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# Photosynthesis Research - Current Research Articles

## Current research articles: Photosynthesis

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... is an international journal open to papers of merit dealing with both basic and applied aspects of photosynthesis.

# Current articles of the journal:

Posted on 1 May 2015 | 2:00 am

## Characterization of a Synechocystis sp. PCC 6803 double mutant lacking the CyanoP and Ycf48 proteins of Photosystem II

### Abstract

Homologs of the Photosystem II (PS II) subunit PsbP are found in plants, algae, and cyanobacteria. In higher plants, PsbP is associated with mature PS II centers, but in cyanobacteria, the homologous CyanoP protein appears sub-stoichiometric to PS II. We have investigated the role of CyanoP by characterizing knockout mutants of the cyanobacterium Synechocystis sp. PCC 6803. Removal of CyanoP resulted in changes to phycobilisome coupling and energy transfer to PS II, but the function of PS II itself remained similar to wild type. We therefore investigated the hypothesis that CyanoP is involved in the biogenesis or repair of PS II by creating a double mutant lacking both CyanoP and the PS II assembly factor Ycf48. This strain exhibited an additive reduction in the amplitude of variable chlorophyll a fluorescence induction relative to either of the single mutants but displayed increased oxygen evolution, slight increases in PS II monomer and dimer levels, and a reduction in accumulation of an early PS II assembly complex containing CP47, compared to the ?Ycf48 strain.

Posted on 1 May 2015 | 2:00 am

## Role of electron transport chain of chloroplasts in oxidative burst of interaction between Erwinia amylovora and host cells

### Abstract

Erwinia amylovora is a necrogenic bacterium, causing the fire blight disease on many rosaceous plants. Triggering oxidative burst by E. amylovora is a key response by which host plants try to restrain pathogen spread. Electron transport chain (ETC) of chloroplasts is known as an inducible source of reactive oxygen species generation in various stresses. This research was performed to assess the role of this ETC in E. amylovora–host interaction using several inhibitors of this chain in susceptible and resistant apple and pear genotypes. All ETC inhibitors delayed appearance of disease necrosis, but the effects of methyl viologen, glutaraldehyde, and DCMU were more significant. In the absence of inhibitors, resistant genotypes showed an earlier and severe H2O2 generation and early suppression of redox dependent, psbA gene. The effects of inhibitors were corresponding to the redox potential of ETC inhibitory sites. In addition, delayed necrosis appearance was associated with the decreased disease severity and delayed H2O2 generation. These results provide evidences for the involvement of this ETC in host oxidative burst and suggest that chloroplast ETC has significant role in E. amylovora–host interaction.

Posted on 1 May 2015 | 2:00 am

## Violaxanthin de-epoxidase disulphides and their role in activity and thermal stability

### Abstract

Violaxanthin de-epoxidase (VDE) catalyses the conversion of violaxanthin to zeaxanthin at the lumen side of the thylakoids during exposure to intense light. VDE consists of a cysteine-rich N-terminal domain, a lipocalin-like domain and a negatively charged C-terminal domain. That the cysteines are important for the activity of VDE is well known, but in what way is less understood. In this study, wild-type spinach VDE was expressed in E. coli as inclusion bodies, refolded and purified to give a highly active and homogenous preparation. The metal content (Fe, Cu, Ni, Mn, Co and Zn) was lower than 1 mol% excluding a metal-binding function of the cysteines. To investigate which of the 13 cysteines that could be important for the function of VDE, we constructed mutants where the cysteines were replaced by serines, one by one. For 12 out of 13 mutants the activity dropped by more than 99.9 %. A quantification of free cysteines showed that only the most N-terminal of these cysteines was in reduced form in the native VDE. A disulphide pattern in VDE of C9–C27, C14–C21, C33–C50, C37–C46, C65–C72 and C118–C284 was obtained after digestion of VDE with thermolysin followed by mass spectroscopy analysis of reduced versus non-reduced samples. The residual activity found for the mutants showed a variation that was consistent with the results obtained from mass spectroscopy. Reduction of the disulphides resulted in loss of a rigid structure and a decrease in thermal stability of 15 °C.

Posted on 1 May 2015 | 2:00 am

## Seasonal changes in the content of dehydrins in mesophyll cells of common pine needles

### Abstract

The appearance of dehydrins (DHNs) in cells is required for the development of cold resistance. DHNs are therefore considered specific markers of cold resistance by some authors. DHNs accumulate in plants concomitantly with a reduction of intracellular water content, and presumably protect membranes and proteins from damage caused by moisture loss. DHN content in pine needles increases in spring and autumn when moisture availability and temperatures are most unfavorable. The present work is focused on seasonal changes in DHN content in various mesophyll-cell compartments of pine (Pinus sylvestris L.) needles in association with changes in environmental factors. In spring, the number of thylakoid membranes per granum was lower than in summer and autumn. An increase in needle content of DHNs with approximate masses of 76, 73, 72, 35, and 17 kD in spring and autumn, associated with needle dehydration during this period, is shown here. The largest increase in DHN content was observed in spring, with the highest amount of DHNs presented in chloroplast membrane system including grana thylakoids, stromal thylakoids, and the two chloroplast envelope membranes and in cell walls. In the autumn, most DHNs were localized in chloroplasts and mitochondria.

Posted on 1 May 2015 | 2:00 am

## How will climate change influence grapevine cv. Tempranillo photosynthesis under different soil textures?

### Abstract

While photosynthetic responses to elevated CO2, elevated temperature, or water availability have previously been reported for grapevine as responses to single stress factors, reports on the combined effect of multiple stress factors are scarce. In the present work, we evaluated effects of simulated climate change [CC; 700 ppm CO2, 28/18 °C, and 33/53 % relative humidity (RH), day/night] versus current conditions (375 ppm CO2, 24/14 °C, and 45/65 % RH), water availability (well-irrigated vs. water deficit), and different types of soil textures (41, 19, and 8 % of soil clay contents) on grapevine (Vitis vinifera L. cv. Tempranillo) photosynthesis. Plants were grown using the fruit-bearing cutting model. CC increased the photosynthetic activity of grapevine plants grown under well-watered conditions, but such beneficial effects of elevated CO2, elevated temperature, and low RH were abolished by water deficit. Under water-deficit conditions, plants subjected to CC conditions had similar photosynthetic rates as those grown under current conditions, despite their higher sub-stomatal CO2 concentrations. As expected, water deficit reduced photosynthetic activity in association with inducing stomatal closure that prevents water loss. Evidence for photosynthetic downregulation under elevated CO2 was observed, with decreases in photosynthetic capacity and leaf N content and increases in the C/N ratio in plants subjected to CC conditions. Soil texture had no marked effects on photosynthesis and did not modify the photosynthetic response to CC and water-deficit conditions. However, in mature well-irrigated plants grown in the soils with the highest sand content, an important decrease in stomatal conductance was observed as well as a slight decrease in the utilization of absorbed light in photosynthetic electron transport (measured as photochemical quenching), possibly related to a low water-retention capacity of these soils even under well-watered conditions.

Posted on 1 May 2015 | 2:00 am

## Light-harvesting regulation from leaf to molecule with the emphasis on rapid changes in antenna size

### Abstract

In the sunlight-fluctuating environment, plants often encounter both light-deficiency and light-excess cases. Therefore, regulation of light harvesting is absolutely essential for photosynthesis in order to maximize light utilization at low light and avoid photodamage of the photosynthetic apparatus at high light. Plants have developed a series of strategies of light-harvesting regulation during evolution. These strategies include rapid responses such as leaf movement and chloroplast movement, state transitions, and reversible dissociation of some light-harvesting complex of the photosystem II (LHCIIs) from PSII core complexes, and slow acclimation strategies such as changes in the protein abundance of light-harvesting antenna and modifications of leaf morphology, structure, and compositions. This review discusses successively these strategies and focuses on the rapid change in antenna size, namely reversible dissociation of some peripheral light-harvesting antennas (LHCIIs) from PSII core complex. It is involved in protective role and species dependence of the dissociation, differences between the dissociation and state transitions, relationship between the dissociation and thylakoid protein phosphorylation, and possible mechanism for thermal dissipation by the dissociated LHCIIs.

Posted on 1 May 2015 | 2:00 am

## Carotenoid–chlorophyll coupling and fluorescence quenching in aggregated minor PSII proteins CP24 and CP29

### Abstract

It is known that aggregation of isolated light-harvesting complex II (LHCII) in solution results in high fluorescence quenching, reduced chlorophyll fluorescence lifetime, and increased electronic coupling of carotenoid (Car) S1 and chlorophyll (Chl) Qy states, as determined by two-photon studies. It has been suggested that this behavior of aggregated LHCII mimics aspects of non-photochemical quenching processes of higher plants and algae. However, several studies proposed that the minor photosystem II proteins CP24 and CP29 also play a significant role in regulation of photosynthesis. Therefore, we use a simple protocol that allows gradual aggregation also of CP24 and CP29. Similarly, as observed for LHCII, aggregation of CP24 and CP29 also leads to increasing fluorescence quenching and increasing electronic Car S1–Chl Qy coupling. Furthermore, a direct comparison of the three proteins revealed a significant higher electronic coupling in the two minor proteins already in the absence of any aggregation. These differences become even more prominent upon aggregation. A red-shift of the Qy absorption band known from LHCII aggregation was also observed for CP29 but not for CP24. We discuss possible implications of these results for the role of CP24 and CP29 as potential valves for excess excitation energy in the regulation of photosynthetic light harvesting.

Posted on 1 May 2015 | 2:00 am

## High prevalence of diffusive uptake of CO 2 by macroalgae in a temperate subtidal ecosystem

### Abstract

Productivity of most macroalgae is not currently considered limited by dissolved inorganic carbon (DIC), as the majority of species have CO2-concentrating mechanisms (CCM) allowing the active uptake of DIC. The alternative, diffusive uptake of CO2 (non-CCM), is considered rare (0–9 % of all macroalgal cover in a given ecosystem), and identifying species without CCMs is important in understanding factors controlling inorganic carbon use by eukaryotic algae. CCM activity has higher energetic requirements than diffusive CO2 uptake, therefore when light is low, CCM activity is reduced in favour of diffusive CO2 uptake. We hypothesized that the proportional cover of macroalgae without CCMs (red and green macroalgae) would be low (<10 %) across four sites in Tasmania, southern Australia at two depths (4–5 and 12–14 m); the proportion of species lacking CCMs would increase with decreasing depth; the ?13C values of macroalgae with CCMs would be more depleted with depth. We found the proportion of non-CCM species ranged from 0 to 90 % and included species from all three macroalgal phyla: 81 % of red (59 species), 14 % of brown (three species) and 29 % of green macroalgae (two species). The proportion of non-CCM species increased with depth at three of four sites. 35 % of species tested had significantly depleted ?13C values at deeper depths. Non-CCM macroalgae are more abundant in some temperate reefs than previously thought. If ocean acidification benefits non-CCM species, the ramifications for subtidal macroalgal assemblages could be larger than previously considered.

Posted on 1 May 2015 | 2:00 am

## Effects of inefficient transcription termination of rbcL on the expression of accD in plastids of Arabidopsis thaliana

### Abstract

The plastid accD gene encodes one subunit of a multimeric acetyl-CoA carboxylase that is required for fatty acid biosynthesis. In Arabidopsis thaliana, the accD gene is transcribed by the nuclear-encoded phage-type RNA polymerase, and the accumulation of accD transcripts is subjected to a dynamic pattern during chloroplast development. However, the mechanisms underlying the regulation of accD expression remain unknown. Here, we showed that the inefficient transcription termination of rbcL due to the absence of RHON1 impaired the developmental profile of accD, resulting in the constitutive expression of accD during chloroplast development. Moreover, the accumulation of accD transcripts accordingly resulted in an increase in accD protein levels, suggesting that transcript abundance is critical for accD gene production. Our study demonstrates that the interplay between accD and upstream rbcL regulates the expression of accD and highlights the significance of transcriptional regulation in plastid gene expression in higher plants.

Posted on 24 May 2015 | 2:00 am

## Multiple regulatory mechanisms in the chloroplast of green algae: relation to hydrogen production

### Abstract

A complex regulatory network in the chloroplast of green algae provides an efficient tool for maintenance of energy and redox balance in the cell under aerobic and anaerobic conditions. In this review, we discuss the structural and functional organizations of electron transport pathways in the chloroplast, and regulation of photosynthesis in the green microalga Chlamydomonas reinhardtii. The focus is on the regulatory mechanisms induced in response to nutrient deficiency stress and anoxia and especially on the role of a hydrogenase-mediated reaction in adaptation to highly reducing conditions and ATP deficiency in the cell.

Posted on 19 May 2015 | 2:00 am

## Correlation between pH dependence of O 2 evolution and sensitivity of Mn cations in the oxygen-evolving complex to exogenous reductants

### Abstract

Effects of pH, Ca2+, and Cl? ions on the extraction of Mn cations from oxygen-evolving complex (OEC) in Ca-depleted photosystem II (PSII(-Ca)) by exogenous reductants hydroquinone (H2Q) and H2O2 were studied. Two of 4 Mn cations are released by H2Q and H2O2 at pHs 5.7, 6.5, and 7.5, and their extraction does not depend on the presence of Ca2+ and Cl? ions. One of Mn cations (“resistant” Mn cation) cannot be extracted by H2Q and H2O2 at any pH. Extraction of 4th Mn ion (“flexible” Mn cation) is sensitive to pH, Ca2+, and Cl?. This Mn cation is released by reductants at pH 6.5 but not at pHs 5.7 and 7.5. A pH dependence curve of the oxygen-evolving activity in PSII(-Ca) membranes (in the presence of exogenous Ca2+) has a bell-shaped form with the maximum at pH 6.5. Thus, the increase in the resistance of flexible Mn cation in OEC to the action of reductants at acidic and alkaline pHs coincides with the decrease in oxygen evolution activity at these pHs. Exogenous Ca2+ protects the extraction of flexible Mn cation at pH 6.5. High concentration of Cl? anions (100 mM) shifts the pH optimum of oxygen evolution to alkaline region (around pH 7.5), while the pH of flexible Mn extraction is also shifted to alkaline pH. This result suggests that flexible Mn cation plays a key role in the water-splitting reaction. The obtained results also demonstrate that only one Mn cation in Mn4 cluster is under strong control of calcium. The change in the flexible Mn cation resistance to exogenous reductants in the presence of Ca2+ suggests that Ca2+ can control the redox potential of this cation.

Posted on 15 May 2015 | 2:00 am

## Recent breakthroughs in the biology of astaxanthin accumulation by microalgal cell

### Abstract

Massive accumulation of the secondary ketokarotenoid astaxanthin is a characteristic stress response of certain microalgal species with Haematococcus pluvialis as an illustrious example. The carotenogenic response confers these organisms a remarkable ability to survive in extremely unfavorable environments and makes them the richest source of natural astaxanthin. Exerting a plethora of beneficial effects on human and animal health, astaxanthin is among the most important bioproducts from microalgae. Though our understanding of astaxanthin biosynthesis, induction, and regulation is far from complete, this gap is filling rapidly with new knowledge generated predominantly by application of advanced “omics” approaches. This review focuses on the most recent progress in the biology of astaxanthin accumulation in microalgae including the genomic, proteomic, and metabolomics insights into the induction and regulation of secondary carotenogenesis and its role in stress tolerance of the photosynthetic microorganisms. Special attention is paid to the coupling of the carotenoid and lipid biosynthesis as well as deposition of astaxanthin in the algal cell. The place of the carotenogenic response among the stress tolerance mechanisms is revisited, and possible implications of the new findings for biotechnological production of astaxanthin from microalgae are considered. The potential use of the carotenogenic microalgae as a source not only of value-added carotenoids, but also of biofuel precursors is discussed.

Posted on 15 May 2015 | 2:00 am

## Photochemical properties in flag leaves of a super-high-yielding hybrid rice and a traditional hybrid rice ( Oryza sativa L.) probed by chlorophyll a fluorescence transient

### Abstract

Chlorophyll a fluorescence of flag leaves in a super-high-yielding hybrid rice (Oryza sativa L.) LYPJ, and a traditional hybrid rice SY63 cultivar with lower grain yield, which were grown in the field, were investigated from emergence through senescence of flag leaves. As the flag leaf matured, there was an increasing trend in photosynthetic parameters such as quantum efficiency of primary photochemistry ( $$\varphi$$ Po) and efficiency of electron transport from PS II to PS I (? Eo). The overall photosynthetic performance index (PIABS) was significantly higher in the high-yielding LYPJ compared to SY63 during the entire reproductive stage of the plant, the same to MDA content. However, $$\varphi$$ Po(=F V/F M), an indicator of the primary photochemistry of the flag leaf, did not display significant changes with leaf age and was not significantly different between the two cultivars, suggesting that PIABS is a more sensitive parameter than $$\varphi$$ Po (=F V/F M) during leaf age for distinguishing between cultivars differing in yield.

Posted on 14 May 2015 | 2:00 am

## Matthias Rögner (ed): Biohydrogen

Posted on 13 May 2015 | 2:00 am

## Derek Bendall (1930–2014)

### Abstract

Derek Bendall carried out pioneering work on photosynthetic electron transport, particularly on protein–protein interactions, cytochromes, and cyclic electron transport, as well as on other topics including the biochemistry of tea. He was a keen musician and a gifted gardener, a devoted family man, and a delightful colleague and friend. The bioenergetics community, especially those working on photosynthesis, will miss him sorely.

Posted on 13 May 2015 | 2:00 am

## Organization of chlorophyll biosynthesis and insertion of chlorophyll into the chlorophyll-binding proteins in chloroplasts

### Abstract

Oxygenic photosynthesis requires chlorophyll (Chl) for the absorption of light energy, and charge separation in the reaction center of photosystem I and II, to feed electrons into the photosynthetic electron transfer chain. Chl is bound to different Chl-binding proteins assembled in the core complexes of the two photosystems and their peripheral light-harvesting antenna complexes. The structure of the photosynthetic protein complexes has been elucidated, but mechanisms of their biogenesis are in most instances unknown. These processes involve not only the assembly of interacting proteins, but also the functional integration of pigments and other cofactors. As a precondition for the association of Chl with the Chl-binding proteins in both photosystems, the synthesis of the apoproteins is synchronized with Chl biosynthesis. This review aims to summarize the present knowledge on the posttranslational organization of Chl biosynthesis and current attempts to envision the proceedings of the successive synthesis and integration of Chl into Chl-binding proteins in the thylakoid membrane. Potential auxiliary factors, contributing to the control and organization of Chl biosynthesis and the association of Chl with the Chl-binding proteins during their integration into photosynthetic complexes, are discussed in this review.

Posted on 9 May 2015 | 2:00 am

## Sll0751 and Sll1041 are involved in acid stress tolerance in Synechocystis sp. PCC 6803

### Abstract

The ATP-binding cassette (ABC) transporter is a multi-subunit membrane protein complex involved in lipid transport and acid stress tolerance in the cyanobacterium Synechocystis sp. PCC 6803. This organism has two sets of three ABC transporter subunits: Slr1045 and Slr1344, Sll0751 and Sll1002, and Sll1001 and Sll1041. We previously found that Slr1045 is essential for survival under acid stress condition (Tahara et al. 2012). In the present study, we examined the participation of other ABC transporter subunits in acid stress tolerance using a deletion mutant series of Synechocystis sp. PCC 6803. Although Slr1344 is highly homologous to Slr1045, ?slr1344 cells were not susceptible to acid stress. ?sll0751 and ?sll1041 cells displayed acid stress sensitivity, whereas ?sll1001/sll1002 double mutant cells grew normally. Under high- and low-temperature stress conditions, the growth rate of ?slr1344 and ?sll1001/sll1002 cells did not differ from WT cells, whereas ?sll0751 and ?sll1041 cells showed significant growth retardation, as previously observed in ?slr1045 cells. Moreover, nile red staining showed more lipid accumulation in ?slr1045, ?sll0751, and ?sll1041 cells than in WT cells. These results suggest that Slr1045, Sll0751, and Sll1041 function together as a lipid transport complex in Synechocystis sp. PCC 6803 and are essential for growth under various stresses.

Posted on 8 May 2015 | 2:00 am

## Relevance of nutrient media composition for hydrogen production in Chlamydomonas

### Abstract

Microalgae are capable of biological H2 photoproduction from water, solar energy, and a variety of organic substrates. Acclimation responses to different nutrient regimes finely control photosynthetic activity and can influence H2 production. Hence, nutrient stresses are an interesting scenario to study H2 production in photosynthetic organisms. In this review, we mainly focus on the H2-production mechanisms in Chlamydomonas reinhardtii and the physiological relevance of the nutrient media composition when producing H2.

Posted on 8 May 2015 | 2:00 am

## Electronic coupling of the phycobilisome with the orange carotenoid protein and fluorescence quenching

### Abstract

Using computational modeling and known 3D structure of proteins, we arrived at a rational spatial model of the orange carotenoid protein (OCP) and phycobilisome (PBS) interaction in the non-photochemical fluorescence quenching. The site of interaction is formed by the central cavity of the OCP monomer in the capacity of a keyhole to the characteristic external tip of the phycobilin-containing domain (PB) and folded loop of the core-membrane linker LCM within the PBS core. The same central protein cavity was shown to be also the site of the OCP and fluorescence recovery protein (FRP) interaction. The revealed geometry of the OCP to the PBLCM attachment is believed to be the most advantageous one as the LCM, being the major terminal PBS fluorescence emitter, gathers, before quenching by OCP, the energy from most other phycobilin chromophores of the PBS. The distance between centers of mass of the OCP carotenoid 3?-hydroxyechinenone (hECN) and the adjacent phycobilin chromophore of the PBLCM was determined to be 24.7 Å. Under the dipole–dipole approximation, from the point of view of the determined mutual orientation and the values of the transition dipole moments and spectral characteristics of interacting chromophores, the time of the direct energy transfer from the phycobilin of PBLCM to the S1 excited state of hECN was semiempirically calculated to be 36 ps, which corresponds to the known experimental data and implies the OCP is a very efficient energy quencher. The complete scheme of OCP and PBS interaction that includes participation of the FRP is proposed.

Posted on 7 May 2015 | 2:00 am

## Andrew A. Benson, 1917–2015

### Abstract

On January 16, 2015, Professor Andrew Alm Benson, one of the leading plant biochemists of the twentieth century, died in La Jolla, California, at the age of 97; he was born on September 24, 1917. Benson was known especially for his pioneering studies on photosynthesis (CO2 assimilation, carbon reduction cycle) and plant lipids (phospholipid phosphatidyl glycerol; and the sulfolipid, sulfoquinovosyl diglyceride). A photograph of Benson is shown in Fig. 1.

Fig. 1
Photograph of Andrew A. Benson. Source: Annual Review of Plant Biology, Vol. 53, 2002, published with permission