Dependence of state transitions on illumination time in arabidopsis and barley plants.

Solar energy absorbed by plants can be redistributed between photosystems in the process termed "state transitions" (ST). ST represents a reversible transition of a part of the PSII light harvesting complex (L-LHCII) between photosystem II (PSII) and photosystem I (PSI) in response to the change in light spectral composition. The present work demonstrates a slower development of the state 1 to state 2 transition, i.e., L-LHCII transition from PSII to PSI, in the leaves of dicotyledonous arabidopsis (Arabidopsis thaliana) than in the leaves of monocotyledonous barley (Hordeum vulgare) plants that was assessed by the measurement of chlorophyll a fluorescence at 77 K and of chlorophyll a fluorescence at room temperature. It is known that the first step of the state 1 to state 2 transition is phosphorylation of Lhcb1 and Lhcb2 proteins; however, we detected no difference in the rate of accumulation of these phosphorylated proteins in the studied plants. Therefore, the parameters, which possibly affect the second step of this transition, i.e., the migration of L-LHCII complexes along the thylakoid membrane, were evaluated. Spin-probe EPR measurements demonstrated that the thylakoid membranes viscosity in arabidopsis was higher compared to that in barley. Moreover, confocal microscopy data evidenced the different size of chloroplasts in the leaves of the studied species being larger in arabidopsis. The obtained results suggest that the observed deference in the development of the state 1 to state 2 transition in arabidopsis and barley is caused by the slower L-LHCII migration rate in arabidopsis than in barley plants rather than by the difference in the Lhcb1 and Lhcb2 phosphorylation.

Peculiarities of DNP-INT and DBMIB as inhibitors of the photosynthetic electron transport.

Inhibitory analysis is a useful tool for studying cytochrome b6f complex in the photosynthetic electron transport chain. Here, we examine the inhibitory efficiency of two widely used inhibitors of the plastoquinol oxidation in the cytochrome b6f complex, namely 2,4-dinitrophenyl ether of 2-iodo-4-nitrothymol (DNP-INT) and 2,5-dibromo-3-methyl-6-isopropylbenzoquinone (DBMIB). Using isolated thylakoids from pea and arabidopsis, we demonstrate that inhibitory activity of DNP-INT and DBMIB is enhanced by increasing irradiance, and this effect is due to the increase in the rate of electron transport. However, the accumulation of protons in the thylakoid lumen at low light intensity has opposite effects on the inhibitory activity of DNP-INT and DBMIB, namely increasing the activity of DNP-INT and restricting the activity of DBMIB. These results allow for the refinement of the conditions under which the use of these inhibitors leads to the complete inhibition of plastoquinol oxidation in the cytochrome b6f complex, thereby broadening our understanding of the operation of the cytochrome b6f complex under conditions of steady-state electron transport.

Features of Electrochemical Hydrogen Pump Based on Irradiated Proton Exchange Membrane.

An electrochemical hydrogen pump (EHP) with a proton exchange membrane (PEM) used as part of fusion cycle systems successfully combines the processes of hydrogen extraction, purification and compression in a single device. This work comprises a novel study of the effect of ionizing radiation on the properties of the PEM as part of the EHP. Radiation exposure leads to nonspecific degradation of membranes, changes in their structure, and destruction of side and matrix chains. The findings from this work reveal that the replacement of sulfate groups in the membrane structure with carboxyl and hydrophilic groups leads to a decrease in conductivity from 0.115 to 0.103 S cm-1, which is reflected in halving the device performance at a temperature of 30 °C. The shift of the ionomer peak of small-angle X-ray scattering curves from 3.1 to 4.4 nm and the absence of changes in the water uptake suggested structural changes in the PEM after the irradiation. Increasing the EHP operating temperature minimized the effect of membrane irradiation on the pump performance, but enhanced membrane drying at low pressure and 50 °C, which caused a current density drop from 0.52 to 0.32 A·cm-2 at 0.5 V.

Superoxide Anion Radical Generation in Photosynthetic Electron Transport Chain.

This review analyzes data available in the literature on the rates, characteristics, and mechanisms of oxygen reduction to a superoxide anion radical at the sites of photosynthetic electron transport chain where this reduction has been established. The existing assumptions about the role of the components of these sites in this process are critically examined using thermodynamic approaches and results of the recent studies. The process of O2 reduction at the acceptor side of PSI, which is considered the main site of this process taking place in the photosynthetic chain, is described in detail. Evolution of photosynthetic apparatus in the context of controlling the leakage of electrons to O2 is explored. The reasons limiting application of the results obtained with the isolated segments of the photosynthetic chain to estimate the rates of O2 reduction at the corresponding sites in the intact thylakoid membrane are discussed.

Features of Photosynthesis in Arabidopsis thaliana Plants with Knocked Out Gene of Alpha Carbonic Anhydrase 2.

The knockout of the At2g28210 gene encoding α-carbonic anhydrase 2 (α-CA2) in Arabidopsis thaliana (Columbia) led to alterations in photosynthetic processes. The effective quantum yields of both photosystem II (PSII) and photosystem I (PSI) were higher in α-carbonic anhydrase 2 knockout plants (α-CA2-KO), and the reduction state of plastoquinone pool was lower than in wild type (WT). The electron transport rate in the isolated thylakoids measured with methyl viologen was higher in α-CA2-KO plants. The amounts of reaction centers of PSII and PSI were similar in WT and α-CA2-KO plants. The non-photochemical quenching of chlorophyll a fluorescence in α-CA2-KO leaves was lower at the beginning of illumination, but became slightly higher than in WT leaves when the steady state was achieved. The degree of state transitions in the leaves was lower in α-CA2-KO than in WT plants. Measurements of the electrochromic carotenoid absorbance shift (ECS) revealed that the light-dependent pH gradient (ΔpH) across the thylakoid membrane was lower in the leaves of α-CA2-KO plants than in WT plants. The starch content in α-CA2-KO leaves was lower than in WT plants. The expression levels of the genes encoding chloroplast CAs in α-CA2-KO changed noticeably, whereas the expression levels of genes of cytoplasmic CAs remained almost the same. It is proposed that α-CA2 may be situated in the chloroplasts.

Controlling Magnetism with Light in a Zero Orbital Angular Momentum Antiferromagnet.

Antiferromagnetic materials feature intrinsic ultrafast spin dynamics, making them ideal candidates for future magnonic devices operating at THz frequencies. A major focus of current research is the investigation of optical methods for the efficient generation of coherent magnons in antiferromagnetic insulators. In magnetic lattices endowed with orbital angular momentum, spin-orbit coupling enables spin dynamics through the resonant excitation of low-energy electric dipoles such as phonons and orbital resonances which interact with spins. However, in magnetic systems with zero orbital angular momentum, microscopic pathways for the resonant and low-energy optical excitation of coherent spin dynamics are lacking. Here, we consider experimentally the relative merits of electronic and vibrational excitations for the optical control of zero orbital angular momentum magnets, focusing on a limit case: the antiferromagnet manganese phosphorous trisulfide (MnPS_{3}), constituted by orbital singlet Mn^{2+} ions. We study the correlation of spins with two types of excitations within its band gap: a bound electron orbital excitation from the singlet orbital ground state of Mn^{2+} into an orbital triplet state, which causes coherent spin precession, and a vibrational excitation of the crystal field that causes thermal spin disorder. Our findings cast orbital transitions as key targets for magnetic control in insulators constituted by magnetic centers of zero orbital angular momentum.

Axion Dark Matter Search around 4.55 µeV with Dine-Fischler-Srednicki-Zhitnitskii Sensitivity.

We report an axion dark matter search at Dine-Fischler-Srednicki-Zhitnitskii sensitivity with the CAPP-12TB haloscope, assuming axions contribute 100% of the local dark matter density. The search excluded the axion-photon coupling g_{aγγ} down to about 6.2×10^{-16} GeV^{-1} over the axion mass range between 4.51 and 4.59 µeV at a 90% confidence level. The achieved experimental sensitivity can also exclude Kim-Shifman-Vainshtein-Zakharov axion dark matter that makes up just 13% of the local dark matter density. The CAPP-12TB haloscope will continue the search over a wide range of axion masses.

Light harvesting regulation: A versatile network of key components operating under various stress conditions in higher plants.

Light harvesting is finetuned through two main strategies controlling energy transfer to the reaction centers of photosystems: i) regulating the amount of light energy at the absorption level, ii) regulating the amount of the absorbed energy at the utilization level. The first strategy is ensured by changes in the cross-section, i.e., the size of the photosynthetic antenna. These changes can occur in a short-term (state transitions) or long-term way (changes in antenna protein biosynthesis) depending on the light conditions. The interrelation of these two ways is still underexplored. Regulating light absorption through the long-term modulation of photosystem II antenna size has been mostly considered as an acclimatory mechanism to light conditions. The present review highlights that this mechanism represents one of the most versatile mechanisms of higher plant acclimation to various conditions including drought, salinity, temperature changes, and even biotic factors. We suggest that H2O2 is the universal signaling agent providing the switch from the short-term to long-term modulation of photosystem II antenna size under these factors. The second strategy of light harvesting is represented by redirecting energy to waste mainly via thermal energy dissipation in the photosystem II antenna in high light through PsbS protein and xanthophyll cycle. In the latter case, H2O2 also plays a considerable role. This circumstance may explain the maintenance of the appropriate level of zeaxanthin not only upon high light but also upon other stress factors. Thus, the review emphasizes the significance of both strategies for ensuring plant sustainability under various environmental conditions.

The Role of Carbonic Anhydrase αCA4 in Photosynthetic Reactions in Arabidopsis thaliana Studied, Using the Cas9 and T-DNA Induced Mutations in Its Gene.

An homozygous mutant line of Arabidopsis thaliana with a knocked out At4g20990 gene encoding thylakoid carbonic anhydrase αCA4 was created using a CRISPR/Cas9 genome editing system. The effects of the mutation were compared with those in two mutant lines obtained by the T-DNA insertion method. In αCA4 knockouts of all three lines, non-photochemical quenching of chlorophyll a fluorescence was lower than in the wild type (WT) plants due to a decrease in its energy-dependent component. The αCA4 knockout also affected the level of expression of the genes encoding all proteins of the PSII light harvesting antennae, the genes encoding cytoplasmic and thylakoid CAs and the genes induced by plant immune signals. The production level of starch synthesis during the light period, as well as the level of its utilization during the darkness, were significantly higher in these mutants than in WT plants. These data confirm that the previously observed differences between insertional mutants and WT plants were not the result of the negative effects of T-DNA insertion transgenesis but the results of αCA4 gene knockout. Overall, the data indicate the involvement of αCA4 in the photosynthetic reactions in the thylakoid membrane, in particular in processes associated with the protection of higher plants' photosynthetic apparatus from photoinhibition.

'Regulation of photosynthesis' session of the Russian Photobiology Society 9th Congress (Shepsi, September 12-19, 2021).

The 'Regulation of Photosynthesis' session was one of the largest events of the Congress, with 17 talks and 9 poster presentations. It covered many topics on the cutting edge of modern photosynthesis research: from the electron transport within the cytochrome b 6 f-complex to the novel role of carbonic anhydrase in the regulation of pH within the thylakoid lumen, the expanding role of retrograde signaling in all aspects of plant life including the regulation of cell-to-cell traffic via plasmodesmata, and the development of new non-invasive techniques to measure photosynthetic response. Here, a short survey of the presented reports is given.

Cooperative pathway of O2 reduction to H2O2 in chloroplast thylakoid membrane: new insight into the Mehler reaction.

Oxygen reduction in chloroplasts in the light was discovered by (Mehler Arch Biochem Biophys 33:65-77, 1951) as production of hydrogen peroxide. Later, it was shown that the primary product of the oxygen reduction is superoxide radical produced in thylakoids by one-electron transfer from reduced components of photosynthetic electron transport chain to O2 molecule. For a long time, the formation of hydrogen peroxide was considered to be a result of disproportionation of superoxide radicals in chloroplast stroma. Here, we overview a growing number of evidence indicating on another one, additional to disproportionation, pathway of hydrogen peroxide formation in chloroplasts, namely its formation in thylakoid membrane due to reaction of superoxide radical generated in the membrane with the reduced plastoquinone molecule, plastohydroquinone. Since various components of photosynthetic electron transport chain (primarily photosystem I) can supply superoxide radicals to this reaction, we refer this two-step O2 photoreduction to H2O2 as a cooperative process. The significance of hydrogen peroxide production via this pathway for redox signaling and scavenging of reactive oxygen species is discussed.

Effect of CO2 Content in Air on the Activity of Carbonic Anhydrases in Cytoplasm, Chloroplasts, and Mitochondria and the Expression Level of Carbonic Anhydrase Genes of the α- and ß-Families in Arabidopsis thaliana Leaves.

The carbonic anhydrase (CA) activities of the preparations of cytoplasm, mitochondria, chloroplast stroma, and chloroplast thylakoids, as well as the expression levels of genes encoding αCA1, αCA2, αCA4, ßCA1, ßCA2, ßCA3, ßCA4, ßCA5, and ßCA6, were measured in the leaves of Arabidopsis thaliana plants, acclimated to different CO2 content in the air: low (150 ppm, lCO2), normal (450 ppm, nCO2), and high (1200 ppm, hCO2). To evaluate the photosynthetic apparatus operation, the carbon assimilation and chlorophyll a fluorescence were measured under the same conditions. It was found that the CA activities of the preparations of cytoplasm, chloroplast stroma, and chloroplast thylakoids measured after two weeks of acclimation were higher, the lower CO2 concentration in the air. That was preceded by an increase in the expression levels of genes encoding the cytoplasmic form of ßCA1, and other cytoplasmic CAs, ßCA2, ßCA3, and ßCA4, as well as of the chloroplast CAs, ßCA5, and the stromal forms of ßCA1 in a short-term range 1-2 days after the beginning of the acclimation. The dependence on the CO2 content in the air was most noticeable for the CA activity of the preparations of the stroma; it was two orders higher in lCO2 plants than in hCO2 plants. The CA activity of thylakoid membranes from lCO2 plants was higher than that in nCO2 and hCO2 plants; however, in these plants, a significant increase in the expression levels of the genes encoding αCA2 and αCA4 located in thylakoid membranes was not observed. The CA activity of mitochondria and the expression level of the mitochondrial ßCA6 gene did not depend on the content of carbon dioxide. Taken together, the data implied that in the higher plants, the supply of inorganic carbon to carboxylation sites is carried out with the cooperative functioning of CAs located in the cytoplasm and CAs located in the chloroplasts.

Exceptional warming over the Barents area.

In recent decades, surface air temperature (SAT) data from Global reanalyses points to maximum warming over the northern Barents area. However, a scarcity of observations hampers the confidence of reanalyses in this Arctic hotspot region. Here, we study the warming over the past 20-40 years based on new available SAT observations and a quality controlled comprehensive SAT dataset from the northern archipelagos in the Barents Sea. We identify a statistically significant record-high annual warming of up to 2.7 °C per decade, with a maximum in autumn of up to 4.0 °C per decade. Our results are compared with the most recent global and Arctic regional reanalysis data sets, as well as remote sensing data records of sea ice concentration (SIC), sea surface temperature (SST) and high-resolution ice charts. The warming pattern is primarily consistent with reductions in sea ice cover and confirms the general spatial and temporal patterns represented by reanalyses. However, our findings suggest even a stronger rate of warming and SIC-SAT relation than was known in this region until now.

Inhibition of plastoquinol oxidation at the cytochrome b6f complex by dinitrophenyl ether of iodonitrothymol (DNP-INT) depends on irradiance and H+ uptake by thylakoid membranes.

Inhibitory analysis is a useful tool for studying reactions in the photosynthetic apparatus. After introducing by Aachim Trebst in 1978, dinitrophenylether of iodonitrothymol (DNP-INT), a competitive inhibitor of plastoquinol oxidation at the cytochrome (cyt.) b6f complex, has been widely applied to study reactions occurring in the plastoquinone pool and the cyt. b6f complex. Here we examine the inhibitory efficiency of DNP-INT by implementing three approaches to estimate the extent of blockage of electron flow from the plastoquinone pool to photosystem I in isolated thylakoids from spinach (Spinacia oleracea). We confirm that DNP-INT is a potent inhibitor of electron flow to photosystem I and demonstrate that inhibitory action of DNP-INT depends on irradiance and H+ uptake by thylakoid membranes. Based on these findings, we infer that affinity of the quinol-oxidizing site of the cyt. b6f complex to DNP-INT is increased in the light due to hydrogen bonding between DNP-INT molecules and acidic amino acid residue(s), which is (are) protonated in the light.

Unsolved Problems of Carbonic Anhydrases Functioning in Photosynthetic Cells of Higher C3 Plants.

The review presents current data on carbonic anhydrases found in various compartments of photosynthetic cells of higher plants. The available data on expression of genes some of carbonic anhydrases and its dependence on environmental factors and plant age are considered. The existing hypotheses on the functions of carbonic anhydrases of plasma membrane, cytoplasm, as well as of stroma and thylakoids of chloroplast, first of all, the hypothesis on participation of these enzymes in supplying carbon dioxide molecules to ribulose-bisphosphate carboxylase (Rubisco) are analyzed. Difficulties of establishing physiological role of the plant cell carbonic anhydrase are discussed in detail.

Phylloquinone is the principal Mehler reaction site within photosystem I in high light.

Photosynthesis is a vital process, responsible for fixing carbon dioxide, and producing most of the organic matter on the planet. However, photosynthesis has some inherent limitations in utilizing solar energy, and a part of the energy absorbed is lost in the reduction of O2 to produce the superoxide radical (O2•-) via the Mehler reaction, which occurs principally within photosystem I (PSI). For decades, O2 reduction within PSI was assumed to take place solely in the distal iron-sulfur clusters rather than within the two asymmetrical cofactor branches. Here, we demonstrate that under high irradiance, O2 photoreduction by PSI primarily takes place at the phylloquinone of one of the branches (the A-branch). This conclusion derives from the light dependency of the O2 photoreduction rate constant in fully mature wild-type PSI from Chlamydomonas reinhardtii, complexes lacking iron-sulfur clusters, and a mutant PSI, in which phyllosemiquinone at the A-branch has a significantly longer lifetime. We suggest that the Mehler reaction at the phylloquinone site serves as a release valve under conditions where both the iron-sulfur clusters of PSI and the mobile ferredoxin pool are highly reduced.

Alpha Carbonic Anhydrase 5 Mediates Stimulation of ATP Synthesis by Bicarbonate in Isolated Arabidopsis Thylakoids.

We studied bicarbonate-induced stimulation of photophosphorylation in thylakoids isolated from leaves of Arabidopsis thaliana plants. This stimulation was not observed in thylakoids of wild-type in the presence of mafenide, a soluble carbonic anhydrase inhibitor, and was absent in thylakoids of two mutant lines lacking the gene encoding alpha carbonic anhydrase 5 (αCA5). Using mass spectrometry, we revealed the presence of αCA5 in stromal thylakoid membranes of wild-type plants. A possible mechanism of the photophosphorylation stimulation by bicarbonate that involves αCA5 is proposed.

Three overlooked photosynthesis papers of Otto Warburg (1883-1970), published in the 1940s in German and in Russian, on light-driven water oxidation coupled to benzoquinone reduction.

After a brief background on Otto Heinrich Warburg (1883-1970), and some of his selected research, we provide highlights, in English, of three of his papers in the 1940s-unknown to many as they were not originally published in English. They are: two brief reports on Photosynthesis, with Wilhelm Lüttgens, originally published in German, in 1944: 'Experiment on assimilation of carbonic acid'; and 'Further experiments on carbon dioxide assimilation'. This is followed by a regular paper, originally published in Russian, in 1946: 'The photochemical reduction of quinone in green granules'. Since the 1944 reports discussed here are very short, their translations are included in the Appendix, but that of the 1946 paper is provided in the Supplementary Material. In all three reports, Warburg provides the first evidence for and elaborates on light-driven water oxidation coupled to reduction of added benzoquinone. These largely overlooked studies of Warburg are in stark contrast to Warburg's well-known error in assigning the origin of the photosynthetically formed dioxygen to carbonate.

Controlling the anisotropy of a van der Waals antiferromagnet with light.

Van der Waals magnets provide an ideal playground to explore the fundamentals of low-dimensional magnetism and open opportunities for ultrathin spin-processing devices. The Mermin-Wagner theorem dictates that as in reduced dimensions isotropic spin interactions cannot retain long-range correlations, the long-range spin order is stabilized by magnetic anisotropy. Here, using ultrashort pulses of light, we control magnetic anisotropy in the two-dimensional van der Waals antiferromagnet NiPS3 Tuning the photon energy in resonance with an orbital transition between crystal field split levels of the nickel ions, we demonstrate the selective activation of a subterahertz magnon mode with markedly two-dimensional behavior. The pump polarization control of the magnon amplitude confirms that the activation is governed by the photoinduced magnetic anisotropy axis emerging in response to photoexcitation of ground state electrons to states with a lower orbital symmetry. Our results establish pumping of orbital resonances as a promising route for manipulating magnetic order in low-dimensional (anti)ferromagnets.

Controlling Magnon Interaction by a Nanoscale Switch.

The ability to control and tune magnetic dissipation is a key concept of emergent spintronic technologies. Magnon scattering processes constitute a major dissipation channel in nanomagnets, redefine their response to spin torque, and hold the promise for manipulating magnetic states on the quantum level. Controlling these processes in nanomagnets, while being imperative for spintronic applications, has remained difficult to achieve. Here, we propose an approach for controlling magnon scattering by a switch that generates nonuniform magnetic field at nanoscale. We provide an experimental demonstration in magnetic tunnel junction nanodevices, consisting of a free layer and a synthetic antiferromagnet. By triggering the spin-flop transition in the synthetic antiferromagnet and utilizing its stray field, magnon interaction in the free layer is toggled. The results open up avenues for tuning nonlinearities in magnetic neuromorphic applications and for engineering coherent magnon coupling in hybrid quantum information technologies.