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Aerobic anoxygenic photosynthesis in Roseobacter clade bacteria from diverse marine habitats.

The marine Roseobacter clade comprises several genera of marine bacteria related to the uncultured SAR83 cluster, the second most abundant marine picoplankton lineage. Cultivated representatives of this clade are physiologically heterogeneous, and only some have the capability for aerobic anoxygenic photosynthesis, a process of potentially great ecological importance in the world's oceans. In an attempt to correlate phylogeny with ecology, we investigated the diversity of Roseobacter clade strains from various marine habitats (water samples, biofilms, laminariae, diatoms, and dinoflagellate cultures) by using the 16S rRNA gene as a phylogenetic marker gene. The potential for aerobic anoxygenic photosynthesis was determined on the genetic level by PCR amplification and sequencing of the pufLM genes of the bacterial photosynthesis reaction center and on the physiological level by detection of bacteriochlorophyll (Bchl) a. A collection of ca. 1,000 marine isolates was screened for members of the marine Roseobacter clade by 16S rRNA gene-directed multiplex PCR and sequencing. The 42 Roseobacter clade isolates found tended to form habitat-specific subclusters. The pufLM genes were detected in two groups of strains from dinoflagellate cultures but in none of the other Roseobacter clade isolates. Strains within the first group (the DFL-12 cluster) also synthesized Bchl a. Strains within the second group (the DFL-35 cluster) formed a new species of Roseovarius and did not produce Bchl a under the conditions investigated here, thus demonstrating the importance of genetic methods for screening of cultivation-dependent metabolic traits. The pufL genes of the dinoflagellate isolates were phylogenetically closely related to pufL genes from Betaproteobacteria, confirming similar previous observations which have been interpreted as indications of gene transfer events.

Aerobiosis↗

Mobilization of the genes for photosynthesis from Rhodopseudomonas capsulata by a promiscuous plasmid.

Plasmid pBLM2, a derivative of RP1 with enhanced chromosome mobilization activity in Rhodopseudomonas capsulata, was isolated by screening rare exconjugant clones for sex factor activity. pBLM2 mobilized all known genes affecting photosynthesis as well as chromosomal genes for streptomycin and rifampin resistance and tryptophan and cytochrome biosynthesis. Tight linkage was exhibited among the genes affecting photosynthesis. The frequency of successful transfer of chromosomal markers reached 6 X 10(-4) per donor cell. R-primes were occasionally formed during conjugation, and a number of R-primes bearing the genes for photosynthesis were isolated by screening R. capsulata exconjugants with complementation phenotypes for the ability to transmit plasmid-borne R. capsulata genes to Escherichia coli cells. These R-primes were unstable in R. capsulata, but stable in E. coli or Pseudomonas fluorescens. Complementation and recombination events that occurred upon introduction of R-primes into R. capsulata mutants with altered photosynthetic apparatuses could be visualized as variations in colony pigmentation. Each R-prime studied complemented all known types of mutation affecting the differentiation of the photosynthetic apparatus, and no other R. capsulata gene was identified on those plasmids. The R. capsulata genes borne on the R-primes were not functional in E. coli or P. fluorescens.

Conjugation, Genetic↗

prrA, a putative response regulator involved in oxygen regulation of photosynthesis gene expression in Rhodobacter sphaeroides.

A new locus, prrA, involved in the regulation of photosynthesis gene expression in response to oxygen, has been identified in Rhodobacter sphaeroides. Inactivation of prrA results in the absence of photosynthetic spectral complexes. The prrA gene product has strong homology to response regulators associated with signal transduction in other prokaryotes. When prrA is present in multiple copies, cells produce light-harvesting complexes under aerobic growth conditions, suggesting that prrA affects photosynthesis gene expression positively in response to oxygen deprivation. Analysis of the expression of puc::lacZ fusions in wild-type and PrrA- cells revealed a substantial decrease in LacZ expression in the absence of prrA under all conditions of growth, especially when cells were grown anaerobically in the dark in the presence of dimethyl sulfoxide. Northern (RNA) and slot blot hybridizations confirmed the beta-galactoside results for puc and revealed additional positive regulation of puf, puhA, and cycA by PrrA. The effect of truncated PrrA on photosynthesis gene expression in the presence of low oxygen levels can be explained by assuming that PrrA may be effective as a multimer. PrrA was found to act on the downstream regulatory sequences (J. K. Lee and S. Kaplan, J. Bacteriol. 174:1146-1157, 1992) of the puc operon regulatory region. Finally, two spontaneous prrA mutations that abolish prrA function by changing amino acids in the amino-terminal domain of the protein were isolated.

Aldehyde Oxidoreductases↗

appA, a novel gene encoding a trans-acting factor involved in the regulation of photosynthesis gene expression in Rhodobacter sphaeroides 2.4.1.

A new gene, the product of which is involved in the regulation of photosynthesis gene expression in the anoxygenic photosynthetic bacterium Rhodobacter sphaeroides 2.4.1, has been identified. The isolation of this gene, designated appA (activation of photopigment and puc expression), was based on its ability, when provided in extra copies, to partially suppress mutations in the two-component PrrB-PrrA regulatory system. The presence of extra copies of the appA gene in either prrB, prrA, or wild-type strains resulted in an activation of puc::lacZ expression under aerobic conditions. Constructed AppA null mutants did not grow photosynthetically and were impaired in the synthesis of both bacteriochlorophyll and carotenoids, as well as the structural proteins of the photosynthetic spectral complexes. When grown anaerobically in the dark, these mutants accumulated bacteriochlorophyll precursors. The expression of lacZ fusions to several photosynthesis genes and operons, including puc, puf, and bchF, was decreased in the AppA mutant strains in comparison with the wild type. To examine the role of AppA involvement in bacteriochlorophyll biosynthesis, we inactivated an early gene, bchE, of the bacteriochlorophyll pathway in both wild-type and AppA- mutant backgrounds. The double mutant, AppA- BchE-, was found to be severely impaired in photosynthesis gene expression, similar to the AppA- BchE+ mutant and in contrast to the AppA+ BchE- mutant. This result indicated that AppA is more likely involved in the regulation of expression of the bch genes than in the biosynthetic pathway per se. The appA gene was sequenced and appears to encode a protein of 450 amino acids with no obvious homology to known proteins.

Aerobiosis↗

Complex regulatory activities associated with the histidine kinase PrrB in expression of photosynthesis genes in Rhodobacter sphaeroides 2.4.1.

Rhodobacter sphaeroides 2.4.1 synthesizes a specialized photosynthetic membrane upon reduction of the O2 tension below threshold levels. The genes prrB and prrA encode a sensor kinase and a response regulator, respectively, of a two-component regulatory system that presumably is involved in transduction of the signal(s) that monitors alterations in oxygen levels. A third gene, prrC, is also involved in this cascade of events. Previously, we described a mutant form of PrrB, namely, PrrB78 (J. M. Eraso and S. Kaplan, J. Bacteriol. 177:2695-2706, 1995), which results in aerobic expression of the photosynthetic apparatus. Here we examine three mutated forms of the prrB gene that have the potential to encode truncated polypeptides containing the N-terminal 6, 63, or 163 amino acids, respectively. The resulting mutant strains showed residual levels of the light-harvesting spectral complexes and had diminished photosynthetic growth rates at high light intensities with no discernible growth under intermediate or low light conditions. When either lacZ transcriptional fusions or direct mRNA determinations were used to monitor specific photosynthesis gene expression, all the mutant strains showed unexpectedly high levels of gene expression when compared to mutant strains affected in prrA. Conversely, when translational fusions were used to monitor photosynthesis gene expression in these mutant strains, expression of both puc and puf operons was reduced, especially puf expression. In light of these studies and those of the PrrB78 mutant, the data suggest that PrrA can be activated in situ by something other than PrrB, and it also appears that PrrB can function as a negative regulator acting through PrrA. Finally, we consider the role of the Prr regulatory system in the posttranscriptional control of photosynthesis gene expression.

Bacterial Proteins↗

Evidence for the role of redox carriers in photosynthesis gene expression and carotenoid biosynthesis in Rhodobacter sphaeroides 2.4.1.

Previous work from this laboratory revealed that alterations in the structure of the ccoNOQP operon of Rhodobacter sphaeroides 2.4.1 could lead to induction of the photosynthetic apparatus under aerobic growth conditions. Immediately downstream of the ccoNOQP operon is the rdxB gene, the first gene of the rdxBHIS cluster. The rdxB gene product is predicted to encode a membrane protein which can bind two [4Fe-4S] clusters. The ccoP gene product is a diheme cytochrome which is a component of the cbb3-type cytochrome oxidase. Under aerobic growth conditions, strains possessing ccoP and rdxB mutations both singly and in combination produced light-harvesting complexes, suggesting that normal functioning of these proteins is required to maintain repression of photosynthesis gene expression in the presence of oxygen. Analysis of the expression of puc::lacZ fusions under aerobic conditions revealed an approximately 12-fold increase in puc operon expression in the RDXB1 and CCOP1 mutant strains compared with that for wild-type 2.4.1. Similarly, puf::lacZ activity was observed to be elevated fourfold above wild-type levels. Further indication of the importance of the RdxB and CcoP proteins was derived from studies of mutant and wild-type cells grown under anoxygenic photosynthetic and nitrogen-fixing conditions. These mutant strains were observed to accumulate spheroidenone to approximately 50% or more of the total carotenoid. In wild-type cultures, spheroidenone normally accumulates to approximately 10 to 20% of the total carotenoid under the same growth conditions. This effect was most pronounced when both the rdxB and the ccoP mutations were present together in cells cultured under nitrogen-fixing photosynthetic growth conditions in which spheroidenone represented approximately 90% of the total carotenoid. We propose that mutations in the rdxB or ccoP gene may lead to changes in a membrane-generated redox signal or the accumulation of a critical redox intermediate in the mutant strains which results in increased photosynthesis gene expression under aerobic conditions by alteration of the activity of a transcriptional regulator(s) of photosynthesis gene expression. Mutations in these genes also appear to posttranscriptionally influence the terminal step of carotenoid biogenesis. Potential regulators interacting with an aberrant redox signal in the mutants and the possible nature of such a redox signal are discussed.

Aerobiosis↗

A redox-responsive pathway for aerobic regulation of photosynthesis gene expression in Rhodobacter sphaeroides 2.4.1.

To further understand the proposed signal transduction pathway involving the presumed redox proteins RdxBH and cbb3 cytochrome oxidase in Rhodobacter sphaeroides 2.4.1, a series of mutants lacking components of both the Prr two-component activation system and the cbb3-type cytochrome oxidase or RdxBH were constructed. We report that under highly aerobic conditions, aberrant photosynthesis gene expression and spectral complex formation typical of cbb3- or RdxBH-deficient mutants were no longer observed when either prrA (encoding the response regulator of the Prr system) or prrB (encoding the presumed sensor kinase) was also deleted. These double-mutant strains are phenotypically identical to single-mutant PrrA and PrrB strains, suggesting that the signal(s) originating from the cbb3 terminal oxidase affects downstream puc and puf operon expression by acting exclusively through the Prr system. When the same double-mutant strains were examined under anaerobic dark dimethyl sulfoxide growth conditions, photosynthesis gene expression was obligatorily linked to the two-component activation system. However, photosynthesis gene expression under the same growth conditions was significantly higher in the cbb3 mutant strain when compared to that in the wild type. Similarly, under anaerobic photosynthetic conditions the high levels of the oxidized carotenoid, spheroidenone, which accumulate in cbb3-deficient mutants were nearly restored to normal in a PrrB- CcoP- double mutant. This observation, together with previously published results, suggests that the regulation of the CrtA-catalyzed reaction possesses both transcriptional and posttranscriptional regulatory effectors. We propose that the cbb3 cytochrome oxidase, which by definition can interact with external oxygen, serves to control the activity of the Prr two-component activation system under both aerobic and anaerobic conditions. Although independent from the cbb3 oxidase, the RdxBH proteins are also required for normal functioning of the Prr two-component activation system and are therefore believed to lie between the cbb3 oxidase in this oxygen-sensing, redox signaling pathway and the Prr activation system.

Aerobiosis↗

A redox-responsive regulator of photosynthesis gene expression in the cyanobacterium Synechocystis sp. Strain PCC 6803.

We have identified genes in the unicellular cyanobacterium Synechocystis sp. strain PCC 6803 that are involved with redox control of photosynthesis and pigment-related genes. The genes, rppA (sll0797) and rppB (sll0798), represent a two-component regulatory system that controls the synthesis of photosystem II (PSII) and PSI genes, in addition to photopigment-related genes. rppA (regulator of photosynthesis- and photopigment-related gene expression) and rppB exhibit strong sequence similarity to prokaryotic response regulators and histidine kinases, respectively. In the wild type, the steady-state mRNA levels of PSII reaction center genes increased when the plastoquinone (PQ) pool was oxidized and decreased when the PQ pool was reduced, whereas transcription of the PSI reaction center genes was affected in an opposite fashion. Such results suggested that the redox poise of the PQ pool is critical for regulation of the photosystem reaction center genes. In Delta rppA, an insertion mutation of rppA, the PSII gene transcripts were highly up-regulated relative to the wild type under all redox conditions, whereas transcription of phycobilisome-related genes and PSI genes was decreased. The higher transcription of the psbA gene in Delta rppA was manifest by higher translation of the D1 protein and a concomitant increase in O(2) evolution. The results demonstrated that RppA is a regulator of photosynthesis- and photopigment-related gene expression, is involved in the establishment of the appropriate stoichiometry between the photosystems, and can sense changes in the PQ redox poise.

Amino Acid Sequence↗

Photosynthesis of overwintering evergreen plants.

In this review we focus on photosynthetic behavior of overwintering evergreens with an emphasis on both the acclimative responses of photosynthesis to cold and the winter behavior of photosynthesis in conifers. Photosynthetic acclimation is discussed in terms of the requirement for a balance between the energy absorbed through largely temperature-insensitive photochemical processes and the energy used for temperature-sensitive biochemical processes and growth. Cold acclimation transforms the xanthophyll-mediated nonphotochemical antenna quenching of absorbed light from a short-term dynamic response to a long-term sustained quenching for the whole winter period. This acclimative response helps protect the evergreen foliage from photooxidative damage during the winter when photosynthesis is restricted or prevented by low temperatures. Although the molecular mechanisms behind the sustained winter excitation quenching are largely unknown, it does involve major alterations in the organization and composition of the photosystem II antenna. In addition, photosystem I may play an important role in overwintering evergreens not only by quenching absorbed light photochemically via its support of cyclic electron transport at low temperatures, but also by nonphotochemical quenching of absorbed light irrespective of temperature. The possible role of photosystem II reaction centers in nonphotochemical quenching of absorbed energy in overwintering evergreens is also discussed. Processes like chlororespiration and cyclic electron transport may also be important for maintaining the functional integrity of the photosynthetic apparatus of overwintering evergreens both during periods of thawing in winter and during recovery from winter stress in spring. We suggest that the photosynthetic acclimation responses of overwintering evergreens represent specific evolutionary adaptations for plant species that invest in the long-term maintenance of leaf structure in cold climatic zones as exemplified by the boreal forests of the Northern Hemisphere.

Acclimatization↗

Single-cell C(4) photosynthesis versus the dual-cell (Kranz) paradigm.

The efficiency of photosynthetic carbon assimilation in higher plants faces significant limitations due to the oxygenase activity of the enzyme Rubisco, particularly under warmer temperatures or water stress. A drop in atmospheric CO(2) and rise in O(2) as early as 300 mya provided selective pressure for the evolution of mechanisms to concentrate CO(2) around Rubisco in order to minimize oxygenase activity and the resultant loss of carbon through photorespiration. It is well established that a carbon-concentrating mechanism occurs in some terrestrial plants through the process of C(4) photosynthesis. These plants are characterized as having Kranz-type leaf anatomy, with two structurally and biochemically specialized photosynthetic cell types, mesophyll and bundle sheath, that function coordinately in carbon assimilation. C(4) photosynthesis has evolved independently many times with great diversity in forms of Kranz anatomy, structure of dimorphic chloroplasts, and biochemistry of the C(4) cycle. The most dramatic variants of C(4) terrestrial plants were discovered recently in two species, Bienertia cycloptera and Borszczowia aralocaspica (family Chenopodiaceae); each has novel compartmentation to accomplish C(4) photosynthesis within a single chlorenchyma cell. This review discusses the amazing diversity in C(4) systems, how the essential features of C(4) are accomplished in single-cell versus Kranz-type C(4) plants, and speculates on why single-cell C(4) plants evolved.

Chloroplasts↗

Exploring photosynthesis evolution by comparative analysis of metabolic networks between chloroplasts and photosynthetic bacteria.

BACKGROUND: Chloroplasts descended from cyanobacteria and have a drastically reduced genome following an endosymbiotic event. Many genes of the ancestral cyanobacterial genome have been transferred to the plant nuclear genome by horizontal gene transfer. However, a selective set of metabolism pathways is maintained in chloroplasts using both chloroplast genome encoded and nuclear genome encoded enzymes. As an organelle specialized for carrying out photosynthesis, does the chloroplast metabolic network have properties adapted for higher efficiency of photosynthesis? We compared metabolic network properties of chloroplasts and prokaryotic photosynthetic organisms, mostly cyanobacteria, based on metabolic maps derived from genome data to identify features of chloroplast network properties that are different from cyanobacteria and to analyze possible functional significance of those features. RESULTS: The properties of the entire metabolic network and the sub-network that consists of reactions directly connected to the Calvin Cycle have been analyzed using hypergraph representation. Results showed that the whole metabolic networks in chloroplast and cyanobacteria both possess small-world network properties. Although the number of compounds and reactions in chloroplasts is less than that in cyanobacteria, the chloroplast's metabolic network has longer average path length, a larger diameter, and is Calvin Cycle -centered, indicating an overall less-dense network structure with specific and local high density areas in chloroplasts. Moreover, chloroplast metabolic network exhibits a better modular organization than cyanobacterial ones. Enzymes involved in the same metabolic processes tend to cluster into the same module in chloroplasts. CONCLUSION: In summary, the differences in metabolic network properties may reflect the evolutionary changes during endosymbiosis that led to the improvement of the photosynthesis efficiency in higher plants. Our findings are consistent with the notion that since the light energy absorption, transfer and conversion is highly efficient even in photosynthetic bacteria, the further improvements in photosynthetic efficiency in higher plants may rely on changes in metabolic network properties.

Arabidopsis↗

High-light-induced changes on photosynthesis, pigments, sugars, lipids and antioxidant enzymes in freshwater (Nostoc spongiaeforme) and marine (Phormidium corium) cyanobacteria.

We studied the effects of high-light exposure (500 micromol m(-2) s(-1) of photosynthetic active radiation) on the cyanobacteria Nostoc spongiaeforme Agardh, a fresh-water alga, and Phormidium corium Agardh (Gomont), a marine alga, with respect to photosynthesis, pigments, sugar content, lipid peroxidation, fatty acids composition, antioxidant enzymes activity and DNA. It was seen that the ratio of variable fluorescence (Fv) to maximum fluorescence (Fm), which is indicative of photosynthetic efficiency, decreased because of the light treatment. The damage to photosynthesis occurred in the antenna system and the photosynthetic II reaction center. Photobleaching of photosynthetic pigments was also observed. High-light treatment also resulted in decreased sugar content, which was probably due to the effect on photosynthesis. Peroxidation of membrane lipids, indicating oxidative damage to lipids and a high level of unsaturation in the cell membrane, was also observed. The activity of antioxidant enzyme superoxide dismutase and ascorbate peroxidase was increased, probably as a result of oxidative damage observed in the form of lipid peroxidation. Quantitative decreases in phospholipid and glycolipid levels were also observed. The level of unsaturated fatty acids in total lipids and glycolipids remained unchanged in both species; however, the level of saturated fatty acids decreased, which slightly changed the ratio in favor of unsaturated fatty acids. Degradation of DNA was also observed in both species. There was a transient plateau 2-4 h after exposure to high-light treatment in the Fv/Fm ratio and in levels of phycobilisome pigments, sugars and antioxidant enzymes after an initial decrease 1 h after the treatment. These findings may indicate a period of partial adaptation to high light that is due to the efficiency of protective processes operational in the two species, which subsequently failed after a longer exposure duration of 4-6 h.

Adaptation, Physiological↗

Optimum and sub-optimal temperature effects on stomata and photosynthesis rate of determinate soybeans.

Stomatal frequency, length and width were studied in two determinate type soybean (Glycine max L. Merr.) cultivars at 15, 20 and 25 degrees C on 60th day after emergence. The stomatal frequency on the adaxial leaf surface did not show any consistent trend for the increase of growing temperatures but on the abaxial surface, stomatal frequency significantly decreased for every increase of 5 degrees C. Akishirome had 503, 454 and 379 stomata mm-2 and Akiyoshi had 471, 442 and 384 mm-2 at 15, 20 and 25 degrees C respectively. The stomatal lengths of both surfaces increased toward optimum temperature and were longer in the adaxial surface Maintaining this trend, the lengths varied between 16.3 to 23.4 micrometers on the adaxial surface and from 14.2 to 21.5 micrometers on the abaxial surface. Width of the whole stomatal apparatus at noon time did not show any significant variation due to environmental temperature. Net photosynthesis rate of 4th leaf from top significantly increased in higher temperatures in both cultivars and showed similar trend at 32nd day after emergence and on 62nd day after emergence. Stomatal conductance increased and dark respiration decreased with increasing temperature. Plants grown in 25 degrees C were transferred to 15, 20 and 30 degrees C temperature chambers. On the 30th day after emergence, 4 hours of treatment resulted similar significant effects on net photosynthesis (ranging between 12.55 and 31.37 micromoles CO2 m-2 s-1 in Akishirome, and between 16.26 to 34.53 micrometers CO2 m-2 s-1 in Akiyoshi). 72 hours of similar treatments at 60 day after emergence also produced identical results. Therefore, higher temperature increased stomatal size but decreased its frequency, and increased net photosynthesis and stomatal conductance.

Cell Respiration↗

Synergistic effects of a photooxidized polycyclic aromatic hydrocarbon and copper on photosynthesis and plant growth: evidence that in vivo formation of reactive oxygen species is a mechanism of copper toxicity.

Heavy metals and polycyclic aromatic hydrocarbons (PAHs) are often cocontaminants in industrialized environments, yet little is known about either the extent or mechanisms of their cotoxicity. To address this shortfall, the combined effects of an oxygenated PAH, 1,2-dihydroxyanthraquinone (1,2-dhATQ), and a heavy metal, Cu2+, on photosynthesis and growth of the duckweed (Lemna gibba) were evaluated. Using assays of chlorophyll a fluorescence and photosystem I activity, 1,2-dhATQ inhibited electron transport at the cytochrome b6/f complex. Conversely, Cu2+ alone (at low concentrations) had little effect on photosynthesis. When Cu2+ was combined with 1,2-dhATQ, an increase in transient and steady-state chlorophyll a fluorescence quenching occurred relative to 1,2-dhATQ alone. Treatment of isolated thylakoid membranes with 1,2-dhATQ inhibited whole-chain linear electron transport, measured as O2 consumption using methyl viologen as the electron acceptor. However, Cu2+ plus 1,2-dhATQ resulted in active O2 consumption with or without methyl viologen as an electron acceptor. From these data, we conclude that 1,2-dhATQ renders the plastoquinone pool to a highly reduced state by inhibiting at cytochrome b6/f. Then, Cu2+ is able to mediate the transfer of electrons from reduced plastoquinone to O2, forming reactive oxygen species. At the whole-organism level, when Cu2+ and 1,2-dhATQ were mixed at concentrations that resulted in the above-mentioned impacts on photosynthesis, synergistic inhibition of plant growth was observed. This suggests a catalytic mechanism of toxicity for redox active metals, a process that could be instrumental in explaining their impacts at low concentrations.

Anthraquinones↗

Acetabulaira mediterranea: circadian rhythms of photosynthesis and associated changes in molecular structure of the thylakoid membranes.

Acetabularia mediterranea algae, grown in three different light-dark regimes, were frozen in liquid nitrogen at c.t.(1) 0 and c.t. 6 and a record made of 77 degrees K fluorescence emission spectra of their chloroplasts. Algae grown under LD cycles exhibited a clear circadian rhythm of oxygen production. The low temperature fluorescence emission spectrum at c.t.0 was different from that at c.t.6 and this difference was increased by submitting the algae to successive "freeze-thaw" treatment. Similar results were obtained in DD, and the photosynthesis rhythm remained fully expressed. Algae grown in LL, where no rhythm of photosynthesis could be detected in the samples because there is a great individual variability in period lenght under these conditions, exhibited a similar difference in their low temperature flourescence emission spectra between c.t.0 and c.t.6. We conclude that the circadian rhythm in low-temperature fluorescence emission of the chloroplasts in Acetabularia is related to the circadian rhythm in photosynthesis.

Acetabularia↗

Excitation energy transfer from phycobilisomes to photosystems: a phenomenon associated with the temporal separation of photosynthesis and nitrogen fixation in a cyanobacterium, Plectonema boryanum.

Plectonema boryanum shows temporal separation of photosynthesis and nitrogen fixation under diazotrophic conditions. Low temperature fluorescence studies have shown that in vivo the nitrogen fixing and photosynthesizing cells are adapted to 'state 2' and 'state 1', respectively. During nitrogen fixation phycobilisomes seem to transfer excitation energy to photosystem I whereas during oxygenic photosynthesis the energy is transferred to photosystem II. The state 2 adapted N-phase cells failed to undergo transition to state 1 while P-phase cells exhibited state 1 to state 2 transition. The nitrogen fixing cells showed a decreased level ofpsbC transcript, lack of CP47 in thylakoid membrane, and presence of the F685 peak but absence of the F695 peak in 77 K fluorescence spectra. These results suggest that the metabolic and molecular changes associated with nitrogen fixation may favor direct energy transfer from the phycobilisomes to photosystem I. This should help the organism to achieve low photosystem II and high photosystem I activity to set temporal separation of nitrogen fixation and photosynthesis for photoautotrophic growth under diazotrophic conditions.

Bacterial Proteins↗

Compartmentation of photosynthesis in cells and tissues of C(4) plants.

Critical to defining photosynthesis in C(4) plants is understanding the intercellular and intracellular compartmentation of enzymes between mesophyll and bundle sheath cells in the leaf. This includes enzymes of the C(4) cycle (including three subtypes), the C(3) pathway and photorespiration. The current state of knowledge of this compartmentation is a consequence of the development and application of different techniques over the past three decades. Initial studies led to some alternative hypotheses on the mechanism of C(4) photosynthesis, and some controversy over the compartmentation of enzymes. The development of methods for separating mesophyll and bundle sheath cells provided convincing evidence on intercellular compartmentation of the key components of the C(4) pathway. Studies on the intracellular compartmentation of enzymes between organelles and the cytosol were facilitated by the isolation of mesophyll and bundle sheath protoplasts, which can be fractionated gently while maintaining organelle integrity. Now, the ability to determine localization of photosynthetic enzymes conclusively, through in situ immunolocalization by confocal light microscopy and transmission electron microscopy, is providing further insight into the mechanism of C(4) photosynthesis and its evolution. Currently, immunological, ultrastructural and cytochemical studies are revealing relationships between anatomical arrangements and photosynthetic mechanisms which are probably related to environmental factors associated with evolution of these plants. This includes interesting variations in the C(4) syndrome in leaves and cotyledons of species in the tribe Salsoleae of the family Chenopodiaceae, in relation to evolution and ecology. Thus, analysis of structure-function relationships using modern techniques is a very powerful approach to understanding evolution and regulation of the photosynthetic carbon reduction mechanisms.

Carbon Dioxide↗

Limitations to photosynthesis of lettuce grown under tropical conditions: alleviation by root-zone cooling.

Aerial parts of lettuce plants were grown under natural tropical fluctuating ambient temperatures, but with their roots exposed to two different root-zone temperatures (RZTs): a constant 20 degrees C-RZT and a fluctuating ambient (A-) RZT from 23-40 degrees C. Plants grown at A-RZT showed lower photosynthetic CO2 assimilation (A), stomatal conductance (gs), midday leaf relative water content (RWC), and chlorophyll fluorescence ratio Fv/Fm than 20 degrees C-RZT plants on both sunny and cloudy days. Substantial midday depression of A and g(s) occurred on both sunny and cloudy days in both RZT treatments, although Fv/Fm did not vary diurnally on cloudy days. Reciprocal temperature transfer experiments investigated the occurrence and possible causes of stomatal and non-stomatal limitations of photosynthesis. For both temperature transfers, light-saturated stomatal conductance (gs sat) and photosynthetic CO2 assimilation (A(sat)) were highly correlated with each other and with midday RWC, suggesting that A was limited by water stress-mediated stomatal closure. However, prolonged growth at A-RZT reduced light- and CO2-saturated photosynthetic O2 evolution (Pmax), indicating non-stomatal limitation of photosynthesis. Tight temporal coupling of leaf nitrogen content and P(max) during both temperature transfers suggested that decreased nutrient status caused this non-stomatal limitation of photosynthesis.

Carbon Dioxide↗