scholarly journals Photosynthesis: basics, history and modelling

2019 ◽  
Vol 126 (4) ◽  
pp. 511-537 ◽  
Author(s):  
Alexandrina Stirbet ◽  
Dušan Lazár ◽  
Ya Guo ◽  
Govindjee Govindjee

Abstract Background With limited agricultural land and increasing human population, it is essential to enhance overall photosynthesis and thus productivity. Oxygenic photosynthesis begins with light absorption, followed by excitation energy transfer to the reaction centres, primary photochemistry, electron and proton transport, NADPH and ATP synthesis, and then CO2 fixation (Calvin–Benson cycle, as well as Hatch–Slack cycle). Here we cover some of the discoveries related to this process, such as the existence of two light reactions and two photosystems connected by an electron transport ‘chain’ (the Z-scheme), chemiosmotic hypothesis for ATP synthesis, water oxidation clock for oxygen evolution, steps for carbon fixation, and finally the diverse mechanisms of regulatory processes, such as ‘state transitions’ and ‘non-photochemical quenching’ of the excited state of chlorophyll a. Scope In this review, we emphasize that mathematical modelling is a highly valuable tool in understanding and making predictions regarding photosynthesis. Different mathematical models have been used to examine current theories on diverse photosynthetic processes; these have been validated through simulation(s) of available experimental data, such as chlorophyll a fluorescence induction, measured with fluorometers using continuous (or modulated) exciting light, and absorbance changes at 820 nm (ΔA820) related to redox changes in P700, the reaction centre of photosystem I. Conclusions We highlight here the important role of modelling in deciphering and untangling complex photosynthesis processes taking place simultaneously, as well as in predicting possible ways to obtain higher biomass and productivity in plants, algae and cyanobacteria.

2017 ◽  
Author(s):  
Tanai Cardona

AbstractWhen and how oxygenic photosynthesis originated remains controversial. Wide uncertainties exist for the earliest detection of biogenic oxygen in the geochemical record or the origin of water oxidation in ancestral lineages of the phylum Cyanobacteria. A unique trait of oxygenic photosynthesis is that the process uses a Type I reaction centre with a heterodimeric core, also known as Photosystem I, made of two distinct but homologous subunits, PsaA and PsaB. In contrast, all other known Type I reaction centres in anoxygenic phototrophs have a homodimeric core. A compelling hypothesis for the evolution of a heterodimeric Type I reaction centre is that the gene duplication that allowed the divergence of PsaA and PsaB was an adaptation to incorporate photoprotective mechanisms against the formation of reactive oxygen species, therefore occurring after the origin of water oxidation to oxygen. Here I show, using sequence comparisons and Bayesian relaxed molecular clocks that this gene duplication event may have occurred in the early Archean more than 3.4 billion years ago, long before the most recent common ancestor of crown group Cyanobacteria and the Great Oxidation Event. If the origin of water oxidation predated this gene duplication event, then that would place primordial forms of oxygenic photosynthesis at a very early stage in the evolutionary history of life.


2014 ◽  
Vol 369 (1640) ◽  
pp. 20130221 ◽  
Author(s):  
Luca Dall'Osto ◽  
Stefano Cazzaniga ◽  
Masamitsu Wada ◽  
Roberto Bassi

Over-excitation of photosynthetic apparatus causing photoinhibition is counteracted by non-photochemical quenching (NPQ) of chlorophyll fluorescence, dissipating excess absorbed energy into heat. The PsbS protein plays a key role in this process, thus making the PsbS-less npq4 mutant unable to carry out qE, the major and most rapid component of NPQ. It was proposed that npq4 does perform qE-type quenching, although at lower rate than WT Arabidopsis . Here, we investigated the kinetics of NPQ in PsbS-depleted mutants of Arabidopsis . We show that red light was less effective than white light in decreasing maximal fluorescence in npq4 mutants. Also, the kinetics of fluorescence dark recovery included a decay component, qM, exhibiting the same amplitude and half-life in both WT and npq4 mutants. This component was uncoupler-sensitive and unaffected by photosystem II repair or mitochondrial ATP synthesis inhibitors. Targeted reverse genetic analysis showed that traits affecting composition of the photosynthetic apparatus, carotenoid biosynthesis and state transitions did not affect qM. This was depleted in the npq4phot2 mutant which is impaired in chloroplast photorelocation, implying that fluorescence decay, previously described as a quenching component in npq4 is, in fact, the result of decreased photon absorption caused by chloroplast relocation rather than a change in the activity of quenching reactions.


1998 ◽  
Vol 76 (6) ◽  
pp. 1018-1024
Author(s):  
Lucina C van Ginkel ◽  
Hidde BA Prins

By the process of pH polarity, several submersed angiosperms can use bicarbonate as carbon source for photosynthesis. Under conditions of relatively high light intensity and low CO2 availability, the pH of the apoplast and unstirred layer becomes acid at one side of the leaf and alkaline at the other. In the acid region, bicarbonate is converted into CO2, which diffuses into the leaf where it is fixed. Previous experiments on the light-dependent reduction of extracellular electron acceptors led to the hypothesis of redox regulation. Under conditions of high light and low CO2, excess reducing power in the chloroplast was supposed to be shuttled to the cytoplasm where it can upregulate the plasma membrane proton pump, leading to activation of polarity. Chlorophyll a fluorescence is an indicator for photosynthetic electron transport, the energization of thylakoids, and the reoxidation of chloroplast NADPH. It was used therefore to test redox regulation in vivo in Potamogeton lucens L. leaves. The fluoresence parameter, qP, an indicator for photochemical quenching and NADPH reoxidation, appeared to be rather insensitive to the inorganic carbon concentration and to the presence or absence of polarity. In contrast, qN, an indicator for non-photochemical quenching related to thylakoid energization, photoinhibition, and state transitions, increased under conditions of low CO2 - high light and polarity. Taken together the data show polarity to be an effective mechanism to make bicarbonate accessible as carbon source and seem to agree with the idea of redox regulation of pH polarity.Key words: bicarbonate utilization, chlorophyll a fluoresence, pH polarity, redox regulation, Potamogeton lucens, submerged aquatic macrophyte.


Author(s):  
Wolfgang Junge

AbstractCyanobacteria and plants carry out oxygenic photosynthesis. They use water to generate the atmospheric oxygen we breathe and carbon dioxide to produce the biomass serving as food, feed, fibre and fuel. This paper scans the emergence of structural and mechanistic understanding of oxygen evolution over the past 50 years. It reviews speculative concepts and the stepped insight provided by novel experimental and theoretical techniques. Driven by sunlight photosystem II oxidizes the catalyst of water oxidation, a hetero-metallic Mn4CaO5(H2O)4 cluster. Mn3Ca are arranged in cubanoid and one Mn dangles out. By accumulation of four oxidizing equivalents before initiating dioxygen formation it matches the four-electron chemistry from water to dioxygen to the one-electron chemistry of the photo-sensitizer. Potentially harmful intermediates are thereby occluded in space and time. Kinetic signatures of the catalytic cluster and its partners in the photo-reaction centre have been resolved, in the frequency domain ranging from acoustic waves via infra-red to X-ray radiation, and in the time domain from nano- to milli-seconds. X-ray structures to a resolution of 1.9 Å are available. Even time resolved X-ray structures have been obtained by clocking the reaction cycle by flashes of light and diffraction with femtosecond X-ray pulses. The terminal reaction cascade from two molecules of water to dioxygen involves the transfer of four electrons, two protons, one dioxygen and one water. A rigorous mechanistic analysis is challenging because of the kinetic enslaving at millisecond duration of six partial reactions (4e−, 1H+, 1O2). For the time being a peroxide-intermediate in the reaction cascade to dioxygen has been in focus, both experimentally and by quantum chemistry. Homo sapiens has relied on burning the products of oxygenic photosynthesis, recent and fossil. Mankind's total energy consumption amounts to almost one-fourth of the global photosynthetic productivity. If the average power consumption equalled one of those nations with the highest consumption per capita it was four times greater and matched the total productivity. It is obvious that biomass should be harvested for food, feed, fibre and platform chemicals rather than for fuel.


2019 ◽  
Vol 476 (20) ◽  
pp. 2981-3018 ◽  
Author(s):  
Petar H. Lambrev ◽  
Parveen Akhtar

Abstract The light reactions of photosynthesis are hosted and regulated by the chloroplast thylakoid membrane (TM) — the central structural component of the photosynthetic apparatus of plants and algae. The two-dimensional and three-dimensional arrangement of the lipid–protein assemblies, aka macroorganisation, and its dynamic responses to the fluctuating physiological environment, aka flexibility, are the subject of this review. An emphasis is given on the information obtainable by spectroscopic approaches, especially circular dichroism (CD). We briefly summarise the current knowledge of the composition and three-dimensional architecture of the granal TMs in plants and the supramolecular organisation of Photosystem II and light-harvesting complex II therein. We next acquaint the non-specialist reader with the fundamentals of CD spectroscopy, recent advances such as anisotropic CD, and applications for studying the structure and macroorganisation of photosynthetic complexes and membranes. Special attention is given to the structural and functional flexibility of light-harvesting complex II in vitro as revealed by CD and fluorescence spectroscopy. We give an account of the dynamic changes in membrane macroorganisation associated with the light-adaptation of the photosynthetic apparatus and the regulation of the excitation energy flow by state transitions and non-photochemical quenching.


2021 ◽  
Vol 12 ◽  
Author(s):  
Gábor Bernát ◽  
Tomáš Zavřel ◽  
Eva Kotabová ◽  
László Kovács ◽  
Gábor Steinbach ◽  
...  

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.


2006 ◽  
Vol 33 (1) ◽  
pp. 9 ◽  
Author(s):  
Dušan Lazár

Chlorophyll a fluorescence rise caused by illumination of photosynthetic samples by high intensity of exciting light, the O–J–I–P (O–I1–I2–P) transient, is reviewed here. First, basic information about chlorophyll a fluorescence is given, followed by a description of instrumental set-ups, nomenclature of the transient, and samples used for the measurements. The review mainly focuses on the explanation of particular steps of the transient based on experimental and theoretical results, published since a last review on chlorophyll a fluorescence induction [Lazár D (1999) Biochimica et Biophysica Acta 1412, 1–28]. In addition to ‘old’ concepts (e.g. changes in redox states of electron acceptors of photosystem II (PSII), effect of the donor side of PSII, fluorescence quenching by oxidised plastoquinone pool), ‘new’ approaches (e.g. electric voltage across thylakoid membranes, electron transport through the inactive branch in PSII, recombinations between PSII electron acceptors and donors, electron transport reactions after PSII, light gradient within the sample) are reviewed. The K-step, usually detected after a high-temperature stress, and other steps appearing in the transient (the H and G steps) are also discussed. Finally, some applications of the transient are also mentioned.


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