Intersystem Electron Transport Research Articles

Natural variation in photosynthesis and water use efficiency of locally adapted Persian walnut populations under drought stress and recovery

Persian walnut is a drought-sensitive species with considerable genetic variation in the photosynthesis and water use efficiency of its populations, which is largely unexplored. Here, we aimed to elucidate changes in the efficiency of photosynthesis and water content using a diverse panel of 60 walnut families which were submitted to a progressive drought for 24 days, followed by two weeks of re-watering. Severe water-withholding reduced leaf relative water content (RWC) by 20%, net photosynthetic rate (Pn) by 50%, stomatal conductance (gs) by 60%, intercellular CO2 concentration (Ci) by 30%, and transpiration rate (Tr) by 50%, but improved water use efficiency (WUE) by 25%. Severe water-withholding also inhibited photosystem II functionality as indicated by reduced quantum yield of intersystem electron transport (φEo) and transfer of electrons per reaction center (ET0/RC), also enhanced accumulation of QA (VJ) resulted in the reduction of the photosynthetic performance (PIABS) and maximal quantum yield of PSII (FV/FM); while elevated quantum yield of energy dissipation (φDo), energy fluxes for absorption (ABS/RC) and dissipated energy flux (DI0/RC) in walnut families. Cluster analysis classified families into three main groups (tolerant, moderately tolerant, and sensitive), with the tolerant group from dry climates exhibiting lesser alterations in assessed parameters than the other groups. Multivariate analysis of phenotypic data demonstrated that RWC and biophysical parameters related to the chlorophyll fluorescence such as FV/FM, φEo, φDo, PIABS, ABS/RC, ET0/RC, and DI0/RC represent fast, robust and non-destructive biomarkers for walnut performance under drought stress. Finally, phenotype-environment association analysis showed significant correlation of some photosynthetic traits with geoclimatic factors, suggesting a key role of climate and geography in the adaptation of walnut to its habitat conditions.

Variation in thermotolerance of photosystem II energy trapping, intersystem electron transport, and photosystem I electron acceptor reduction for diverse cotton genotypes

Cotton breeding programs have focused on agronomically-desirable traits. Without targeted selection for tolerance to high temperature extremes, cotton will likely be more vulnerable to environment-induced yield loss. Recently-developed methods that couple chlorophyll fluorescence induction measurements with temperature response experiments could be used to identify genotypic variation in photosynthetic thermotolerance of specific photosynthetic processes for field-grown plants. It was hypothesized that diverse cotton genotypes would differ significantly in photosynthetic thermotolerance, specific thylakoid processes would exhibit differential sensitivities to high temperature, and that the most heat tolerant process would exhibit substantial genotypic variation in thermotolerance plasticity. A two-year field experiment was conducted at Tifton and Athens, Georgia, USA. Experiments included 10 genotypes in 2020 and 11 in 2021. Photosynthetic thermotolerance for field-collected leaf samples was assessed by determining the high temperature threshold resulting in a 15% decline in photosynthetic efficiency (T15) for energy trapping by photosystem II (ΦPo), intersystem electron transport (ΦEo), and photosystem I end electron acceptor reduction (ΦRo). Significant genotypic variation in photosynthetic thermotolerance was observed, but the response was dependent on location and photosynthetic parameter assessed. ΦEo was substantially more heat sensitive than ΦPo or ΦRo. Significant genotypic variation in thermotolerance plasticity of ΦEo was also observed. Identifying the weakest link in photosynthetic tolerance to high temperature will facilitate future selection efforts by focusing on the most heat-susceptible processes. Given the genotypic differences in environmental plasticity observed here, future research should evaluate genotypic variation in acclimation potential in controlled environments.

The cytochrome b6f complex: plastoquinol oxidation and regulation of electron transport in chloroplasts.

In oxygenic photosynthetic systems, the cytochrome b6f (Cytb6f) complex (plastoquinol:plastocyanin oxidoreductase) is a heart of the hub that provides connectivity between photosystems (PS) II and I. In this review, the structure and function of the Cytb6f complex are briefly outlined, being focused on the mechanisms of a bifurcated (two-electron) oxidation of plastoquinol (PQH2). In plant chloroplasts, under a wide range of experimental conditions (pH and temperature), a diffusion of PQH2 from PSII to the Cytb6f does not limit the intersystem electron transport. The overall rate of PQH2 turnover is determined mainly by the first step of the bifurcated oxidation of PQH2 at the catalytic site Qo, i.e., the reaction of electron transfer from PQH2 to the Fe2S2 cluster of the high-potential Rieske iron-sulfur protein (ISP). This point has been supported by the quantum chemical analysis of PQH2 oxidation within the framework of a model system including the Fe2S2 cluster of the ISP and surrounding amino acids, the low-potential heme b6L, Glu78 and 2,3,5-trimethylbenzoquinol (the tail-less analog of PQH2). Other structure-function relationships and mechanisms of electron transport regulation of oxygenic photosynthesis associated with the Cytb6f complex are briefly outlined: pH-dependent control of the intersystem electron transport and the regulatory balance between the operation of linear and cyclic electron transfer chains.

Integrative Transcriptome and Chlorophyll Fluorescence Test Analysis Shed New Light on the Leaf Senescence Mechanism of Zoysia japonica

Zoysia japonica is an important warm-season turfgrass used worldwide. The decreased aesthetic quality and functionality during leaf senescence hamper its further utilization. However, information about the transcriptional mechanism and genes involved in leaf senescence in Z. japonica needs to be more extensive. Therefore, to better understand leaf senescence in Z. japonica, we investigated the integrated analysis of chlorophyll fluorescence test (JIP-test) and RNA sequencing (RNA-seq) of mature and senescent leaves. First, we identified 22,049 genes, of which 4038 were differentially expressed genes (DEGs). The results for gene expression profiles were evaluated using quantitative real-time PCR. A total of 2515 genes have homologous genes in other plants. The matched known-function SAGs are mainly involved in chlorophyll degradation and plant hormone response. A total of 539 differentially expressed transcription factor genes, including AP2/ERF-ERF, NAC, WRKY, bHLH, and MYB, were identified to be associated with leaf senescence. Next, senescence represses chlorophyll biosynthesis while upregulating chlorophyll degradation. Senescence harms the integrity and functionality of PSII, PSI, and the intersystem electron transport chain. In addition, IAA biosynthesis was inhibited, whereas ABA and ET biosynthesis were activated in leaf senescence, and senescence activates signal transduction of IAA, ABA, and ET. These findings add to our understanding of the regulatory mechanism of leaf senescence. The senescence-associated genes are candidate targets for providing new insight into leaf senescence modeling in Z. japonica. They provided a theoretical foundation to reveal the functions of senescence-associated genes and chlorophyll catabolic genes involved in leaf senescence.

The noninvasive monitoring of the redox status of photosynthetic electron transport chains in Hibiscus rosa-sinensis and Tradescantia leaves

We present an approach to the noninvasive determination of the electron capacity of the intersystem pool of electron carriers in chloroplasts in situ. As apt experimental models, we used the leaves of Hibiscus rosa-sinensis and Tradescantia species. Electron paramagnetic resonance and optical response of P700 (the primary electron donor in Photosystem I) were applied to measuring electron transport in chloroplasts. Electron capacities of the intersystem electron transport chain (ETC) were determined from redox transients of P700 upon chromatic transitions (white light → far-red light). During the induction period, we observed the nonmonotonic changes in the number of electron equivalents in the intersystem ETC per P700 (parameter Q). In Hibiscus rosa-sinensis, the light-induced rise of Q from ≈2.5 (in the dark) to Q ≈ 12 was followed by its decrease to Q ≈ 6. The data obtained are discussed in the context of pH-dependent regulation of electron transport in chloroplasts, which provides the well-balanced operation of the intersystem ETC. The decay of Q is explained by the attenuation of Photosystem II activity due to the lumen acidification and the acceleration of plastoquinol re-oxidation as a result of the Calvin-Benson cycle activation. Our computer model of electron and proton transport coupled to ATP synthesis in chloroplasts was used to analyze the up and down feedbacks responsible for pH-dependent regulation of electron transport in chloroplasts. The procedures introduced here may be important for subsequent works aimed at defining the plastoquinone participation in regulation of photosynthetic processes in chloroplasts in situ.

Low Light Increases the Abundance of Light Reaction Proteins: Proteomics Analysis of Maize (Zea mays L.) Grown at High Planting Density.

Maize (Zea mays L.) is usually planted at high density, so most of its leaves grow in low light. Certain morphological and physiological traits improve leaf photosynthetic capacity under low light, but how light absorption, transmission, and transport respond at the proteomic level remains unclear. Here, we used tandem mass tag (TMT) quantitative proteomics to investigate maize photosynthesis-related proteins under low light due to dense planting, finding increased levels of proteins related to photosystem II (PSII), PSI, and cytochrome b6f. These increases likely promote intersystem electron transport and increased PSI end electron acceptor abundance. OJIP transient curves revealed increases in some fluorescence parameters under low light: quantum yield for electron transport (φEo), probability that an electron moves beyond the primary acceptor QA− (ψo), efficiency/probability of electron transfer from intersystem electron carriers to reduction end electron acceptors at the PSI acceptor side (δRo), quantum yield for reduction of end electron acceptors at the PSI acceptor side (φRo), and overall performance up to the PSI end electron acceptors (PItotal). Thus, densely planted maize shows elevated light utilization through increased electron transport efficiency, which promotes coordination between PSII and PSI, as reflected by higher apparent quantum efficiency (AQE), lower light compensation point (LCP), and lower dark respiration rate (Rd).

Extreme temperatures affect seedling growth and photosynthetic performance of advanced cotton genotypes

Growth temperature and genotype can influence seedling vigor in cotton (Gossypium hirsutum L.), and identifying genotypes that can perform well under different temperature extremes may broaden the range of temperatures over which optimum growth could be obtained. To this end, a controlled environment study was conducted to evaluate the response of advanced breeding lines to growth temperature and to evaluate the utility of rapid, fluorescence-based measurements (the OJIP test) as indicators of plant growth response to day/night temperature regimes (20/15, 30/20, 35/25, and 40/30 °C). At two weeks after planting, growth analysis, chlorophyll fluorescence measurements, and pigment concentrations were obtained. Significant genotype, temperature, and interaction effects were observed for seedling growth parameters, and some of the key components of the thylakoid reactions. Specifically, the 35/25 °C treatment had the highest values for all growth parameters, with genotypic differences in growth primarily being observed at this regime. Energy trapping by photosystem II (PSII) (φPO), intersystem electron transport (φEO), and photosystem I end electron acceptor reduction (φRO) were significantly affected by temperature, but each component differed in heat sensitivity. All growth parameters were significantly correlated with a number of OJIP parameters including quantum efficiencies and performance indices. However, the strongest positive associations were observed between quantum efficiencies and growth metrics, with φEO exhibiting the strongest correlations with growth. Our observations indicate that rapid OJIP assessments can potentially be used as indicators of early season growth responses to temperature.

Functional characterization of the chlorophyll b reductase gene NYC1 associated with chlorophyll degradation and photosynthesis in Zoysia japonica

Chlorophyll plays essential roles in photosynthesis and its degradation should be regulated precisely. Chlorophyll b reductase encoded by the gene NYC1 is responsible for catalyzing chlorophyll b degradation and maintaining the stability of photosystems. Although NYC1s have been characterized in many plants, their function in the photosynthetic machinery and the underlying regulatory mechanisms at the whole transcriptome level remain elusive. In this study, the ZjNYC1 gene was cloned from the important warm-season turfgrass Zoysia japonica. Analysis of the subcellular localization revealed that ZjNYC1 was localized in the chloroplasts, and results of the bimolecular fluorescence complementation (BiFC) assay indicated that ZjNYC1 could interact in vivo with the NYC1-like protein, ZjNOL. Overexpression of ZjNYC1 accelerated the degradation of chlorophyll, and promoted senescence with accumulation of ABA and reactive oxygen species (ROS) in Arabidopsis. Expression of ZjNYC1 rescued the stay-green phenotype of the Arabidopsis nyc1 mutant. An integrated analysis of chlorophyll fluorescence test (JIP-test) and RNA-sequencing revealed that ZjNYC1 negatively affected the efficiency of the PSII, PSI, and the intersystem electron transport chain. Furthermore, the Calvin cycle and photorespiration were suppressed, but ABA and ethylene synthesis and signal transduction pathways were activated in the transgenic lines. Collectively, these results demonstrated that ZjNYC1 plays important roles in accelerating chlorophyll degradation and senescence, and caused negative affect on the integrity and functionality of the photosystems.

Photomorphogenesis in the Picocyanobacterium Cyanobium gracile Includes Increased Phycobilisome Abundance Under Blue Light, Phycobilisome Decoupling Under Near Far-Red Light, and Wavelength-Specific Photoprotective Strategies.

Photomorphogenesis is a process by which photosynthetic organisms perceive external light parameters, including light quality (color), and adjust cellular metabolism, growth rates and other parameters, in order to survive in a changing light environment. In this study we comprehensively explored the light color acclimation of Cyanobium gracile, a common cyanobacterium in turbid freshwater shallow lakes, using nine different monochromatic growth lights covering the whole visible spectrum from 435 to 687 nm. According to incident light wavelength, C. gracile cells performed great plasticity in terms of pigment composition, antenna size, and photosystem stoichiometry, to optimize their photosynthetic performance and to redox poise their intersystem electron transport chain. In spite of such compensatory strategies, C. gracile, like other cyanobacteria, uses blue and near far-red light less efficiently than orange or red light, which involves moderate growth rates, reduced cell volumes and lower electron transport rates. Unfavorable light conditions, where neither chlorophyll nor phycobilisomes absorb light sufficiently, are compensated by an enhanced antenna size. Increasing the wavelength of the growth light is accompanied by increasing photosystem II to photosystem I ratios, which involve better light utilization in the red spectral region. This is surprisingly accompanied by a partial excitonic antenna decoupling, which was the highest in the cells grown under 687 nm light. So far, a similar phenomenon is known to be induced only by strong light; here we demonstrate that under certain physiological conditions such decoupling is also possible to be induced by weak light. This suggests that suboptimal photosynthetic performance of the near far-red light grown C. gracile cells is due to a solid redox- and/or signal-imbalance, which leads to the activation of this short-term light acclimation process. Using a variety of photo-biophysical methods, we also demonstrate that under blue wavelengths, excessive light is quenched through orange carotenoid protein mediated non-photochemical quenching, whereas under orange/red wavelengths state transitions are involved in photoprotection.

Temperature-dependent regulation of electron transport and ATP synthesis in chloroplasts in vitro and in silico.

The significance of temperature-dependent regulation of photosynthetic apparatus (PSA) is determined by the fact that plant temperature changes with environmental temperature. In this work, we present a brief overview of temperature-dependent regulation of photosynthetic processes in class B chloroplasts (thylakoids) and analyze these processes using a computer model that takes into account the key stages of electron and proton transport coupled to ATP synthesis. The rate constants of partial reactions were parametrized on the basis of experimental temperature dependences of partial photosynthetic processes: (1) photosystem II (PSII) turnover and plastoquinone (PQ) reduction, (2) the plastoquinol (PQH2) oxidation by the cytochrome (Cyt) b6f complex, (3) the ATP synthase activity, and (4) the proton leak from the thylakoid lumen. We consider that PQH2 oxidation is the rate-limiting step in the intersystem electron transport. The parametrization of the rate constants of these processes is based on earlier experimental data demonstrating strong correlations between the functional and structural properties of thylakoid membranes that were probed with the lipid-soluble spin labels embedded into the membranes. Within the framework of our model, we could adequately describe a number of experimental temperature dependences of photosynthetic reactions in thylakoids. Computer modeling of electron and proton transport coupled to ATP synthesis supports the notion that PQH2 oxidation by the Cyt b6f complex and proton pumping into the lumen are the basic temperature-dependent processes that determine the overall electron flux from PSII to molecular oxygen and the net ATP synthesis upon variations of temperature. The model describes two branches of the temperature dependence of the post-illumination reduction of [Formula: see text] characterized by different activation energies (about 60 and ≤ 3.5kJmol-1). The model predicts the bell-like temperature dependence of ATP formation, which arises from the balance of several factors: (1) the thermo-induced acceleration of electron transport through the Cyt b6f complex, (2) deactivation of PSII photochemistry at sufficiently high temperatures, and (3) acceleration of the passive proton outflow from the thylakoid lumen bypassing the ATP synthase complex. The model describes the temperature dependence of experimentally measured parameter P/2e, determined as the ratio between the rates of ATP synthesis and pseudocyclic electron transport (H2O → PSII → PSI → O2).

Melatonin induced changes in photosynthetic efficiency as probed by OJIP associated with improved chromium stress tolerance in canola (Brassica napus L.)

Chromium toxicity is considered as a major problem for agricultural soil that reduced crop productivity by affecting photosynthetic tissues. Exogenous application of melatonin can alleviate the adverse effects of chromium toxicity on plant growth. However, little is known about its effect on thylakoidal protein complexes responsible for conversion of solar energy to biochemical energy. Chlorophyll fluorescence a transients considered one of the best non-invasive and rapid method for the evaluation of photosynthetic (Photosystem II) efficiency of plants and plant health under environmental stress conditions. In the present study, three-week old plants of two canola cultivars AC-Excel and DGL were applied to melatonin (0, 1, 5, 10 μM) when grown under chromium stress (0, 50 and 100 μM) for further two weeks. Chromium stress reduced the growth (fresh and dry weights of shoots and roots) of both canola cultivars and exogenous application of 5 and 10 μM melatonin improved the growth of canola at 50 or 100 μM chromium stress. This improvement was greater in cv DGL than in AC-Excel. Increasing chromium decreased the photosynthetic pigments (chlorophyll a and chlorophyll b). However, 5 and 10 μM melatonin application improved chlorophyll a at 50 μM chromium stress. Structural stability and efficiency of photosystem II (PSII) measured as performance index (PIABS) and ratios of fluorescence (Fv/Fm, Fv/Fo) Fv decreased due to chromium stress. JIP-test parameters showed that chromium stress increased the absorption and trapping fluxes with decrease in electron transport fluxes which caused the damage to reaction centers (RC), detachment of oxygen evolving complex (OEC) from RC or inefficiency of electron transfer from OEC to RC. Such adverse effects were greater in cv AC-Excel. However exogenous application of melatonin improved PIABS, electron transport per reaction center (ET/RC), reduced variable fluorescence at J step (VJ) reflecting melatonin protected PSII from chromium stress induced damage by protecting OEC. Thus, OJIP fluorescence transients are quite helpful for understanding the intersystem electron transport beyond photosystem II in canola cultivars due to melatonin application under chromium stress. FindingsExogenous application of melatonin alleviated toxic effects of chromium on plant growth of canola by modulating photosynthesis, enhanced photosystem II efficiency and regulation of electron transport flux to protect photo-inhibition of PSII from oxidative damage.

Revisiting cyanobacterial state transitions

Photosynthetic organisms are exposed to a fluctuating environment in which light intensity and quality change continuously. Specific illumination of either photosystem (PSI or PSII) creates an energy imbalance, leading to the reduction or oxidation of the intersystem electron transport chain. This redox imbalance could trigger the formation of dangerous reactive oxygen species. Cyanobacteria, like plants and algae, have developed a mechanism to re-balance this preferential excitation of either reaction center, called state transitions. State transitions are triggered by changes in the redox state of the membrane-soluble plastoquinone (PQ) pool. In plants and green algae, these changes in redox potential are sensed by Cytochrome b6f, which interacts with a specific kinase that triggers the movement of the main PSII antenna (the light-harvesting complex II). By contrast, although cyanobacterial state transitions have been studied extensively, there is still no agreement about the molecular mechanism, the PQ redox state sensor and the signaling pathways involved. In this review, we aimed to critically evaluate the results published on cyanobacterial state transitions, and discuss the “new” and “old” models in the subject. The phycobilisome and membrane contributions to this physiological process were addressed and the current hypotheses regarding its signaling transduction pathway were discussed.

Chromium effects on photosynthetic electron transport in pea (Pisum sativum L.).

Components of the photosynthetic electron transport chain in pea (Pisum sativum L.) leaves under in vivo conditions showed the following sensitivity to the inhibitory action of chromium(VI): intersystem electron transport > photosystem I > photosystem II. Inhibitory effects of chromium (VI) (K2Cr2O7, Cr) on the light reactions of photosynthesis were studied in vivo in Pisum sativum L.by using Multi-function Plant Efficiency Analyser (M-PEA-2). Photosynthetic parameters related to photosystem (PS) II, PSI and intersystem electron carriers were calculated from the light-induced kinetics of prompt chlorophyll a fluorescence (OJIP transient), delayed chlorophyll a fluorescence (DF), and 820nm modulated reflection (MR). We showed that the I2 step of DF induction is sensitive to inhibition of the Q0 site of the cytochrome b6f complex. Such parameters as δRo of the JIP test related to the functional state of photosynthetic reactions beyond the PQ pool, Vred of the MR induction assigned to the overall rate of P700+ and plastocyanin reduction, and I2 step of the DF induction were significantly altered in the presence of low-dose Cr(VI). Moderate doses of Cr affected mainly PSI-related parameters including Vox and ΔMR parameters of the MR induction, whereas high-dose treatment influenced JIP test parameters φPo(= FV/FM) and ψEo related to PSII. The obtained results showed that the earliest Cr(VI) effect on the photosynthetic electron transport chain manifests itself by inhibition of the intersystem electron transport, rather, at the level of the cytochrome b6f complex. Inhibitory effects of Cr on PSI were more pronounced than those on PSII. Sensitivity of the used kinetic parameters toward the functional state of photosynthetic reactions makes this approach suitable for early diagnostics of toxic action of pollutants on plants.

Early Brassica Crops Responses to Salinity Stress: A Comparative Analysis Between Chinese Cabbage, White Cabbage, and Kale.

Soil salinity is severely affecting crop productivity in many countries, particularly in the Mediterranean area. To evaluate early plant responses to increased salinity and characterize tolerance markers, three important Brassica crops – Chinese cabbage (Brassica rapa ssp. pekinensis), white cabbage (B. oleracea var. capitata) and kale (B. oleracea var. acephala) were subjected to short-term (24 h) salt stress by exposing them to NaCl at concentrations of 50, 100, or 200 mM. Physiological (root growth, photosynthetic performance parameters, and Na+/K+ ratio) and biochemical parameters (proline content and lipid peroxidation as indicated by malondialdehyde, MDA, levels) in the plants’ roots and leaves were then measured. Photosynthetic parameters such as the total performance index PItotal (describing the overall efficiency of PSI, PSII and the intersystem electron transport chain) appeared to be the most salinity-sensitive parameter and informative stress marker. This parameter was decreased more strongly in Chinese cabbage than in white cabbage and kale. It indicated that salinity reduced the capacity of the photosynthetic system for efficient energy conversion, particularly in Chinese cabbage. In parallel with the photosynthetic impairments, the Na+/K+ ratio was highest in Chinese cabbage leaves and lowest in kale leaves while kale root is able to keep high Na+/K+ ratio without a significant increase in MDA. Thus Na+/K+ ratio, high in root and low in leaves accompanying with low MDA level is an informative marker of salinity tolerance. The crops’ tolerance was positively correlated with levels of the stress hormone abscisic acid (ABA) and negatively correlated with levels of jasmonic acid (JA), and jasmonoyl-L-isoleucine (JA-Ile). Furthermore, salinity induced contrasting changes in levels of the growth-promoting hormones brassinosteroids (BRs). The crop’s tolerance was positively correlated with levels of BR precursor typhasterol while negatively with the active BR brassinolide. Principal Component Analysis revealed correlations in observed changes in phytohormones, biochemical, and physiological parameters. Overall, the results show that kale is the most tolerant of the three species and Chinese cabbage the most sensitive to salt stress, and provide holistic indications of the spectrum of tolerance mechanisms involved.

Effect of soil salinity on growth, metal distribution and photosynthetic performance of two Lycium species

The effect of salinity on gas exchange and fluorescence characteristics in Lycium barbarum and Lycium chinense grown on nonsaline and saline soils was investigated. The distribution of Na+ between three types of substrates and different organs showed acropetal concentration gradient for Na+ accumulation only in L. barbarum grown on peat-moss-perlite and saline soil. Selective uptake of K+ over Na+ in the L. barbarum leaves was established. All plants grown on saline soil showed a slight inhibition of PSII activity as revealed by the quantum yield of the primary photochemical reaction. Intersystem electron transport, reduction of the end electron acceptors, and the performance index of photosynthesis declined to a greater extent in L. barbarum than that in L. chinense. The higher tolerance of L. chinense to salt stress was accompanied with the higher overall photosynthetic performance.

OJIP-fluorescence parameters as rapid indicators of cotton (Gossypium hirsutum L.) seedling vigor under contrasting growth temperature regimes

Vigorous seedling growth in cotton is desirable because it minimizes the negative impact of multiple early season stresses, and seedling vigor can be impacted by early season growth temperature or cultivar. OJIP fluorescence provides rapid information on a broad range of photosynthetic component processes and may be a useful surrogate for seeding vigor, but this possibility has not been evaluated previously in cotton. To this end, a controlled environment study was conducted with six cultivars selected based on seed characteristics that are widely indicative of vigor and under two growth temperature regimes (sub-optimal = 20/15 °C day/night temperature; optimal = 30/20 °C) for the first two weeks after seed germination. Thereafter multiple whole-plant vigor assessments were conducted along with extensive OJIP-fluorescence characterization in cotyledons. Growth temperature was the primary factor influencing multiple plant responses. Specifically, all whole-plant indicators of seedling vigor were negatively impacted by sub-optimal temperature as were all photosynthetic performance indices and quantum efficiencies. By comparison, most photosynthetic structural indicators or reaction center-specific fluxes were either unaffected or positively impacted by low growth temperature, largely because PSII antenna size increased. The performance index, PIABS, and the quantum efficiency, φEo, were the most sensitive to low growth temperature and exhibited the strongest relationships with whole-plant seedling vigor. Thus, OJIP parameters incorporating intersystem electron transport beyond PSII but not additional downstream processes may represent the most useful surrogates for whole-plant seedling vigor in cotton.

Sub-optimal emergence temperature alters thermotolerance of thylakoid component processes in cotton seedlings

Cotton (Gossypium hirsutum) is often planted under sub-optimal early season temperatures, yet it is unknown whether this increases susceptibility to acute high temperature exposure. To address this, cotton seedlings were grown under optimal (30/20 °C) and sub-optimal (20/15 °C) growth temperature regimes, and comprehensive fluorescence analysis was conducted for cotyledons from both temperature regimes in response to incubation temperatures ranging from 30 to 50 °C. Results indicated that low growth temperature did not alter reaction center density (RC/CS0), but increased antenna size (ratio of antenna chlorophyll to reaction center chlorophyll; ABS/RC) and decreased the amplitude of the I to P phase of the fluorescence transient [ΔVIP; associated with the photosystem (PS) I content]. Photosynthetic performance indices (PItotal and PIABS), quantum yield of energy trapping (φPo), the contribution of the thylakoid reactions to PIABS (Fv/F0), electron transport to intersystem electron acceptors (φE0) and reduction of PSI end electron acceptors (φRo) were all substantially lower in plants grown under sub-optimal temperature than in plants grown at optimal temperature. Furthermore, thermotolerance of all the aforementioned parameters was significantly reduced by low early season growth temperatures. Among different OJIP-derived parameters, PIABS and PItotal were more heat sensitive than other fluorescence-based parameters, and φPo and φRo were the most heat tolerant. In addition, thermotolerance of PItotal (overall photosynthetic thermotolerance) was significantly correlated with thermotolerance of all other OJIP-derived parameters. Regression analysis indicated that heat tolerance trends for φE0 were more consistent with the changes in thermotolerance of PItotal. We conclude that 1) low growth temperature in the seedling phase of cotton results in decreased photosynthetic thermotolerance and 2) overall photosynthetic thermotolerance is primarily constrained by the ability of intersystem electron transport to acclimate to growth environment.

The cytochrome b6f complex: DFT modeling of the first step of plastoquinol oxidation by the iron-sulfur protein

In chloroplasts, the cytochrome (Cyt) b6f complex (plastoquinol-plastocyanin-oxidoreductase) provides connectivity between photosystems (PS) II and I, oxidizing plastoquinol (PQH2) formed in PSII and reducing plastocyanin (electron donor to PSI). The overall rate of the intersystem electron transport is determined by PQH2 oxidation by the Cyt b6f complex. In this work, using the DFT method, we have modeled the first step of PQH2 oxidation by the iron-sulfur protein (ISP) of the Cyt b6f complex. The model system contained the iron–sulfur cluster [Fe2S2], surrounding amino acid residues, and 2,3,5-trimethylbenzoquinol (TMBQH2), the tailless analog of PQH2. The energy profiles of the H atom transfer from TMBQH2 to the iron-sulfur protein (ISP) were calculated for two modes of the H-transfer, “diabatic” and “adiabatic”. The energies of transient states were estimated as 18.4 and 14.4 kcal mol−1. The energy effects of the reaction were evaluated as 7.7 and −0.2 kcal mol−1, respectively. The analysis of partial spin densities and electric charges on the atoms of the model system supports the bidirectional mechanism of the H-transfer reaction: an electron is directed to the Fe(1) atom of the [Fe2S2] cluster of the ISP, and a proton is accepted by the Nε atom of the His155 residue liganding the Fe(1) atom. Using the results of DFT computations of the energy profiles for the H-transfer reaction, we estimated the rate constants of quinol oxidation within the framework of the Marcus-Dutton-Crofts approach. Our analysis supports the diabatic model of the H-transfer, which implies that the elementary steps of electron and proton transfer occur much more rapidly than the concomitant changes in the system geometry. The rate constants estimated for this mode of H-transfer are in reasonable agreement with experimental data on the intersystem electron transport in chloroplasts.

The Cytochrome b 6 f Complex: Biophysical Aspects of Its Functioning in Chloroplasts.

This chapter presents an overview of structural properties of the cytochrome (Cyt) b 6 f complex and its functioning in chloroplasts. The Cyt b 6 f complex stands at the crossroad of photosynthetic electron transport pathways, providing connectivity between Photosystem (PSI) and Photosysten II (PSII) and pumping protons across the membrane into the thylakoid lumen. After a brief review of the chloroplast electron transport chain, the consideration is focused on the structural organization of the Cyt b 6 f complex and its interaction with plastoquinol (PQH2, reduced form of plastoquinone), a mediator of electron transfer from PSII to the Cyt b 6 f complex. The processes of PQH2 oxidation by the Cyt b 6 f complex have been considered within the framework of the Mitchell's Q-cycle. The overall rate of the intersystem electron transport is determined by PQH2 turnover at the quinone-binding site Qo of the Cyt b 6 f complex. The rate of PQH2 oxidation is controlled by the intrathylakoid pHin, which value determines the protonation/deprotonation events in the Qo-center. Two other regulatory mechanisms associated with the Cyt b 6 f complex are briefly overviewed: (i) redistribution of electron fluxes between alternative (linear and cyclic) pathways, and (ii) "state transitions" related to redistribution of solar energy between PSI and PSII.

Delayed fluorescence emission as a probe for the response of photosynthetic organisms to high temperature exposure: A comparative study

The effects of a short-term exposure (40s) to 50°C on the prompt fluorescence (PF), delayed fluorescence (DF) and 820nm reflection (MR) signals in three different photosynthetic organisms: Pisum sativum (angiosperm plant), Ginkgo biloba (gymnosperm tree) and thalli of Parmelina tiliacea (Hoffm.) Ach (lichen) were investigated. The aim of this study was to understand the relationship between prompt PF, DF and MR when the oxygen evolving complex (OEC) is destroyed. The signals of PF, DF and MR were measured simultaneously in plants in vivo to probe heat-induced damages in the photosynthetic electron transfer. PF is characterised by a polyphasic fluorescence rise OJIPSMT and DF induction curve has several maxima (In) and numbered in sequence according to their appearance during illumination. In control condition (25°C) leaves of pea plants yield a typical OJIP rise with a flat P-step (0.2–2s) and in G. biloba and P. tiliacea thalli, the P-level was split into two peaks, the first peak P (0.2s) is followed by a decrease of the fluorescence intensity to a dip (S) and a subsequent rise to a second peak (M). An antiparallel relation between J–I–P–S and DF curves was also observed. Methyl viologen (MV) reduced the amplitude of the I–P phase of the PF curve and this change was accompanied by a disappearance of the PSM transient. In the presence of MV, the DF intensity remained higher; and the I4 peak disappeared. We suggested that P-S-M rise from PF and I4 peak might be related to an acceleration of intersystem electron transport. After 50°C incubation, significant changes appeared in the PF, DF and MR transients and that were dependent on the photosynthetic sample. Heat treatment provoked a down regulation of electrons donation by the OEC in the three plant species and the re-reduction of P700+ and PC+ started earlier and began already after 10 ms of illumination. Inhibition of the donation of electrons by OEC after 50°C incubation indicates that DF induction depends on the concentrations of QA− and P680+ and the proton gradient.