of 32 h.

Journal ArticleDOI
TL;DR: Based on a comparison of the response of many species to different irradiances during growth, it is generally the case that the proportion of thylakoid nitrogen increases for leaves grown under lower irradiance so as to maintain a constant ratio of photosynthetic capacity to total leaf nitrogen.
Abstract: Acclimation by the photosynthetic system to the gradient in irradiance through a leaf canopy was investigated with a plot of lucerne (Medicago sativa L. cv. Aurora). The aims were to determine the extent to which acclimation occurred in a natural canopy and to quantify the changes in the partitioning of nitrogen within the leaf that are associated with acclimation. The canopy grew up around light sensors placed at 10 cm height increments which logged the irradiance at 1 min intervals for the 4 days that preceded sampling. Photosynthetic capacity was measured with leaf disc oxygen electrodes and the chlorophyll, soluble protein and nitrogen contents of the leaves were determined. Daily irradiance declined exponentially down through the canopy. Nitrogen content and photosynthetic capacity both declined down through the canopy. Photosynthetic acclimation by the lower leaves was evident from the lower chlorophyll a/b ratios and reduced photosynthetic capacity per unit chlorophyll. The lower photosynthetic capacity per unit of chlorophyll was offset by an increased proportion of leaf nitrogen present in the thylakoids. Consequently, the photosynthetic capacity per unit leaf nitrogen was nearly independent of irradiance. Based on a comparison of the response of many species to different irradiances during growth, it is generally the case that the proportion of thylakoid nitrogen increases for leaves grown under lower irradiance so as to maintain a constant ratio of photosynthetic capacity to total leaf nitrogen. However, the ratio of photosynthetic capacity to total leaf nitrogen varies widely between species.

Journal ArticleDOI
TL;DR: For C 3 plants, higher CO 2 levels favour the photosynthetic carbon reduction cycle over the photorespiratory cycle, resulting in higher rates of carbohydrate production and plant productivity as discussed by the authors.
Abstract: The atmospheric CO 2 concentration has risen from the preindustrial level of approximately 290 μl l -1 to more than 350 μl l -1 in 1993. The current rate of rise is such that concentrations of 420 μl l -1 are expected in the next 20 years. For C 3 plants, higher CO 2 levels favour the photosynthetic carbon reduction cycle over the photorespiratory cycle, resulting in higher rates of carbohydrate production and plant productivity. The change in balance between the two photosynthetic cycles appears to alter nitrogen and carbon metabolism in the leaf, possibly causing decreases in nitrogen concentrations in the leaf


Journal ArticleDOI
TL;DR: It is concluded that the level of SPS in the leaves plays a pivotal role in carbon partitioning and high SPS levels have the potential to boost photosynthetic rates under favorable conditions.
Abstract: The expression of a sucrose-phosphate synthase (SPS) gene from maize (Zea mays, a monocotyledon) in tomato (Lycopersicon esculentum, a dicotyledon) resulted in marked increases in extractable SPS activity in the light and the dark. Diurnal modulation of the native tomato SPS activity was found. However, when the maize enzyme was present the tomato leaf cells were unable to regulate its activation state. No detrimental effects were observed and total dry matter production was unchanged. However, carbon allocation within the plants was modified such that in shoots it increased, whereas in roots it decreased. There was, therefore, a change in the shoot:root dry weight ratio favoring the shoot. This was positively correlated with increased SPS activity in leaves. SPS was a major determinant of the amount of starch in leaves as well as sucrose. There was a strong positive correlation between the ratio of sucrose to starch and SPS activity in leaves. Therefore, SPS activity is a major determinant of the partitioning of photosynthetically fixed carbon in the leaf and in the whole plant. The photosynthetic rate in air was not significantly increased as a result of elevated leaf SPS activity. However, the light- and CO2-saturated rate of photosynthesis was increased by about 20% in leaves expressing high SPS. In addition, the temporary enhancement of the photosynthetic rate following brief exposures to low light was increased in the high SPS plants relative to controls. We conclude that the level of SPS in the leaves plays a pivotal role in carbon partitioning. Furthermore, high SPS levels have the potential to boost photosynthetic rates under favorable conditions.

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TL;DR: In this paper, Cd markedly decreased ferredoxin(Fd)-dependent NADP+ photoreduction, while it had no effect on electron transport from 2.6dichlorophenolindophenol to methyl viologen, indicating that the metal interferred with electron transport on the reducing side of photosystem I.
Abstract: Photosystem I activity of chloroplasts isolated from 21 days old maize seedlings (Zea mays L. cv. Hidosil) cultivated in a nutrient solution containing different concentrations of Cd (10,20,30μM) was investigated. Cd markedly decreased ferredoxin(Fd)-dependent NADP+ photoreduction, while it had no effect on electron transport from 2. 6-dichlorophenolindophenol to methyl viologen, indicating that the metal interferred with electron transport on the reducing side of photosystem I. The decrease in electron transport correlated with a low Fd content, which in turn was correlated with a low Fe concentration, suggesting Cd-induced Fe deficiency. In in vitro experiments direct Cd inhibition of Fd-dependent NADP+ photoreduction required much higher Cd concentrations than those observed in Cd-treated plants.

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TL;DR: Transgenic potato plants were constructed that have a reduced expression of the TPT at both the RNA and protein level due to antisense inhibition, which leads to a reduction of maximal photosynthesis and a change in carbon partitioning into starch at the expense of sucrose and amino acids.
Abstract: The major chloroplast envelope membrane protein E29 is central for the communication between chloroplasts and cytosol. It has been identified as the triose phosphate translocator (TPT) exporting the primary products of the Calvin cycle (i.e., triose phosphates and 3-phosphoglycerate) out of the chloroplast in a strict counter exchange for Pi. To study the in vivo role of the TPT, transgenic potato plants were constructed that have a reduced expression of the TPT at both the RNA and protein level due to antisense inhibition. Chloroplasts isolated from these plants show a 20-30% reduction with respect to their ability to import Pi. The reduced TPT activity leads to a reduction of maximal photosynthesis by 40-60%, to a change in carbon partitioning into starch at the expense of sucrose and amino acids, and to an increase of the leaf starch content by a factor of approximately 3. At early developmental stages the inhibited plants are retarded in growth compared to the wild type.


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TL;DR: Mass spectrometric analysis showed that O2 uptake was higher in the light than in the dark for both species and in both cases wasHigher inGracilaria sp.
Abstract: The influence of elevated CO2 concentrations on growth and photosynthesis ofGracilaria sp. andG. chilensis was investigated in order to procure information on the effective utilization of CO2. Growth of both was enhanced by CO2 enrichment (air + 650 ppm CO2, air + 1250 ppm CO2, the enhancement being greater inGracilaria sp. Both species increased uptake of NO3 − with CO2 enrichment. Photosynthetic inorganic carbon uptake was depressed inG. chilensis by pre-culture (15 days) with CO2 enrichment, but little affected inGracilaria sp. Mass spectrometric analysis showed that O2 uptake was higher in the light than in the dark for both species and in both cases was higher inGracilaria sp. The higher growth enhancement inGracilaria sp. was attributed to greater depression of photorespiration by the enrichment of CO2 in culture.