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Current work in photosynthetic engineering is progressing along the lines of cyanobacterial, microalgal, and plant research. These are interconnected through the fundamental mechanisms of photosynthesis and advances in one field can often be leveraged to improve another. It is worthwhile for researchers specializing in one or more of these systems to be aware of the work being done across the entire research space as parallel advances of techniques and experimental approaches can often be applied across the field of photosynthesis research. This review focuses on research published in recent years related to the light reactions of photosynthesis in cyanobacteria, eukaryotic algae, and plants. Highlighted are attempts to improve photosynthetic efficiency, and subsequent biomass production. Also discussed are studies on cross-field heterologous expression, and related work on augmented and novel light capture systems. This is reviewed in the context of translatability in research across diverse photosynthetic organisms. [ABSTRACT FROM AUTHOR]

Current work in photosynthetic engineering is progressing along the lines of cyanobacterial, microalgal, and plant research. These are interconnected through the fundamental mechanisms of photosynthesis and advances in one field can often be leveraged to improve another. It is worthwhile for researchers specializing in one or more of these systems to be aware of the work being done across the entire research space as parallel advances of techniques and experimental approaches can often be applied across the field of photosynthesis research. This review focuses on research published in recent years related to the light reactions of photosynthesis in cyanobacteria, eukaryotic algae, and plants. Highlighted are attempts to improve photosynthetic efficiency, and subsequent biomass production. Also discussed are studies on cross-field heterologous expression, and related work on augmented and novel light capture systems. This is reviewed in the context of translatability in research across diverse photosynthetic organisms.

Summary: Drought is a major abiotic stress that impairs plant growth and development. Despite this, a comprehensive understanding of drought effects on the photosynthetic apparatus is lacking. In this work, we studied the consequences of 14‐d drought treatment on Arabidopsis thaliana.We used biochemical and spectroscopic methods to examine photosynthetic membrane composition and functionality.Drought led to the disassembly of PSII supercomplexes and the degradation of PSII core. The light‐harvesting complexes (LHCII) instead remain in the membrane but cannot act as an antenna for active PSII, thus representing a potential source of photodamage. This effect can also be observed during nonphotochemical quenching (NPQ) induction when even short pulses of saturating light can lead to photoinhibition. At a later stage, under severe drought stress, the PSI antenna size is also reduced and the PSI‐LHCI supercomplexes disassemble. Surprisingly, although we did not observe changes in the PSI core protein content, the functionality of PSI is severely affected, suggesting the accumulation of nonfunctional PSI complexes.We conclude that drought affects both photosystems, although at a different stage, and that the operative quantum efficiency of PSII (ΦPSII) is very sensitive to drought and can thus be used as a parameter for early detection of drought stress. [ABSTRACT FROM AUTHOR]

Photosynthesis is the largest mass- and energy-conversion process on Earth, and it is the material basis for almost all biological activities. The efficiency of converting absorbed light energy into energy substances during photosynthesis is very low compared to theoretical values. Based on the importance of photosynthesis, this article summarizes the latest progress in improving photosynthesis efficiency from various perspectives. The main way to improve photosynthetic efficiency is to optimize the light reactions, including increasing light absorption and conversion, accelerating the recovery of non-photochemical quenching, modifying enzymes in the Calvin cycle, introducing carbon concentration mechanisms into C3 plants, rebuilding the photorespiration pathway, de novo synthesis, and changing stomatal conductance. These developments indicate that there is significant room for improvement in photosynthesis, providing support for improving crop yields and mitigating changes in climate conditions. [ABSTRACT FROM AUTHOR]

Dramatic climate change has led to an increase in the intensity and frequency of drought episodes and, together with the high light conditions of the Mediterranean area, detrimentally influences crop production. Salicylic acid (SA) has been shown to supress phototoxicity, offering photosystem II (PSII) photoprotection. In the current study, we attempted to reveal the mechanism by which SA is improving PSII efficiency in oregano seedlings under moderate drought stress (MoDS). Foliar application of SA decreased chlorophyll content under normal growth conditions, but under MoDS increased chlorophyll content, compared to H2O-sprayed oregano seedlings. SA improved the PSII efficiency of oregano seedlings under normal growth conditions at high light (HL), and under MoDS, at both low light (LL) and HL. The mechanism by which, under normal growth conditions and HL, SA sprayed oregano seedlings compared to H2O-sprayed exhibited a more efficient PSII photochemistry, was the increased (17%) fraction of open PSII reaction centers (qp), and the increased (7%) efficiency of these open reaction centers (Fv′/Fm′), which resulted in an enhanced (24%) electron transport rate (ETR). SA application under MoDS, by modulating chlorophyll content, resulted in optimized antenna size and enhanced effective quantum yield of PSII photochemistry (ΦPSII) under both LL (7%) and HL (25%), compared to non-SA-sprayed oregano seedlings. This increased effective quantum yield of PSII photochemistry (ΦPSII) was due to the enhanced efficiency of the oxygen evolving complex (OEC), and the increased fraction of open PSII reaction centers (qp), which resulted in an increased electron transport rate (ETR) and a lower amount of singlet oxygen (1O2) production with less excess excitation energy (EXC). [ABSTRACT FROM AUTHOR]

SUMMARY: When C4 leaves are exposed to low light, the CO2 concentration in the bundle sheath (BS) cells decreases, causing an increase in photorespiration relative to assimilation, and a consequent reduction in biochemical efficiency. These effects can be mitigated by complex acclimation syndromes, which are of primary importance for crop productivity but are not well studied. We unveil an acclimation strategy involving the coordination of electron transport processes. First, we characterize the anatomy, gas exchange and electron transport of C4Setaria viridis grown under low light. Through a purposely developed biochemical model, we resolve the photon fluxes and reaction rates to explain how the concerted acclimation strategies sustain photosynthetic efficiency. Our results show that a smaller BS in low‐light‐grown plants limited leakiness (the ratio of CO2 leak rate out of the BS over the rate of supply via C4 acid decarboxylation) but sacrificed light harvesting and ATP production. To counter ATP shortage and maintain high assimilation rates, plants facilitated light penetration through the mesophyll and upregulated cyclic electron flow in the BS. This shade tolerance mechanism, based on the optimization of light reactions, is possibly more efficient than the known mechanisms involving the rearrangement of carbon metabolism, and could potentially lead to innovative strategies for crop improvement. Significance Statement: We mechanistically link the optical cross section of leaf compartments with the rate of electron transport, the engagement of cyclic electron flow, the relative rate of ATP and NADPH generation and fluxes through the carbon metabolism. The striking capacity of Setaria viridis to counter the decrease in light absorption in the bundle sheath with an increase of cyclic electron flow presents perhaps the most efficient mechanism of shade acclimation. [ABSTRACT FROM AUTHOR]

Salicylic acid (SA), an essential plant hormone, has received much attention due to its role in modulating the adverse effects of biotic and abiotic stresses, acting as an antioxidant and plant growth regulator. However, its role in photosynthesis under non stress conditions is controversial. By chlorophyll fluorescence imaging analysis, we evaluated the consequences of foliar applied 1 mM SA on photosystem II (PSII) efficiency of tomato (Solanum lycopersicum L.) plants and estimated the reactive oxygen species (ROS) generation. Tomato leaves sprayed with 1 mM SA displayed lower chlorophyll content, but the absorbed light energy was preferentially converted into photochemical energy rather than dissipated as thermal energy by non-photochemical quenching (NPQ), indicating photoprotective effects provided by the foliar applied SA. This decreased NPQ, after 72 h treatment by 1 mM SA, resulted in an increased electron transport rate (ETR). The molecular mechanism by which the absorbed light energy was more efficiently directed to photochemistry in the SA treated leaves was the increased fraction of the open PSII reaction centers (qp), and the increased efficiency of open reaction centers (Fv'/Fm'). SA induced a decrease in chlorophyll content, resulting in a decrease in non-regulated energy dissipated in PSII (ΦNO) under high light (HL) treatment, suggesting a lower amount of triplet excited state chlorophyll (3Chl*) molecules available to produce singlet oxygen (1O2). Yet, the increased efficiency, compared to the control, of the oxygen evolving complex (OEC) on the donor side of PSII, associated with lower formation of hydrogen peroxide (H2O2), also contributed to less creation of ROS. We conclude that under non stress conditions, foliar applied SA decreased chlorophyll content and suppressed phototoxicity, offering PSII photoprotection; thus, it can be regarded as a mechanism that reduces photoinhibition and photodamage, improving PSII efficiency in crop plants. [ABSTRACT FROM AUTHOR]

Avocado is a commercially important fruit tree which is sold worldwide. Originating in subtropical regions of the South America, this species is now grown worldwide and is sometimes exposed to cold temperatures. Specifically, frost stress harms the crop yield and its quality. While it is known in general that the photosynthetic apparatus changes in response to cold conditions, there is still not much information regarding the photosynthetic apparatus response to sporadic frost stress. In this study, we tracked the photosynthetic apparatus' light reaction of 'Hass' and 'Ettinger' avocado cultivars to frost stress, with Ettinger being known to be more resilient to cold than Hass. We found that in avocado trees, the photosynthetic apparatus' response to frost occurs at the level of photosystem II (PSII) itself, rather than a photoprotective response to a stress. The Hass apparatus incorrectly interprets the reduction in electron transport rate activity and by that increases its light harvesting complex size at the expense of its reaction centers which then increases the apparatus' probability to generate reactive oxygen species. The results of this study open opportunities to further research the process which regulates the feedback mechanism that controls the photosynthetic unit's size in Hass when compared to the Ettinger cultivar, and whether it is part of a feedback regulation from the carbon assimilation step or indirectly from a stomatal limitation which arises in these subtropical species. While corroborating past studies performed on avocados, this study suggests using advanced chlorophyll a fluorescence protocols when researching natural variation in crops. [ABSTRACT FROM AUTHOR]

Photosynthesis started to evolve some 3.5 billion years ago CO2 is the substrate for photosynthesis and in the past 200–250 years, atmospheric levels have approximately doubled due to human industrial activities. However, this time span is not sufficient for adaptation mechanisms of photosynthesis to be evolutionarily manifested. Steep increases in human population, shortage of arable land and food, and climate change call for actions, now. Thanks to substantial research efforts and advances in the last century, basic knowledge of photosynthetic and primary metabolic processes can now be translated into strategies to optimize photosynthesis to its full potential in order to improve crop yields and food supply for the future. Many different approaches have been proposed in recent years, some of which have already proven successful in different crop species. Here, we summarize recent advances on modifications of the complex network of photosynthetic light reactions. These are the starting point of all biomass production and supply the energy equivalents necessary for downstream processes as well as the oxygen we breathe. [ABSTRACT FROM AUTHOR]

Dramatic climate change has led to an increase in the intensity and frequency of drought episodes and, together with the high light conditions of the Mediterranean area, detrimentally influences crop production. Salicylic acid (SA) has been shown to supress phototoxicity, offering photosystem II (PSII) photoprotection. In the current study, we attempted to reveal the mechanism by which SA is improving PSII efficiency in oregano seedlings under moderate drought stress (MoDS). Foliar application of SA decreased chlorophyll content under normal growth conditions, but under MoDS increased chlorophyll content, compared to H2O-sprayed oregano seedlings. SA improved the PSII efficiency of oregano seedlings under normal growth conditions at high light (HL), and under MoDS, at both low light (LL) and HL. The mechanism by which, under normal growth conditions and HL, SA sprayed oregano seedlings compared to H2O-sprayed exhibited a more efficient PSII photochemistry, was the increased (17%) fraction of open PSII reaction centers (qp), and the increased (7%) efficiency of these open reaction centers (Fv′/Fm′), which resulted in an enhanced (24%) electron transport rate (ETR). SA application under MoDS, by modulating chlorophyll content, resulted in optimized antenna size and enhanced effective quantum yield of PSII photochemistry (ΦPSII) under both LL (7%) and HL (25%), compared to non-SA-sprayed oregano seedlings. This increased effective quantum yield of PSII photochemistry (ΦPSII) was due to the enhanced efficiency of the oxygen evolving complex (OEC), and the increased fraction of open PSII reaction centers (qp), which resulted in an increased electron transport rate (ETR) and a lower amount of singlet oxygen (1O2) production with less excess excitation energy (EXC).

Photosynthetic organisms have adapted to survive a myriad of extreme environments from the earth’s deserts to its poles, yet the proteins that carry out the light reactions of photosynthesis are highly conserved from the cyanobacteria to modern day crops. To investigate adaptations of the photosynthetic machinery in cyanobacteria to excessive light stress, we isolated a new strain of cyanobacteria, Cyanobacterium aponinum 0216, from the extreme light environment of the Sonoran Desert. Here we report the biochemical characterization and the 2.7 Å resolution structure of trimeric photosystem I from this high-light-tolerant cyanobacterium. The structure shows a new conformation of the PsaL C-terminus that supports trimer formation of cyanobacterial photosystem I. The spectroscopic analysis of this photosystem I revealed a decrease in far-red absorption, which is attributed to a decrease in the number of long- wavelength chlorophylls. Using these findings, we constructed two chimeric PSIs in Synechocystis sp. PCC 6803 demonstrating how unique structural features in photosynthetic complexes can change spectroscopic properties, allowing organisms to thrive under different environmental stresses.

Chemical thinning of apple fruitlets is an important practice as it reduces the natural fruit load and, therefore, increases the size of the final fruit for commercial markets. In apples, one chemical thinner used is Metamitron, which is sold as the commercial product Brevis® (Adama, Ashdod, Israel). This thinner inhibits the electron transfer between Photosystem II and Quinone-b within light reactions of photosynthesis. In this study, we investigated the responses of two apple cultivars—Golden Delicious and Top Red—and photosynthetic light reactions after administration of Brevis®. The analysis revealed that the presence of the inhibitor affects both cultivars’ energetic status. The kinetics of the photoprotective mechanism’s sub-processes are attenuated in both cultivars, but this seems more severe in the Top Red cultivar. State transitions of the antenna and Photosystem II repair cycle are decreased substantially when the Metamitron concentration is above 0.6% in the Top Red cultivar but not in the Golden Delicious cultivar. These attenuations result from a biased absorbed energy distribution between photochemistry and photoprotection pathways in the two cultivars. We suggest that Metamitron inadvertently interacts with photoprotective mechanism-related enzymes in chloroplasts of apple tree leaves. Specifically, we hypothesize that it may interact with the kinases responsible for the induction of state transitions and the Photosystem II repair cycle.

Summary: Recent progress in observing sun‐induced Chl fluorescence (SIF) provides an unprecedented opportunity to advance photosynthesis research in natural environments. However, we still lack an analytical framework to guide SIF studies and integration with the well‐developed active fluorescence approaches. Here, we derive a set of coupled fundamental equations to describe the dynamics of SIF and its relationship with C3 and C4 photosynthesis. These equations show that, although SIF is dynamically as complex as photosynthesis, the measured SIF simplifies photosynthetic modeling from the perspective of light reactions by integrating over the dynamic complexities of photosynthesis. Specifically, the measured SIF contains direct information about the actual electron transport from photosystem II to photosystem I, giving a quantifiable link between light and dark reactions. With much‐reduced requirements on inputs and parameters, the light‐reactions‐centric, SIF‐based biophysical model complements the traditional, dark‐reactions‐centric biochemical model of photosynthesis. The SIF–photosynthesis relationship, however, is nonlinear. This is because photosynthesis saturates at high light whereas SIF has a stronger tendency to keep increasing, as fluorescence quantum yield has a relatively muted sensitivity to light levels. Successful applications of the SIF‐based model of photosynthesis will depend on a predictive understanding of several previously underexplored physiological and biophysical processes. Advances can be facilitated by coordinated efforts in plant physiology, remote sensing, and eddy covariance flux observations. [ABSTRACT FROM AUTHOR]

For modeling approaches in systems biology, knowledge of the absolute abundances of cellular proteins is essential. One way to gain this knowledge is the use of quantification concatamers (QconCATs), which are synthetic proteins consisting of proteotypic peptides derived from the target proteins to be quantified. The QconCAT protein is labeled with a heavy isotope upon expression in E. coli and known amounts of the purified protein are spiked into a whole cell protein extract. Upon tryptic digestion, labeled and unlabeled peptides are released from the QconCAT protein and the native proteins, respectively, and both are quantified by LC-MS/MS. The labeled Q-peptides then serve as standards for determining the absolute quantity of the native peptides/proteins. Here, we have applied the QconCAT approach to Chlamydomonas reinhardtii for the absolute quantification of the major proteins and protein complexes driving photosynthetic light reactions in the thylakoid membranes and carbon fixation in the pyrenoid. We found that with 25.2 attomol/cell the Rubisco large subunit makes up 6.6% of all proteins in a Chlamydomonas cell and with this exceeds the amount of the small subunit by a factor of 1.56. EPYC1, which links Rubisco to form the pyrenoid, is eight times less abundant than RBCS, and Rubisco activase is 32-times less abundant than RBCS. With 5.2 attomol/cell, photosystem II is the most abundant complex involved in the photosynthetic light reactions, followed by plastocyanin, photosystem I and the cytochrome b6/f complex, which range between 2.9 and 3.5 attomol/cell. The least abundant complex is the ATP synthase with 2 attomol/cell. While applying the QconCAT approach, we have been able to identify many potential pitfalls associated with this technique. We analyze and discuss these pitfalls in detail and provide an optimized workflow for future applications of this technique.

For modeling approaches in systems biology, knowledge of the absolute abundances of cellular proteins is essential. One way to gain this knowledge is the use of quantification concatamers (QconCATs), which are synthetic proteins consisting of proteotypic peptides derived from the target proteins to be quantified. The QconCAT protein is labeled with a heavy isotope upon expression in E. coli and known amounts of the purified protein are spiked into a whole cell protein extract. Upon tryptic digestion, labeled and unlabeled peptides are released from the QconCAT protein and the native proteins, respectively, and both are quantified by LC-MS/MS. The labeled Q-peptides then serve as standards for determining the absolute quantity of the native peptides/proteins. Here, we have applied the QconCAT approach to Chlamydomonas reinhardtii for the absolute quantification of the major proteins and protein complexes driving photosynthetic light reactions in the thylakoid membranes and carbon fixation in the pyrenoid. We found that with 25.2 attomol/cell the Rubisco large subunit makes up 6.6% of all proteins in a Chlamydomonas cell and with this exceeds the amount of the small subunit by a factor of 1.56. EPYC1, which links Rubisco to form the pyrenoid, is eight times less abundant than RBCS, and Rubisco activase is 32-times less abundant than RBCS. With 5.2 attomol/cell, photosystem II is the most abundant complex involved in the photosynthetic light reactions, followed by plastocyanin, photosystem I and the cytochrome b6/ f complex, which range between 2.9 and 3.5 attomol/cell. The least abundant complex is the ATP synthase with 2 attomol/cell. While applying the QconCAT approach, we have been able to identify many potential pitfalls associated with this technique. We analyze and discuss these pitfalls in detail and provide an optimized workflow for future applications of this technique. [ABSTRACT FROM AUTHOR]

Thioredoxins (TRXs) are protein oxidoreductases that control the structure and function of cellular proteins by cleavage of a disulphide bond between the side chains of two cysteine residues. Oxidized thioredoxins are reactivated by thioredoxin reductases (TR) and a TR-dependent reduction of TRXs is called a thioredoxin system. Thiol-based redox regulation is an especially important mechanism to control chloroplast proteins involved in biogenesis, in regulation of light harvesting and distribution of light energy between photosystems, in photosynthetic carbon fixation and other biosynthetic pathways, and in stress responses of plants. Of the two plant plastid thioredoxin systems, the ferredoxin-dependent system relays reducing equivalents from photosystem I via ferredoxin and ferredoxin-thioredoxin reductase (FTR) to chloroplast proteins, while NADPH-dependent thioredoxin reductase (NTRC) forms a complete thioredoxin system including both reductase and thioredoxin domains in a single polypeptide. Chloroplast thioredoxins transmit environmental light signals to biochemical reactions, which allows fine tuning of photosynthetic processes in response to changing environmental conditions. In this paper we focus on the recent reports on specificity and networking of chloroplast thioredoxin systems and evaluate the prospect of improving photosynthetic performance by modifying the activity of thiol regulators in plants. [ABSTRACT FROM AUTHOR]

Although some elements of the photosynthetic light reactions might appear to be ideal, the overall efficiency of light conversion to biomass has not been optimized during evolution. Because crop plants are depleted of genetic diversity for photosynthesis, efforts to enhance its efficiency with respect to light conversion to yield must generate new variation. In principle, three sources of natural variation are available: (i) rare diversity within extant higher plant species, (ii) photosynthetic variants from algae, and (iii) reconstruction of no longer extant types of plant photosynthesis. Here, we argue for a novel approach that outsources crop photosynthesis to a cyanobacterium that is amenable to adaptive evolution. This system offers numerous advantages, including a short generation time, virtually unlimited population sizes and high mutation rates, together with a versatile toolbox for genetic manipulation. On such a synthetic bacterial platform, 10 000 years of (crop) plant evolution can be recapitulated within weeks. Limitations of this system arise from its unicellular nature, which cannot reproduce all aspects of crop photosynthesis. But successful establishment of such a bacterial host for crop photosynthesis promises not only to enhance the performance of eukaryotic photosynthesis but will also reveal novel facets of the molecular basis of photosynthetic flexibility. [ABSTRACT FROM AUTHOR]

Light induced photosystem (PS)II photoinhibition inactivates and irreversibly damages the reaction center protein(s) but the light harvesting complexes continue the collection of light energy. Here we addressed the consequences of such a situation on thylakoid light harvesting and electron transfer reactions. For this purpose, Arabidopsis thaliana leaves were subjected to investigation of the function and regulation of the photosynthetic machinery after a distinct portion of PSII centers had experienced photoinhibition in the presence and absence of Lincomycin (Lin), a commonly used agent to block the repair of damaged PSII centers. In the absence of Lin, photoinhibition increased the relative excitation of PSII and decreased NPQ, together enhancing the electron transfer from still functional PSII centers to PSI. In contrast, in the presence of Lin, PSII photoinhibition increased the relative excitation of PSI and led to strong oxidation of the electron transfer chain. We hypothesize that plants are able to minimize the detrimental effects of high-light illumination on PSII by modulating the energy and electron transfer, but lose such a capability if the repair cycle is arrested. It is further hypothesized that dynamic regulation of the LHCII system has a pivotal role in the control of excitation energy transfer upon PSII damage and repair cycle to maintain the photosynthesis safe and efficient. • The consequences of PSII photoinhibition are mitigated by the regulatory mechanisms of photosynthetic light reactions • When the PSII repair cycle is functional, the mechanisms to mitigate the consequences of photoinhibition are efficient • When the repair cycle is arrested, the ability to mitigate the consequences of PSII photoinhibition is lost • Photoinhibition-related quenching (qI) is developed by different mechanisms depending on the function of the PSII repair cycle [ABSTRACT FROM AUTHOR]

Chemical thinning of apple fruitlets is an important practice as it reduces the natural fruit load and, therefore, increases the size of the final fruit for commercial markets. In apples, one chemical thinner used is Metamitron, which is sold as the commercial product Brevis® (Adama, Ashdod, Israel). This thinner inhibits the electron transfer between Photosystem II and Quinone-b within light reactions of photosynthesis. In this study, we investigated the responses of two apple cultivars—Golden Delicious and Top Red—and photosynthetic light reactions after administration of Brevis®. The analysis revealed that the presence of the inhibitor affects both cultivars' energetic status. The kinetics of the photoprotective mechanism's sub-processes are attenuated in both cultivars, but this seems more severe in the Top Red cultivar. State transitions of the antenna and Photosystem II repair cycle are decreased substantially when the Metamitron concentration is above 0.6% in the Top Red cultivar but not in the Golden Delicious cultivar. These attenuations result from a biased absorbed energy distribution between photochemistry and photoprotection pathways in the two cultivars. We suggest that Metamitron inadvertently interacts with photoprotective mechanism-related enzymes in chloroplasts of apple tree leaves. Specifically, we hypothesize that it may interact with the kinases responsible for the induction of state transitions and the Photosystem II repair cycle. [ABSTRACT FROM AUTHOR]