Photosynthesis Research (v.131, #3)
Frederick Loring Crane (1925–2016): Discoverer of coenzyme Q10 and rediscoverer of plastoquinone by Richard A. Dilley (237-239).
Drought stress memory in the photosynthetic mechanisms of an invasive CAM species, Aptenia cordifolia by Marta Pintó-Marijuan; Alba Cotado; Eva Fleta-Soriano; Sergi Munné-Bosch (241-253).
Plants are known for their high capacity to acclimatise to fluctuating environmental conditions. A wide range of environmental conditions can lead to suboptimal physiological efficiency. However, recent studies have shown that plants can withstand repeated periods of stress. To find out how they do it, we studied photosynthetic adjustments to repeated water stress in Aptenia cordifolia: a facultative, invasive CAM species. Plants were subjected to three cycles of water deficit, and photosynthetic parameters and chloroplast antioxidants were quantified to gain an understanding of the mechanisms by which they cope with repeated stress periods. Significant modification of the photosystems’ antenna and reaction centres was observed in plants subjected to previous water stress cycles, and this led to higher PSII efficiency than in plants challenged with drought for the first time. These findings underline the biological significance of stress memory and show how plants can adjust their photosynthetic apparatus to fluctuating environmental conditions and thus optimise photosynthesis and photoprotection under drought conditions.
Keywords: Abiotic stress; Chloroplast pigments; Crassulacean acid metabolism; Photoprotection; Photosynthesis; Vitamin E
Modular antenna of photosystem I in secondary plastids of red algal origin: a Nannochloropsis oceanica case study by David Bína; Zdenko Gardian; Miroslava Herbstová; Radek Litvín (255-266).
Photosystem I (PSI) is a multi-subunit integral pigment–protein complex that performs light-driven electron transfer from plastocyanin to ferredoxin in the thylakoid membrane of oxygenic photoautotrophs. In order to achieve the optimal photosynthetic performance under ambient irradiance, the absorption cross section of PSI is extended by means of peripheral antenna complexes. In eukaryotes, this role is played mostly by the pigment–protein complexes of the LHC family. The structure of the PSI-antenna supercomplexes has been relatively well understood in organisms harboring the primary plastid: red algae, green algae and plants. The secondary endosymbiotic algae, despite their major ecological importance, have so far received less attention. Here we report a detailed structural analysis of the antenna-PSI association in the stramenopile alga Nannochloropsis oceanica (Eustigmatophyceae). Several types of PSI-antenna assemblies are identified allowing for identification of antenna docking sites on the PSI core. Instances of departure of the stramenopile system from the red algal model of PSI-Lhcr structure are recorded, and evolutionary implications of these observations are discussed.
Keywords: Photosystem I; Electron microscopy; Light-harvesting complex; Nannochloropsis ; Stramenopila
Synechocystis sp. PCC 6803 CruA (sll0147) encodes lycopene cyclase and requires bound chlorophyll a for activity by Wei Xiong; Gaozhong Shen; Donald A. Bryant (267-280).
The genome of the model cyanobacterium, Synechococcus sp. PCC 7002, encodes two paralogs of CruA-type lycopene cyclases, SynPCC7002_A2153 and SynPCC7002_A0043, which are denoted cruA and cruP, respectively. Unlike the wild-type strain, a cruA deletion mutant is light-sensitive, grows slowly, and accumulates lycopene, γ-carotene, and 1-OH-lycopene; however, this strain still produces β-carotene and other carotenoids derived from it. Expression of cruA from Synechocystis sp. PCC 6803 (cruA 6803) in Escherichia coli strains that synthesize either lycopene or γ-carotene did not lead to the synthesis of either γ-carotene or β-carotene, respectively. However, expression of this orthologous cruA 6803 gene (sll0147) in the Synechococcus sp. PCC 7002 cruA deletion mutant produced strains with phenotypic properties identical to the wild type. CruA6803 was purified from Synechococcus sp. PCC 7002 by affinity chromatography, and the purified protein was pale yellow-green due to the presence of bound chlorophyll (Chl) a and β-carotene. Native polyacrylamide gel electrophoresis of the partly purified protein in the presence of lithium dodecylsulfate at 4 °C confirmed that the protein was yellow-green in color. When purified CruA6803 was assayed in vitro with either lycopene or γ-carotene as substrate, β-carotene was synthesized. These data establish that CruA6803 is a lycopene cyclase and that it requires a bound Chl a molecule for activity. Possible binding sites for Chl a and the potential regulatory role of the Chl a in coordination of Chl and carotenoid biosynthesis are discussed.
Keywords: Carotenoids; Carotenogenesis; Lycopene cyclase; β-Carotene; Chlorophyll; Photosynthesis
Effects of co-overexpression of the genes of Rubisco and transketolase on photosynthesis in rice by Yuji Suzuki; Eri Kondo; Amane Makino (281-289).
Metabolome analyses have indicated an accumulation of sedoheptulose 7-phosphate in transgenic rice plants with overproduction of Rubisco (Suzuki et al. in Plant Cell Environ 35:1369–1379, 2012. doi: 10.1111/j.1365-3040.2012.02494.x ). Since Rubisco overproduction did not quantitatively enhance photosynthesis even under CO2-limited conditions, it is suspected that such an accumulation of sedoheptulose 7-phosphate hampers the improvement of photosynthetic capacity. In the present study, the gene of transketolase, which is involved in the metabolism of sedoheptulose 7-phosphate, was co-overexpressed with the Rubisco small subunit gene in rice. Rubisco and transketolase were successfully overproduced in comparison with those in wild-type plants by 35–53 and 39–84 %, respectively. These changes in the amounts of the proteins were associated with those of the mRNA levels. However, the rate of CO2 assimilation under high irradiance and different [CO2] did not differ between co-overexpressed plants and wild-type plants. Thus, co-overproduction of Rubisco and transketolase did not improve photosynthesis in rice. Transketolase was probably not a limiting factor of photosynthesis as overproduction of transketolase alone by 80–94 % did not affect photosynthesis.
Keywords: Rubisco; Transketolase; Overproduction; CO2 assimilation; Rice
New insights into the photochemistry of carotenoid spheroidenone in light-harvesting complex 2 from the purple bacterium Rhodobacter sphaeroides by Dariusz M. Niedzwiedzki; Preston L. Dilbeck; Qun Tang; Elizabeth C. Martin; David F. Bocian; C. Neil Hunter; Dewey Holten (291-304).
Light-harvesting complex 2 (LH2) from the semi-aerobically grown purple phototrophic bacterium Rhodobacter sphaeroides was studied using optical (static and time-resolved) and resonance Raman spectroscopies. This antenna complex comprises bacteriochlorophyll (BChl) a and the carotenoid spheroidenone, a ketolated derivative of spheroidene. The results indicate that the spheroidenone-LH2 complex contains two spectral forms of the carotenoid: (1) a minor, “blue” form with an S2 (11B u + ) spectral origin band at 522 nm, shifted from the position in organic media simply by the high polarizability of the binding site, and (2) the major, “red” form with the origin band at 562 nm that is associated with a pool of pigments that more strongly interact with protein residues, most likely via hydrogen bonding. Application of targeted modeling of excited-state decay pathways after carotenoid excitation suggests that the high (92%) carotenoid-to-BChl energy transfer efficiency in this LH2 system, relative to LH2 complexes binding carotenoids with comparable double-bond conjugation lengths, derives mainly from resonance energy transfer from spheroidenone S2 (11B u + ) state to BChl a via the Qx state of the latter, accounting for 60% of the total transfer. The elevated S2 (11B u + ) → Qx transfer efficiency is apparently associated with substantially decreased energy gap (increased spectral overlap) between the virtual S2 (11B u + ) → S0 (11A g − ) carotenoid emission and Qx absorption of BChl a. This reduced energetic gap is the ultimate consequence of strong carotenoid–protein interactions, including the inferred hydrogen bonding.
Keywords: Carotenoids; Energy transfer; Ultrafast spectroscopy; Light-harvesting complex; Purple bacteria
Identification of large variation in the photosynthetic induction response among 37 soybean [Glycine max (L.) Merr.] genotypes that is not correlated with steady-state photosynthetic capacity by M. A. Soleh; Y. Tanaka; S. Y. Kim; S. C. Huber; K. Sakoda; T. Shiraiwa (305-315).
Irradiance continuously fluctuates during the day in the field. The speed of the induction response of photosynthesis in high light affects the cumulative carbon gain of the plant and could impact growth and yield. The photosynthetic induction response and its relationship with the photosynthetic capacity under steady-state conditions (P max) were evaluated in 37 diverse soybean [Glycine max (L.) Merr.] genotypes. The induction response of leaf photosynthesis showed large variation among the soybean genotypes. After 5 min illumination with strong light, genotype NAM23 had the highest leaf photosynthetic rate of 33.8 µmol CO2 m−2 s−1, while genotype NAM12 showed the lowest rate at 4.7 µmol CO2 m−2 s−1. Cumulative CO2 fixation (CCF) during the first 5 min of high light exposure ranged from 5.5 mmol CO2 m−2 for NAM23 to 0.81 mmol CO2 m−2 for NAM12. The difference in the induction response among genotypes was consistent throughout the growth season. However, there was no significant correlation between CCF and P max among genotypes suggesting that different mechanisms regulate P max and the induction response. The observed variation in the induction response was mainly attributed to ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) activation, but soybean lines differing in the induction response did not differ in the leaf content of Rubisco activase α- and β-proteins. Future studies will be focused on identifying molecular determinants of the photosynthetic induction response and determining whether this trait could be an important breeding target to achieve improved growth of soybeans in the field.
Keywords: Soybean [Glycine max (L.) Merr.]; Photosynthetic induction response; Rubisco activase; Photosynthetic capacity
An enzyme kinetics study of the pH dependence of chloride activation of oxygen evolution in photosystem II by Sergei Baranov; Alice Haddy (317-332).
Oxygen evolution by photosystem II (PSII) involves activation by Cl− ion, which is regulated by extrinsic subunits PsbQ and PsbP. In this study, the kinetics of chloride activation of oxygen evolution was studied in preparations of PSII depleted of the PsbQ and PsbP subunits (NaCl-washed and Na2SO4/pH 7.5-treated) over a pH range from 5.3 to 8.0. At low pH, activation by chloride was followed by inhibition at chloride concentrations >100 mM, whereas at high pH activation continued as the chloride concentration increased above 100 mM. Both activation and inhibition were more pronounced at lower pH, indicating that Cl− binding depended on protonation events in each case. The simplest kinetic model that could account for the complete data set included binding of Cl− at two sites, one for activation and one for inhibition, and four protonation steps. The intrinsic (pH-independent) dissociation constant for Cl− activation, K S, was found to be 0.9 ± 0.2 mM for both preparations, and three of the four pK as were determined, with the fourth falling below the pH range studied. The intrinsic inhibition constant, K I, was found to be 64 ± 2 and 103 ± 7 mM for the NaCl-washed and Na2SO4/pH7.5-treated preparations, respectively, and is considered in terms of the conditions likely to be present in the thylakoid lumen. This enzyme kinetics analysis provides a more complete characterization of chloride and pH dependence of O2 evolution activity than has been previously presented.
Keywords: Chloride; Enzyme kinetics; Oxygen evolution; pH dependence; Photosystem II; Substrate inhibition
Impact of elevated temperatures on specific leaf weight, stomatal density, photosynthesis and chlorophyll fluorescence in soybean by Kanchan Jumrani; Virender Singh Bhatia; Govind Prakash Pandey (333-350).
High-temperature stress is a major environmental stress and there are limited studies elucidating its impact on soybean (Glycine max L. Merril.). The objectives of present study were to quantify the effect of high temperature on changes in leaf thickness, number of stomata on adaxial and abaxial leaf surfaces, gas exchange, chlorophyll fluorescence parameters and seed yield in soybean. Twelve soybean genotypes were grown at day/night temperatures of 30/22, 34/24, 38/26 and 42/28 °C with an average temperature of 26, 29, 32 and 35 °C, respectively, under greenhouse conditions. One set was also grown under ambient temperature conditions where crop season average maximum, minimum and mean temperatures were 28.0, 22.4 and 25.2 °C, respectively. Significant negative effect of temperature was observed on specific leaf weight (SLW) and leaf thickness. Rate of photosynthesis, stomatal conductance and water use efficiency declined as the growing temperatures increased; whereas, intercellular CO2 and transpiration rate were increased. With the increase in temperature chlorophyll fluorescence parameters such as Fv/Fm, qP and PhiPSII declined while there was increase in qN. Number of stomata on both abaxial and adaxial surface of leaf increased significantly with increase in temperatures. The rate of photosynthesis, PhiPSII, qP and SPAD values were positively associated with leaf thickness and SLW. This indicated that reduction in photosynthesis and associated parameters appears to be due to structural changes observed at higher temperatures. The average seed yield was maximum (13.2 g/pl) in plants grown under ambient temperature condition and declined by 8, 14, 51 and 65% as the temperature was increased to 30/22, 34/24, 38/26 and 42/28 °C, respectively.
Keywords: Chlorophyll fluorescence; Crop yield; Leaf thickness; Photosynthesis; Stomatal density; Temperature
Electron flow from water to NADP+ with students acting as molecules in the chain: a Z-scheme drama in a classroom by Sagarika Jaiswal; Misha Bansal; Shirshanya Roy; Adyasha Bharati; Barnali Padhi (351-359).