Photosynthetic Antenna Size Regulation as an Essential Mechanism of Higher Plants Acclimation to Biotic and Abiotic Factors: The Role of the Chloroplast Plastoquinone Pool and Hydrogen Peroxide

The present chapter describes the mechanisms of reactive oxygen species formation in photosynthetic reactions and the functional significance of reactive oxygen species as signal messengers in photosynthetic cells of plants. Attention is given to the acclimation mechanisms of higher plants to abiotic and biotic factors such as increased light, drought, soil salinity and colonization of plants by rhizosphere microorganisms. Special attention is paid to the reactions of reactive oxygen species with the components of the chloroplasts plastoquinone pool leading to production of hydrogen peroxide as a signal molecule, which is involved in acclimation of plants to these stress conditions. The chapter also presents the data demonstrating that regulation of the size of the light-harvesting antenna of photosystem II is one of the universal mechanisms of the structural and functional reorganization of the photosynthetic apparatus of higher plants exposed to the abiotic and biotic factors. These data were obtained for both model Arabidopsis (Arabidopsis thaliana) plants as well as for agricultural barley (Hordeum vulgare) plants. It is hypothesized that hydrogen peroxide, produced with involvement of the plastoquinone pool components, plays the role of a signaling molecule for regulation of the photosystem II antenna size in higher plants when environmental conditions change

O2 reduction by photosystem I involves phylloquinone under steady-state illumination

AbstractO2 reduction was investigated in photosystem I (PSI) complexes isolated from cyanobacteria Synechocystis sp. PCC 6803 wild type (WT) and menB mutant strain, which is unable to synthesize phylloquinone and contains plastoquinone at the quinone-binding site A1. PSI complexes from WT and menB mutant exhibited different dependencies of O2 reduction on light intensity, namely, the values of O2 reduction rate in WT did not reach saturation at high intensities, in contrast to the values in menB mutant. The obtained results suggest the immediate phylloquinone involvement in the light-induced O2 reduction by PSI

Features of Photosynthesis in <i>Arabidopsis thaliana</i> 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

Long-term acclimatory response to excess excitation energy: evidence for a role of hydrogen peroxide in the regulation of photosystem II antenna size

Higher plants possess the ability to trigger a long-term acclimatory response to different environmental light conditions through the regulation of the light-harvesting antenna size of photosystem II. The present study provides an insight into the molecular nature of the signal which initiates the high light-mediated response of a reduction in antenna size. Using barley (Hordeum vulgare) plants, it is shown (i) that the light-harvesting antenna size is not reduced in high light with a low hydrogen peroxide content in the leaves; and (ii) that a decrease in the antenna size is observed in low light in the presence of an elevated concentration of hydrogen peroxide in the leaves. In particular, it has been demonstrated that the ability to reduce the antenna size of photosystem II in high light is restricted to photosynthetic apparatus with a reduced level of the plastoquinone pool and with a low hydrogen peroxide content. Conversely, the reduction of antenna size in low light is induced in photosynthetic apparatus possessing elevated hydrogen peroxide even when the reduction level of the plastoquinone pool is low. Hydrogen peroxide affects the relative abundance of the antenna proteins that modulate the antenna size of photosystem II through a down-regulation of the corresponding lhcb mRNA levels. This work shows that hydrogen peroxide contributes to triggering the photosynthetic apparatus response for the reduction of the antenna size of photosystem II by being the molecular signal for the long-term acclimation of plants to high light