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Limits of photosynthesis in extrasolar planetary systems for earth-like planets.

We present a general modeling scheme for investigating the possibility of photosynthesis-based life on extrasolar planets. The scheme focuses on the identification of the habitable zone in main-sequence-star planetary systems with planets of Earth mass and size. Our definition of habitability is based on the long-term possibility of photosynthetic biomass production as a function of mean planetary surface temperature and atmospheric CO2-content. All the astrophysical, climatological, biogeochemical, and geodynamic key processes involved in the generation of photosynthesis-driven life conditions are taken into account. Implicitly, a co-genetic origin of the central star and the orbiting planet is assumed. The numerical solution of an advanced geodynamic model yields realistic look-up diagrams for determining the limits of photosynthesis in extrasolar planetary systems, assuming minimum CO2 levels set by the demand of C4 photosynthesis.

Atmosphere↗

Functional genomics of plant photosynthesis in the fast lane using Chlamydomonas reinhardtii.

Oxygenic photosynthesis by algae and plants supports much of life on Earth. Several model organisms are used to study this vital process, but the unicellular green alga Chlamydomonas reinhardtii offers significant advantages for the genetic dissection of photosynthesis. Recent experiments with Chlamydomonas have substantially advanced our understanding of several aspects of photosynthesis, including chloroplast biogenesis, structure-function relationships in photosynthetic complexes, and environmental regulation. Chlamydomonas is therefore the organism of choice for elucidating detailed functions of the hundreds of genes involved in plant photosynthesis.

Animals↗

Chloroplast redox signals: how photosynthesis controls its own genes.

The photosynthetic apparatus of higher plants and algae is composed of plastid- and nuclear-encoded components, therefore the expression of photosynthesis genes needs to be highly coordinated. Expression is regulated by various factors, one of the most important of which is light. Photosynthesis functions as a sensor for such light signals, and the redox state of photosynthetic electron transport components and redox-active soluble molecules act as regulating parameters. This provides a feedback response loop in which the expression of photosynthesis genes is coupled to the function of the photosynthetic process, and highlights the dual role of photosynthesis in energy fixation and the reception of environmental information.

Animals↗

Bacterial photosynthesis in surface waters of the open ocean.

The oxidation of the global ocean by cyanobacterial oxygenic photosynthesis, about 2,100 Myr ago, is presumed to have limited anoxygenic bacterial photosynthesis to oceanic regions that are both anoxic and illuminated. The discovery of oxygen-requiring photosynthetic bacteria about 20 years ago changed this notion, indicating that anoxygenic bacterial photosynthesis could persist under oxidizing conditions. However, the distribution of aerobic photosynthetic bacteria in the world oceans, their photosynthetic competence and their relationship to oxygenic photoautotrophs on global scales are unknown. Here we report the first biophysical evidence demonstrating that aerobic bacterial photosynthesis is widespread in tropical surface waters of the eastern Pacific Ocean and in temperate coastal waters of the northwestern Atlantic. Our results indicate that these organisms account for 2-5% of the photosynthetic electron transport in the upper ocean.

Bacteria↗

Photosynthesis genes in marine viruses yield proteins during host infection.

Cyanobacteria, and the viruses (phages) that infect them, are significant contributors to the oceanic 'gene pool'. This pool is dynamic, and the transfer of genetic material between hosts and their phages probably influences the genetic and functional diversity of both. For example, photosynthesis genes of cyanobacterial origin have been found in phages that infect Prochlorococcus and Synechococcus, the numerically dominant phototrophs in ocean ecosystems. These genes include psbA, which encodes the photosystem II core reaction centre protein D1, and high-light-inducible (hli) genes. Here we show that phage psbA and hli genes are expressed during infection of Prochlorococcus and are co-transcribed with essential phage capsid genes, and that the amount of phage D1 protein increases steadily over the infective period. We also show that the expression of host photosynthesis genes declines over the course of infection and that replication of the phage genome is a function of photosynthesis. We thus propose that the phage genes are functional in photosynthesis and that they may be increasing phage fitness by supplementing the host production of these proteins.

Bacteriophages↗

The cyanobacterial genome core and the origin of photosynthesis.

Comparative analysis of 15 complete cyanobacterial genome sequences, including "near minimal" genomes of five strains of Prochlorococcus spp., revealed 1,054 protein families [core cyanobacterial clusters of orthologous groups of proteins (core CyOGs)] encoded in at least 14 of them. The majority of the core CyOGs are involved in central cellular functions that are shared with other bacteria; 50 core CyOGs are specific for cyanobacteria, whereas 84 are exclusively shared by cyanobacteria and plants and/or other plastid-carrying eukaryotes, such as diatoms or apicomplexans. The latter group includes 35 families of uncharacterized proteins, which could also be involved in photosynthesis. Only a few components of cyanobacterial photosynthetic machinery are represented in the genomes of the anoxygenic phototrophic bacteria Chlorobium tepidum, Rhodopseudomonas palustris, Chloroflexus aurantiacus, or Heliobacillus mobilis. These observations, coupled with recent geological data on the properties of the ancient phototrophs, suggest that photosynthesis originated in the cyanobacterial lineage under the selective pressures of UV light and depletion of electron donors. We propose that the first phototrophs were anaerobic ancestors of cyanobacteria ("procyanobacteria") that conducted anoxygenic photosynthesis using a photosystem I-like reaction center, somewhat similar to the heterocysts of modern filamentous cyanobacteria. From procyanobacteria, photosynthesis spread to other phyla by way of lateral gene transfer.

Bacterial Proteins↗

Transcriptional regulation and DNA methylation of nuclear genes for photosynthesis in nongreen plant cells.

The transcripts of nuclear genes for the small subunit of ribulose-1,5-bisphosphate carboxylase/oxygenase (rbcS), chlorophyll a/b-binding protein (cab), and extrinsic 33-kDa protein involved in photosystem II water oxidation (woxA) were not detectable in the white wild cultured cells of sycamore (Acer pseudoplatanus), in contrast to their high levels in the sibling green mutant cells and the constitutive expression of actin genes (act) in both cell types. We have examined the template activities of nuclear DNAs using the HeLa cell in vitro transcription system. All of the three photosynthesis genes from the green cell line and act from both cell types were well transcribed in vitro, but these photosynthesis genes from the white cell line were not, indicating that the transcriptional regulation is ascribable to DNA templates. Digestion of nuclear DNA with methyl-sensitive and -insensitive isoschizomeric endonucleases and the subsequent Southern hybridization showed that each gene has the identical recognition sites of restriction enzymes in the green and white cell lines, but some of the sites were methylated only in the photosynthesis genes in the white cells. There was observed a clear inverse relationship between the level of expressed transcripts and the extent of DNA methylation. Thus, it is inferred that the selective methylation of DNA is a likely mechanism for suppressing transcription of nuclear genes for photosynthesis in the nonphotosynthetic plant cells.

Base Composition↗

Evidences from action spectra for a specific participation of chlorophyll b in photosynthesis.

Rate of oxygen evolution in photosynthesis was measured as the current from a polarized platinum electrode covered by a thin layer of Chlorella. The arrangement gave a reproducibly measurable rate of photosynthesis proportional to light intensity at the low levels used and gave rapid response to changes in illumination. Two phenomena have been explored. The Emerson effect was observed as an enhancement of photosynthesis in long wavelength red light (700 mmicro) when shorter wavelengths were added. Two light beams of wavelengths 653 and 700 mmicro when presented together gave a photosynthetic rate about 25 per cent higher than the sum of the rates obtained separately. Large and reproducible transients in rate of oxygen evolution were observed accompanying change in illumination between two wavelengths adjusted in intensity to support equal steady rates of photosynthesis. The transients were found not to be specifically related to long wavelength red light. Both enhancement and the transients have identical action spectra which are interpreted as demonstrating a specific photochemical participation of chlorophyll b.

Chlorophyll↗

A coupled model of photosynthesis, stomatal conductance and transpiration for a rose leaf (Rosa hybrida L.).

The following three models were combined to predict simultaneously photosynthesis, stomatal conductance, transpiration and leaf temperature of a rose leaf: the biochemical model of photosynthesis of Farquhar, von Caemmerer and Berry (1980, Planta 149: 78-90), the stomatal conductance model of Ball, Woodrow and Berry (In: Biggens J, ed. Progress in photosynthesis research. The Netherlands: Martinus Nijhoff Publishers), and an energy balance model. The photosynthetic parameters: maximum carboxylation rate, potential rate of electron transport and rate of triose phosphate utilization, and their temperature dependence were determined using gas exchange data of fully expanded, young, sunlit leaves. The stomatal conductance model was calibrated independently. Prediction of net photosynthesis by the coupled model agreed well with the validation data, but the model tended to underestimate rates of stomatal conductance and transpiration. The coupled model developed in this study can be used to assist growers making environmental control decisions in glasshouse production.

Carbon Dioxide↗

Underwater photosynthesis in flooded terrestrial plants: a matter of leaf plasticity.

BACKGROUND: Flooding causes substantial stress for terrestrial plants, particularly if the floodwater completely submerges the shoot. The main problems during submergence are shortage of oxygen due to the slow diffusion rates of gases in water, and depletion of carbohydrates, which is the substrate for respiration. These two factors together lead to loss of biomass and eventually death of the submerged plants. Although conditions under water are unfavourable with respect to light and carbon dioxide supply, photosynthesis may provide both oxygen and carbohydrates, resulting in continuation of aerobic respiration. SCOPE: This review focuses on evidence in the literature that photosynthesis contributes to survival of terrestrial plants during complete submergence. Furthermore, we discuss relevant morphological and physiological responses of the shoot of terrestrial plant species that enable the positive effects of light on underwater plant performance. CONCLUSIONS: Light increases the survival of terrestrial plants under water, indicating that photosynthesis commonly occurs under these submerged conditions. Such underwater photosynthesis increases both internal oxygen concentrations and carbohydrate contents, compared with plants submerged in the dark, and thereby alleviates the adverse effects of flooding. Additionally, several terrestrial species show high plasticity with respect to their leaf development. In a number of species, leaf morphology changes in response to submergence, probably to facilitate underwater gas exchange. Such increased gas exchange may result in higher assimilation rates, and lower carbon dioxide compensation points under water, which is particularly important at the low carbon dioxide concentrations observed in the field. As a result of higher internal carbon dioxide concentrations in submergence-acclimated plants, underwater photorespiration rates are expected to be lower than in non-acclimated plants. Furthermore, the regulatory mechanisms that induce the switch from terrestrial to submergence-acclimated leaves may be controlled by the same pathways as described for heterophyllous aquatic plants.

Acclimatization↗

Stem photosynthesis not pressurized ventilation is responsible for light-enhanced oxygen supply to submerged roots of alder (Alnus glutinosa).

BACKGROUND AND AIMS: Claims that submerged roots of alder and other wetland trees are aerated by pressurized gas flow generated in the stem by a light-induced thermo-osmosis have seemed inconsistent with root anatomy. Our aim was to seek a verification using physical root-stem models, stem segments with or without artificial roots, and rooted saplings. METHODS: Radial O2 loss (ROL) from roots was monitored polarographically as the gas space system of the models, and stems were pressurized artificially. ROL and internal pressurization were also measured when stems were irradiated and the xylem stream was either CO2 enriched or not. Stem photosynthesis and respiration were measured polarographically. Stem and root anatomy were examined by light and fluorescence microscopy. KEY RESULTS: Pressurizing the models and stems to <or=10 kPa, values much higher than those reportedly generated by thermo-osmosis, created only a negligible density-induced increase in ROL, but ROL increased rapidly when ambient O2 concentrations were raised. Internal pressures rose by several kPa when shoots were exposed to high light flux and ROL increased substantially, but both were due to O2 accumulation from stem photosynthesis using internally sourced CO2. Increased stem pressures had little effect on O2 transport, which remained largely diffusive. Oxygen flux from stems in high light periods indicated a net C gain by stem photosynthesis. Chloroplasts were abundant in the secondary cortex and secondary phloem, and occurred throughout the secondary xylem rays and medulla of 3-year-old stems. Diurnal patterns of ROL, most marked when light reached submerged portions of the stem, were modified by minor variations in light flux and water level. Low root temperatures also helped improve root aeration. CONCLUSIONS: Pressurized gas flow to submerged roots does not occur to any significant degree in alder, but stem photosynthesis, using internally sourced CO2 from respiration and the transpiration stream, may play an important role in root aeration in young trees and measurably affect the overall carbon balance of this and other species.

Alnus↗

QTL analysis of photosynthesis and water status traits in sunflower (Helianthus annuus L.) under greenhouse conditions.

The identification of QTL for several physiological traits in sunflower is described. Traits related to photosynthesis (leaf chlorophyll concentration, net photosynthesis and internal CO(2) concentration) and water status (stomatal conductance, transpiration, predawn leaf water potential, and relative water content) were evaluated in a population of recombinant inbred lines under greenhouse conditions. Narrow-sense heritabilities were low to average. Using an AFLP linkage map, 19 QTL were detected explaining 8.8-62.9% of the phenotypic variance for each trait. Among these, two major QTL for net photosynthesis were identified on linkage group IX. One QTL co-location was found on linkage group VIII for stomatal movements and water status. Coincident locations for QTL regulating photosynthesis, transpiration and leaf water potential were described on linkage group XIV. These results lead to the first description of the organization of genomic regions related to yield in sunflower.

Carbon Dioxide↗

Protecting cotton photosynthesis during moderate chilling at high light intensity by increasing chloroplastic antioxidant enzyme activity.

This study examined the effect of increasing chloroplastic superoxide dismutase (SOD), ascorbate peroxidase (APX), or glutathione reductase (GR) activity via plant transformation of cotton on the initial recovery of photosynthesis following exposures to 10 degrees C and high photon flux density (PFD). Growing wild-type or non-expressing segregate plants (controls) and transformants at two PFDs (600 micromol m(-2) s(-1) and full sun) resulted in a range of total antioxidant enzyme activities. Total SOD activities above that for control leaves grown in full sun did not substantially improve the recoveries of CO(2)-saturated photosynthesis, especially for stress treatments lasting more than 1 h, while elevated APX or GR activity did improve recoveries after 1-3 h of the chilling treatment. No synergistic effects were noted when the activities of more than one antioxidant enzyme were elevated in transgenic hybrids. Although these results suggest that the protection of photosynthesis can be realized by reducing either superoxide or H(2)O(2) levels, thereby reducing the possibility of hydroxyl radical formation, the situation is complicated, since elevated APX or GR activity can improve recoveries even when additional SOD activity has no effect. In conclusion, to enhance the protection of photosynthesis using stroma-targeted antioxidant enzymes, enhancing metabolism associated with H(2)O(2) is more effective than enhancing the capacity for superoxide scavenging. Although small, the improvement in the protection of photosynthetic capacity may be sufficient to improve cotton yield in temperate regions with large diurnal temperature fluctuations.

Antioxidants↗

C(4) photosynthesis: principles of CO(2) concentration and prospects for its introduction into C(3) plants.

C(4) photosynthesis has a number of distinct properties that enable the capture of CO(2) and its concentration in the vicinity of Rubisco, so as to reduce the oxygenase activity of Rubisco, and hence the rate of photorespiration. The aim of this review is to discuss the properties of this CO(2)-concentrating mechanism, and thus to indicate the minimum requirements of any genetically-engineered system. In particular, the Kranz leaf anatomy of C(4) photosynthesis and the division of the C(4)-cycle between two cell types involves intercellular co-operation that requires modifications in regulation and transport to make C(4) photosynthesis work. Some examples of these modifications are discussed. Comparisons are made with the C(4)-type photosynthesis found in single-celled C(4)-type CO(2)-concentrating mechanisms, such as that found in the aquatic plant, Hydrilla verticillata and the single-celled C(4) system found in the terrestrial chenopod Borszczowia aralocaspica. The outcome of recent attempts to engineer C(4) enzymes into C(3) plants is discussed.

Adenosine Triphosphate↗

Variation in acclimation of photosynthesis in Trifolium repens after eight years of exposure to Free Air CO2 Enrichment (FACE).

The initial stimulation of photosynthesis observed on elevation of [CO2] in grasslands has been predicted to be a transient phenomenon constrained by the loss of photosynthetic capacity due to other limitations, notably nutrients and sinks for carbohydrates. Legumes might be expected partially to escape these feedbacks through symbiotic N2 fixation. The Free-Air Carbon dioxide Enrichment (FACE) experiment at Eschikon, Switzerland, has been the longest running investigation of the effects of open-air elevation of [CO2] on vegetation. The prediction of a long-term loss of photosynthetic capacity was tested by analysing photosynthesis in Trifolium repens L. (cv. Milkanova) in the spring and autumn of the eighth, ninth and tenth years of treatment. A high and low N treatment also allowed a test of the significance of exogenous N-supply in maintaining a stimulation of photosynthetic capacity in the long-term. Prior work in this Free Air CO2 Enrichment (FACE) experiment has revealed that elevated [CO2] increased both vegetative and reproductive growth of T. repens independent of N treatment. It is shown here that the photosynthetic response of T. repens was also independent of N fertilization under both current ambient and elevated (600 micro mol mol-1) [CO2]. There was a strong effect of season on photosynthesis, with light-saturated rates (Asat) 37% higher in spring than in autumn. Higher Asat in the spring was supported by higher maximum Rubisco carboxylation rates (Vc,max) and maximum rates of electron transport (Jmax) contributing to RuBP regeneration. Elevated [CO2] increased Asat by 37% when averaged across all measurement periods and both N fertilization levels, and decreased stomatal conductance by 25%. In spring, there was no effect of elevated [CO2] on photosynthetic capacity of leaves, but in autumn both Vc,max and Jmax were reduced by approximately 20% in elevated [CO2]. The results show that acclimation of photosynthetic capacity can occur in a nitrogen-fixing species, in the field where there are no artificial restrictions on sink capacity. However, even with acclimation there was a highly significant increase in photosynthesis at elevated [CO2].

Acclimatization↗

Temperature acclimation of photosynthesis: mechanisms involved in the changes in temperature dependence of photosynthetic rate.

Growth temperature alters temperature dependence of the photosynthetic rate (temperature acclimation). In many species, the optimal temperature that maximizes the photosynthetic rate increases with increasing growth temperature. In this minireview, mechanisms involved in changes in the photosynthesis-temperature curve are discussed. Based on the biochemical model of photosynthesis, change in the photosynthesis-temperature curve is attributable to four factors: intercellular CO2 concentration, activation energy of the maximum rate of RuBP (ribulose-1,5-bisphosphate) carboxylation (Vc max), activation energy of the rate of RuBP regeneration (Jmax), and the ratio of Jmax to Vc max. In the survey, every species increased the activation energy of Vc max with increasing growth temperature. Other factors changed with growth temperature, but their responses were different among species. Among these factors, activation energy of Vc max may be the most important for the shift of optimal temperature of photosynthesis at ambient CO2 concentrations. Physiological and biochemical causes for the change in these parameters are discussed.

Acclimatization↗

Differences between maize and rice in N-use efficiency for photosynthesis and protein allocation.

The N-use efficiency for photosynthesis was higher in a C(4) plant, maize, than in a C(3) plant, rice, including rbcS antisense rice with optimal ribulose-1,5-bisphosphate carboxylase (Rubisco) content for CO(2)-saturated photosynthesis, even when photosynthesis was measured under saturating CO(2) conditions. The N cost for the C(4) cycle enzymes in maize was not large, and the lower amount of Rubisco allowed a greater N investment in the thylakoid components. This greater content of the thylakoid components as well as the CO(2) concentrating mechanism may support higher N-use efficiency for photosynthesis in maize.

Oryza↗

Distinct roles of the cytochrome pathway and alternative oxidase in leaf photosynthesis.

In illuminated leaves, mitochondria are thought to play roles in optimizing photosynthesis. However, the roles of the cytochrome pathway (CP) and alternative oxidase (AOX) in photosynthesis, in particular in the redox state of the photosynthetic electron transport chain, are not separately characterized. We examined the effects of specific inhibition of two respiratory pathways, CP and AOX, on photosynthetic oxygen evolution and the redox state of the photosynthetic electron transport chain in broad bean (Vicia faba L.) leaves under various light intensities. Under saturating photosynthetic photon flux density (PPFD; 700 micromol photon m(-2) s(-1)), inhibition of either pathway caused a decrease in the steady-state levels of the photosynthetic O(2) evolution rate and the PSII operating efficiency, Phi(II). Because these inhibitors, at the concentrations applied to the leaves, had little effect on photosynthesis in the intact chloroplasts, two respiratory pathways are essential for maintenance of high photosynthetic rates at saturating PPFD. CP or AOX inhibition affected to Chl fluorescence parameters (e.g. photochemical quenching and non-photochemical quenching) differently, suggesting that CP and AOX contribute to photosynthesis in different ways. At low PPFD (100 micromol photon m(-2) s(-1)), only the inhibition of AOX, not CP, lowered the photosynthetic rate and Phi(II). AOX inhibition also decreased the Phi(II)/Phi(I) ratio even at low PPFD levels. These data suggest that AOX inhibition caused the over-reduction of the photosynthetic electron transport chain and induced the cyclic electron flow around PSI (CEF-PSI) even at the low PPFD. Based on these results, we discuss possible roles for CP and AOX in the light.

Antimycin A↗