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A plant notices insect egg deposition and changes its rate of photosynthesis.

Scots pine (Pinus sylvestris) is known to change its terpenoid metabolism in response to egg deposition by the sawfly Diprion pini (Hymenoptera, Diprionidae). Three days after egg deposition, parts of the pine twig adjacent to the egg-laden one are induced to emit volatiles, which attract egg parasitoids. In this study, we investigated whether egg deposition by this sawfly affects pine photosynthesis. Measurements of photosynthesis were taken from untreated control twigs and from pine twigs adjacent to egg-laden ones (i.e. systemically oviposition-induced twigs) for a period of 3 d starting after egg deposition. The net photosynthetic rate of oviposition-induced pine twigs was lower than that of untreated control twigs, whereas the respiration rate of pine twigs was not affected by egg deposition. CO2 response curves of oviposition-induced twigs tended to be lower than those of controls. The potential rate of electron transport (J(max)) and the maximum rate of Rubisco activity (V(cmax)) were calculated from the data of the CO2 response curves. J(max) of oviposition-induced twigs was significantly lower than that of controls at day 1 after egg deposition, while the difference diminished from day 2 to day 3. A similar pattern was observed for V(cmax). Light response curves of oviposition-induced twigs were significantly lower than those of untreated ones during 3 d of measurements. Stomatal conductance was slightly lowered by egg deposition. When considering photosynthetic activity as a physiological currency to measure costs of induction of plant defense, the effects of insect egg deposition on gas exchange of pine are discussed with respect to known effects of insect feeding on the photosynthesis activity of plants.

Acclimatization↗

Carbonic anhydrase and its influence on carbon isotope discrimination during C4 photosynthesis. Insights from antisense RNA in Flaveria bidentis.

In C4 plants, carbonic anhydrase (CA) facilitates both the chemical and isotopic equilibration of atmospheric CO2 and bicarbonate (HCO3-) in the mesophyll cytoplasm. The CA-catalyzed reaction is essential for C4 photosynthesis, and the model of carbon isotope discrimination (Delta13C) in C4 plants predicts that changes in CA activity will influence Delta13C. However, experimentally, the influence of CA on Delta13C has not been demonstrated in C4 plants. Here, we compared measurements of Delta13C during C4 photosynthesis in Flaveria bidentis wild-type plants with F. bidentis plants with reduced levels of CA due to the expression of antisense constructs targeted to a putative mesophyll cytosolic CA. Plants with reduced CA activity had greater Delta13C, which was also evident in the leaf dry matter carbon isotope composition (delta13C). Contrary to the isotope measurements, photosynthetic rates were not affected until CA activity was less than 20% of wild type. Measurements of Delta13C, delta13C of leaf dry matter, and rates of net CO2 assimilation were all dramatically altered when CA activity was less than 5% of wild type. CA activity in wild-type F. bidentis is sufficient to maintain net CO2 assimilation; however, reducing leaf CA activity has a relatively large influence on Delta13C, often without changes in net CO2 assimilation. Our data indicate that the extent of CA activity in C4 leaves needs to be taken into account when using Delta13C and/or delta13C to model the response of C4 photosynthesis to changing environmental conditions.

Carbon Dioxide↗

Two genes encode highly similar chloroplastic NADP-malic enzymes in Flaveria. Implications for the evolution of C4 photosynthesis.

To gain an understanding of the molecular events underlying the evolution of C4 photosynthesis, we have undertaken as detailed study of the NADP-malic enzyme gene family in C4 and C3 species of Flaveria. Three genomic clones form the C4 species Flaveria bidentis were characterized and found to encode two highly similar chloroplastic forms of NADP-malic enzyme, termed ME1 and ME2. Genomic southern blotting with gene-specific probes showed that both Me1 and Me2 are found in Flaveria trinervia (C4) and Flaveria pringlei (C3) as well as in F. bidentis. Northern blots demonstrated that Me1 expression in leaves parallels the degree of C4 photosynthesis in seven Flaveria species. Furthermore, whereas Me2 was expressed at a low level in both roots and leaves of F. bidentis, Me1 expression was seen only in leaves and was light-regulated. We discuss these results in the context of the evolution of C4 photosynthesis in Flaveria.

Base Sequence↗

Growth and photosynthesis under high and low irradiance of Arabidopsis thaliana antisense mutants with reduced ribulose-1,5-bisphosphate carboxylase/oxygenase activase content.

Photosynthesis and growth to maturity of antisense ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) activase Arabidopsis thaliana with reduced concentrations of activase relative to wild-type (Wt) plants were measured under low (200 mumol m-2 s-1) and high (600 mumol m-2 s-1) photosynthetic photon flux density growing conditions. Both growth and photosynthesis were significantly reduced in an Arabidopsis clone (R100) with 30 to 40% Wt activase, an effect that was more pronounced in high light. The aboveground biomass of the antisense clone R100 reached 80% of Wt under low light and 65% of Wt under high light. Decreased growth in the antisense plants was attributed to reduced relative rates of growth and leaf area expansion early in development; all plants attained similar values of relative rates of growth and leaf elongation by 21 d after planting. Reductions in photosynthesis were attributed to decreased Rubisco activation in the antisense plants. Rubisco constituted about 40% of total soluble protein in both Wt and clone R100 under both light regimes. Activase content was 5% and 1.4% of total soluble protein in Wt and clone R100, respectively, and also was unaffected by growth irradiance. The stoichiometry of Rubisco to activase was estimated at 20 Rubisco active sites per activase tetramer in Wt Arabidopsis and 60 to 80 in the transgenic clone R100. We conclude that Wt Arabidopsis does not contain Rubisco activase in great excess of the amount required for optimal growth.

Antisense Elements (Genetics)↗

Differential expression of photosynthesis and nitrogen fixation genes in the cyanobacterium Plectonema boryanum.

The filamentous non-heterocystous cyanobacterium Plectonema boryanum fixes dinitrogen at a high rate during microaerobic growth in continuous illumination by temporal separation of oxygen-evolving photosynthesis and oxygen-sensitive dinitrogen fixation. The onset of nitrogen fixation is preceded by a depression in photosynthesis that establishes a sufficiently low level of dissolved oxygen in the growth medium. A several-fold reduction in the level of transcripts coding for phycocyanin (cpcBA) and the chlorophyll a binding protein of photosystem II (psbC) and psbA accompanied the depression in photosynthetic oxygen evolution. Unlike most of the other organisms examined to date, in P. boryanum, psbC and psbD do not appear to be co-transcribed. The psbC transcripts were down-regulated several fold, while the psbD transcript declined marginally during the nitrogen fixation phase. A decrease in dissolved oxygen and a dramatic increase in the level of nifH transcripts and the enzyme activity of nitrogenase were characteristic of the nitrogen fixation phase. The level of transcript for glnA, which encodes glutamine synthetase, was not altered. Reciprocal regulation of gene expression was well orchestrated with the alternating cycles of photosynthesis and nitrogen fixation in P. boryanum.

Cyanobacteria↗

Targeted inactivation of the plastid ndhB gene in tobacco results in an enhanced sensitivity of photosynthesis to moderate stomatal closure.

The ndh genes encoding for the subunits of NAD(P)H dehydrogenase complex represent the largest family of plastid genes without a clearly defined function. Tobacco (Nicotiana tabacum) plastid transformants were produced in which the ndhB gene was inactivated by replacing it with a mutant version possessing translational stops in the coding region. Western-blot analysis indicated that no functional NAD(P)H dehydrogenase complex can be assembled in the plastid transformants. Chlorophyll fluorescence measurements showed that dark reduction of the plastoquinone pool by stromal reductants was impaired in ndhB-inactivated plants. Both the phenotype and photosynthetic performance of the plastid transformants was completely normal under favorable conditions. However, an enhanced growth retardation of ndhB-inactivated plants was revealed under humidity stress conditions causing a moderate decline in photosynthesis via stomatal closure. This distinctive phenotype was mimicked under normal humidity by spraying plants with abscisic acid. Measurements of CO(2) fixation demonstrated an enhanced decline in photosynthesis in the mutant plants under humidity stress, which could be restored to wild-type levels by elevating the external CO(2) concentration. These results suggest that the plastid NAD(P)H:plastoquinone oxidoreductase in tobacco performs a significant physiological role by facilitating photosynthesis at moderate CO(2) limitation.

Abscisic Acid↗

Increased sensitivity of photosynthesis to antimycin A induced by inactivation of the chloroplast ndhB gene. Evidence for a participation of the NADH-dehydrogenase complex to cyclic electron flow around photosystem I.

Tobacco (Nicotiana tabacum var Petit Havana) ndhB-inactivated mutants (ndhB-) obtained by plastid transformation (E.M. Horvath, S.O. Peter, T. Joët, D. Rumeau, L. Cournac, G.V. Horvath, T.A. Kavanagh, C. Schäfer, G. Peltier, P. MedgyesyHorvath [2000] Plant Physiol 123: 1337-1350) were used to study the role of the NADH-dehydrogenase complex (NDH) during photosynthesis and particularly the involvement of this complex in cyclic electron flow around photosystem I (PSI). Photosynthetic activity was determined on leaf discs by measuring CO2 exchange and chlorophyll fluorescence quenchings during a dark-to-light transition. In the absence of treatment, both non-photochemical and photochemical fluorescence quenchings were similar in ndhB- and wild type (WT). When leaf discs were treated with 5 microM antimycin A, an inhibitor of cyclic electron flow around PSI, both quenchings were strongly affected. At steady state, maximum photosynthetic electron transport activity was inhibited by 20% in WT and by 50% in ndhB-. Under non-photorespiratory conditions (2% O2, 2,500 microL x L(-1) CO2), antimycin A had no effect on photosynthetic activity of WT, whereas a 30% inhibition was observed both on quantum yield of photosynthesis assayed by chlorophyll fluorescence and on CO2 assimilation in ndhB-. The effect of antimycin A on ndhB- could not be mimicked by myxothiazol, an inhibitor of the mitochondrial cytochrome bc1 complex, therefore showing that it is not related to an inhibition of the mitochondrial electron transport chain but rather to an inhibition of cyclic electron flow around PSI. We conclude to the existence of two different pathways of cyclic electron flow operating around PSI in higher plant chloroplasts. One of these pathways, sensitive to antimycin A, probably involves ferredoxin plastoquinone reductase, whereas the other involves the NDH complex. The absence of visible phenotype in ndhB- plants under normal conditions is explained by the complement of these two pathways in the supply of extra-ATP for photosynthesis.

Antimycin A↗

Studies on the biochemistry and fine structure of silica shell formation in diatoms. Photosynthesis and respiration in silicon-starvation synchrony of Navicula pelliculosa.

Rates of photosynthesis, measured by oxygen electrode or by (14)CO(2) fixation, dark respiration and (32)P-phosphate incorporation are reported for the silicon-starvation synchrony of the fresh water diatom Navicula pelliculosa. During late exponential growth the rates were consistent with increase in carbon mass. During silicon starvation, rates of carbon dioxide fixation, oxygen evolution and (32)P incorporation fell, and the saturating light intensity decreased from 27,000 lux to 5000 lux. Reintroduction of silicon led to immediate transients in all parameters studied, followed by a prolonged increase in rate of dark respiration and a gradual increase in apparent photosynthesis. During release of daughter cells, the rates of dark respiration decreased as photosynthesis and (32)P incorporation increased. These results are discussed in relation to effects of silicon on the energy metabolism of the diatom.

Carbon Dioxide↗

Biphasic inhibition of photosynthesis in powdery mildewed barley.

A careful restudy of the photosynthetic process of barley (Hordeum vulgare) infected with powdery mildew (Erysiphe graminis f. sp. hordei) is reported. Unlike previous reports, which indicated a stimulation in infected host tissue photosynthesis during early stages of the disease followed by a rapid decline in activity, this study observed no stimulation but instead a biphasic inhibition in infected host photosynthesis under physiological concentrations of CO(2). Under high CO(2) (1.0%) a stimulation was observed during early stages of infection. The inhibition in low CO(2) and stimulation in high CO(2) of infected host photosynthesis was shown to occur in healthy host tissue when in the presence of 10(-2)m alpha-hydroxy-2-pyridinemethanesulfonic acid. The possible significance of these similar results is discussed.

Ascomycota↗

Overexpression of Arabidopsis hexokinase in tomato plants inhibits growth, reduces photosynthesis, and induces rapid senescence.

Sugars are key regulatory molecules that affect diverse processes in higher plants. Hexokinase is the first enzyme in hexose metabolism and may be a sugar sensor that mediates sugar regulation. We present evidence that hexokinase is involved in sensing endogenous levels of sugars in photosynthetic tissues and that it participates in the regulation of senescence, photosynthesis, and growth in seedlings as well as in mature plants. Transgenic tomato plants overexpressing the Arabidopsis hexokinase-encoding gene AtHXK1 were produced. Independent transgenic plants carrying single copies of AtHXK1 were characterized by growth inhibition, the degree of which was found to correlate directly to the expression and activity of AtHXK1. Reciprocal grafting experiments suggested that the inhibitory effect occurred when AtHXK1 was expressed in photosynthetic tissues. Accordingly, plants with increased AtHXK1 activity had reduced chlorophyll content in their leaves, reduced photosynthesis rates, and reduced photochemical quantum efficiency of photosystem II reaction centers compared with plants without increased AtHXK1 activity. In addition, the transgenic plants underwent rapid senescence, suggesting that hexokinase is also involved in senescence regulation. Fruit weight, starch content in young fruits, and total soluble solids in mature fruits were also reduced in the transgenic plants. The results indicate that endogenous hexokinase activity is not rate limiting for growth; rather, they support the role of hexokinase as a regulatory enzyme in photosynthetic tissues, in which it regulates photosynthesis, growth, and senescence.

Arabidopsis↗

Light-dependent regulation of photosynthesis genes in Rhodobacter sphaeroides 2.4.1 is coordinately controlled by photosynthetic electron transport via the PrrBA two-component system and the photoreceptor AppA.

Formation of the photosynthetic apparatus in Rhodobacter is regulated by oxygen tension and light intensity. Here we show that in anaerobically grown Rhodobacter cells a light-dependent increase in expression of the puc and puf operons encoding structural proteins of the photosynthetic complexes requires an active photosynthetic electron transport. The redox-sensitive CrtJ/PpsR repressor of photosynthesis genes, which was suggested to mediate electron transport-dependent signals, is not involved in this light-dependent signal chain. Our data reveal that the signal initiated in the photosynthetic reaction centre is transmitted via components of the electron transport chain and the PrrB/PrrA two-component system in Rhodobacter sphaeroides. Under blue light illumination in the absence of oxygen this signal leads to activation of photosynthesis genes and interferes with a blue-light repression mediated by the AppA photoreceptor and the PpsR transcriptional repressor in R. sphaeroides. Thus, light either sensed by a photoreceptor or initiating photosynthetic electron transport has opposite effects on the transcription of photosynthesis genes. Both signalling pathways involve redox-dependent steps that finally determine the effect of light on gene expression.

Anaerobiosis↗

Can improvement in photosynthesis increase crop yields?

The yield potential (Yp) of a grain crop is the seed mass per unit ground area obtained under optimum growing conditions without weeds, pests and diseases. It is determined by the product of the available light energy and by the genetically determined properties: efficiency of light capture (epsilon i), the efficiency of conversion of the intercepted light into biomass (epsilon c) and the proportion of biomass partitioned into grain (eta). Plant breeding brings eta7 and epsilon i close to their theoretical maxima, leaving epsilon c, primarily determined by photosynthesis, as the only remaining major prospect for improving Yp. Leaf photosynthetic rate, however, is poorly correlated with yield when different genotypes of a crop species are compared. This led to the viewpoint that improvement of leaf photosynthesis has little value for improving Yp. By contrast, the many recent experiments that compare the growth of a genotype in current and future projected elevated [CO2] environments show that increase in leaf photosynthesis is closely associated with similar increases in yield. Are there opportunities to achieve similar increases by genetic manipulation? Six potential routes of increasing epsilon c by improving photosynthetic efficiency were explored, ranging from altered canopy architecture to improved regeneration of the acceptor molecule for CO2. Collectively, these changes could improve epsilon c and, therefore, Y p by c. 50%. Because some changes could be achieved by transgenic technology, the time of the development of commercial cultivars could be considerably less than by conventional breeding and potentially, within 10-15 years.

Biomass↗

Water relations, thallus structure and photosynthesis in Negev Desert lichens.

The role of lichen thallus structure in water relations and photosynthesis was studied in Ramalina maciformis (Del.) Bory and Teloschistes lacunosus (Rupr.) Sav. Water-vapour adsorption and photosynthesis are dependent upon thallus integrity and are significantly lower in crushed thalli. Cultured phycobiont (Trebouxia sp.) cells are capable of photosynthesis over the same relative humidity range (> 80% RH) as are intact lichens. Thus, water-vapour adsorption by the thallus and physiological adaptation of the phycobiont contribute to the ability of these lichens to photosynthesize in an arid environment. Despite differences in their anatomical structure and water-uptake characteristics, their CO2 incorporation is similar. The two lichens use liquid water differently and they occupy different niches.

Carbon Dioxide↗

Pigments, photosynthesis and photoinhibition in two amphibious plants: consequences of varying carbon availability.

In the present study, we investigated the effects of CO(2) availability on photosynthesis, photoinhibition and pigmentation in two species of amphibious plants, Lobelia cardinalis and Nesaea crassicaulis. The plants were grown emergent under atmospheric conditions and submerged under low and high CO(2) availability. Compared with Lobelia, Nesaea had thin leaves and few stomata in all CO(2) treatments. While Lobelia expressed no variation in anthocyanin content among treatments, Nesaea produced high concentrations of anthocyanin when submerged. Lobelia photosynthesis increased in response to increasing CO(2) availability, and photoinhibition was negatively related to xanthophyll content. By contrast, Nesaea photosynthesis was highest under submerged conditions, and there was no relationship between photoinhibition and the xanthophyll content. We conclude that the response of Lobelia to varying CO(2) availability is similar to that of terrestrial plants and that this species relies on the xanthophyll cycle for nonphotochemical quenching (NPQ) and protection against photoinhibition. By contrast, the thin leaves, few stomata and low levels of chlorophylls and accessory pigments in Nesaea, relative to Lobelia, suggest adaptation to a submerged habitat. While Nesaea does not seem to rely on the xanthophyll cycle or other xanthophylls for NPQ, some role of anthocyanins in the protection against photoinhibition cannot be ruled out, owing to its effect as a sunscreen and as an efficient quencher of free radicals.

Anthocyanins↗

Alleviation of CO2-Induced Reductions in Tomato Photosynthesis Under Deficit Irrigation by Purple Nonsulfur Photosynthetic Bacteria.

The stimulatory effect of elevated CO2 (eCO2) on photosynthesis in most C3 crops under water deficit often declines over time due to photosynthetic acclimation. An exception occurs in plants inoculated with symbiotic nitrogen-fixing bacteria. Photosynthetic bacteria (PSB), specifically anoxygenic purple nonsulfur bacteria (Rhodopseudomonas palustris in this study), a group of nitrogen-fixing bacteria, are effective in enhancing crop photosynthesis. Therefore, this study investigated the synergistic effects of PSB and eCO2 in alleviating the effects of deficit irrigation and enhancing photosynthetic capacity in tomato plants during prolonged exposure. Our results showed that photosynthetic efficiency was significantly reduced in noninoculated plants under eCO2, and this reduction was more pronounced under water deficit. Proteomic analysis revealed that in eCO2-treated plants, the downregulation of cell wall proteins increased mesophyll resistance to CO2 diffusion, while the suppression of the photosynthetic apparatus impaired electron transport capacity, ultimately reducing CO2 assimilation efficiency. In contrast, these negative effects were alleviated by PSB inoculation. PSB promoted the upregulation of proteins involved in photosynthesis under deficit irrigation, as well as proteins related to chlorophyll biosynthesis, components of photosystem I and II, and light-harvesting complex proteins. These proteins contributed to improved photosynthetic efficiency during deficit irrigation and photosynthetic acclimation. Physiological analyses further confirmed that PSB inoculation enhanced nitrogen content, electron transport capacity, chlorophyll biosynthesis, and overall photosynthetic performance under eCO2 and deficit irrigation, resulting in improved plant growth. These findings suggest that PSB inoculation is a promising strategy to sustain and enhance the CO2 fertilization effect on crop productivity under water-limited conditions.

Photosynthesis↗

Segregation of nitrogen fixation and oxygenic photosynthesis in the marine cyanobacterium Trichodesmium.

In the modern ocean, a significant amount of nitrogen fixation is attributed to filamentous, nonheterocystous cyanobacteria of the genus Trichodesmium. In these organisms, nitrogen fixation is confined to the photoperiod and occurs simultaneously with oxygenic photosynthesis. Nitrogenase, the enzyme responsible for biological N2 fixation, is irreversibly inhibited by oxygen in vitro. How nitrogenase is protected from damage by photosynthetically produced O2 was once an enigma. Using fast repetition rate fluorometry and fluorescence kinetic microscopy, we show that there is both temporal and spatial segregation of N2 fixation and photosynthesis within the photoperiod. Linear photosynthetic electron transport protects nitrogenase by reducing photosynthetically evolved O2 in photosystem I (PSI). We postulate that in the early evolutionary phase of oxygenic photosynthesis, nitrogenase served as an electron acceptor for anaerobic heterotrophic metabolism and that PSI was favored by selection because it provided a micro-anaerobic environment for N2 fixation in cyanobacteria.

Aerobiosis↗

Tight coupling of root-associated nitrogen fixation and plant photosynthesis in the salt marsh grass Spartina alterniflora and carbon dioxide enhancement of nitrogenase activity.

The coupling of root-associated nitrogen fixation and plant photosynthesis was examined in the salt marsh grass Spartina alterniflora. In both field experiments and hydroponic assay chambers, nitrogen fixation associated with the roots was rapidly enhanced by stimulating plant photosynthesis. A kinetic analysis of acetylene reduction activity (ARA) showed that a five-to sixfold stimulation occurred within 10 to 60 min after the plant leaves were exposed to light or increased CO2 concentrations (with the light held constant). In field experiments, CO2 enrichment increased plant-associated ARA by 27%. Further evidence of the dependence of ARA on plant photosynthate was obtained when activity in excised roots was shown to decrease after young greenhouse plants were placed in the dark. Seasonal variation in the ARA of excised plant roots from field cores appears to be related to the annual cycle of net photosynthesis in S. alterniflora.

Acetylene↗

Proline-rich peptide from the coral pathogen Vibrio shiloi that inhibits photosynthesis of Zooxanthellae.

The coral-bleaching bacterium Vibrio shiloi biosynthesizes and secretes an extracellular peptide, referred to as toxin P, which inhibits photosynthesis of coral symbiotic algae (zooxanthellae). Toxin P was produced during the stationary phase when the bacterium was grown on peptone or Casamino Acids media at 29 degrees C. Glycerol inhibited the production of toxin P. Toxin P was purified to homogeneity, yielding the following 12-residue peptide: PYPVYAPPPVVP (molecular weight, 1,295.54). The structure of toxin P was confirmed by chemical synthesis. In the presence of 12.5 mM NH(4)Cl, pure natural or synthetic toxin P (10 microM) caused a 64% decrease in the photosynthetic quantum yield of zooxanthellae within 5 min. The inhibition was proportional to the toxin P concentration. Toxin P bound avidly to zooxanthellae, such that subsequent addition of NH(4)Cl resulted in rapid inhibition of photosynthesis. When zooxanthellae were incubated in the presence of NH(4)Cl and toxin P, there was a rapid decrease in the pH (pH 7.8 to 7.2) of the bulk liquid, suggesting that toxin P facilitates transport of NH(3) into the cell. It is known that uptake of NH(3) into cells can destroy the pH gradient and block photosynthesis. This mode of action of toxin P can help explain the mechanism of coral bleaching by V. shiloi.

Animals↗