Increase in the rate of photosynthetic linear electron transport in cyanobacteruim <Emphasis Type="Italic">Synechocystis</Emphasis> sp. PCC 6803 lacking phycobilisomes

Compensating changes in the pigment apparatus of photosynthesis that resulted from a complete loss of phycobilisomes (PBS) were investigated in the cells of a PAL mutant of cyanobacterium Synechocystis sp. PCC 6803. The ratio PBS/chlorophyll calculated on the basis of the intensity of bands in the action spectra of photosynthetic activity of two photosystems in the wild strain was 1: 70 for PSII and 1: 300 for PSI. Taking into consideration the number of chlorophyll molecules per reaction center in each photosystem, these ratios could be interpreted as association of PBS with dimers of PSII and trimers of PSI as well as greater dependence of PSII as compared with PSI on light absorption by PBS. The ratio PSI/PSII determined by photochemical cross-section of the reactions of two photosystems was 3.5: 1.0 for wild strain of Synechocystis sp. PCC 6803 and 0.7: 1.0 for the PAL mutant. A fivefold increase in the relative content of PSII in pigment apparatus corresponds to a 5-fold increase in the intensity of bands at 685 and 695 nm as related to the band of PSI at 726 nm recorded in low-temperature fluorescence spectrum of the PAL mutant. Inhibition of PSII with diuron resulted in a pronounced stimulation of chlorophyll fluorescence in the PAL mutant as compared to the wild strain of Synechocystis sp. PCC 6803; these data suggested an activation of electron transfer between PSII and PSI in the mutant cells. Thus, the lack of PBS in the mutant strain of Synechocystis sp. PCC 6803 was compensated for by the higher relative content of PSII in the pigment apparatus of photosynthesis and by a rise in the rate of linear electron transport.     

Disturbed structure and function of chloroplasts during blocked biosynthesis of 5-aminolevulinic acid in the light

Xantha-702 mutant of cotton (Gossypium hirsutum L.) proved to have blocked synthesis of 5-aminolevulinic acid in the light. Accordingly, mutant leaves accumulated 2–5% chlorophyll of baseline. Mutant plants demonstrated disturbed production of pigment-protein complexes of photosystems I (PSI) and II (PSII) and generation of the chloroplast membrane system blocked at the early stages, largely, at the stages of vesicles and single short thylakoid. The functional activity of the PSI and PSII reaction centers was close to zero. Only the chlorophyll a/b light-harvesting complexes of PSI and PSII with the chlorophyll fluorescence peaks at 728 and 681 nm, respectively, were produced in the xantha-702 mutant. We propose that the genetic block of 5-aminolevunilic acid biosynthesis in the light in the xantha-702 mutant disturbs the formation and activity of the complexes of the reaction centers of PS-I and PS-II and inhibits the development of the whole membrane system of chloroplasts.     

Effects of heat stress on photosynthetic electron transport in a marine cyanobacterium <Emphasis Type="Italic">Arthrospira</Emphasis> sp.

Arthrospira (Spirulina) is widely used as human health food and animal feed. In cultures grown outdoors in open ponds, Arthrospira cells are subjected to various environmental stresses, such as high temperature. A better understanding of the effects of high temperature on photosynthesis may help optimize the productivity of Arthrospira cultures. In this study, the effects of heat stress on photosynthetic rate, chlorophyll a fluorescence transients, and photosystem (PS) II, PSI activities in a marine cyanobacterium Arthrospira sp. were examined. Arthrospira cells grown at 25 °C were treated for 30 min at 25 (control), 30, 34, 37, or 40 °C in the dark. Heat stress (30–37 °C) enhanced net photosynthetic O₂ evolution rate. Heat stress caused over-reduction PSII acceptor side, damage of donor side of PSII, decrease in the energetic connectivity of PSII units, and decrease in the performance of PSII. When the temperature changed from 25 to 37 °C, PSII activity decreased, while PSI activity increased, the enhancement of photosynthetic O₂ evolution was synchronized with the increase in PSI activity. When temperature was further increased to 40 °C, it induced a decrease in photosynthetic O₂ evolution rate and a more severe decrease in PSII activity, but an increase in PSI activity. These results suggest that PSI activity was the decisive factor determining the change of photosynthetic O₂ evolution when Arthrospira was exposed to a temperature from 25 to 37 °C, but then, PSII activity became the decisive factor adjusting the change of photosynthetic O₂ evolution when the temperature was increased to 40 °C.     

Growth under Red Light Enhances Photosystem II Relative to Photosystem I and Phycobilisomes in the Red Alga Porphyridium cruentum

Acclimation of the photosynthetic apparatus to light absorbed primarily by photosystem I (PSI) or by photosystem II (PSII) was studied in the unicellular red alga Porphyridium cruentum (ATCC 50161). Cultures grown under green light of 15 microeinsteins per square meter per second (PSII light; absorbed predominantly by the phycobilisomes) exhibited a PSII/PSI ratio of 0.26 ± 0.05. Under red light (PSI light; absorbed primarily by chlorophyll) of comparable quantum flux, cells contained nearly five times as many PSII per PSI (1.21 ± 0.10), and three times as many PSII per cell. About 12% of the chlorophyll was attributed to PSII in green light, 22% in white light, and 39% in red light-grown cultures. Chlorophyll antenna sizes appeared to remain constant at about 75 chlorophyll per PSII and 140 per PSI. Spectral quality had little effect on cell content or composition of the phycobilisomes, thus the number of PSII per phycobilisome was substantially greater in red light-grown cultures (4.2 ± 0.6) than in those grown under green (1.6 ± 0.3) or white light (2.9 ± 0.1). Total photosystems (PSI + PSII) per phycobilisome remained at about eight in each case. Carotenoid content and composition was little affected by the spectral composition of the growth light. Zeaxanthin comprised more than 50% (mole/mole), β-carotene about 40%, and cryptoxanthin about 4% of the carotenoid pigment. Despite marked changes in the light-harvesting apparatus, red and green light-grown cultures have generation times equal to that of cultures grown under white light of only one-third the quantum flux.     

Effects of Chromatic Adaptation on the Photochemical Apparatus of Photosynthesis in Porphyridium cruentum

Cells of Porphyridium cruentum were grown in different colors of light which would be absorbed primarily by chlorophyll (Chl) (red and blue light) or by the phycobilisomes (green or two intensities of cool-white fluorescent light), and samples of these cells were frozen to −196 C for measurements of absorption and fluorescence emission spectra. Cells grown in the high intensity white light had least of all of the photosynthetic pigments, a higher ratio of carotenoid/Chl, but essentially the same ratio of phycobilin to Chl as cells grown in the low intensity white light. The ratio of photosystem II (PSII) to photosystem I (PSI) pigments was affected by light quality; the ratios of phycobilin to Chl and of short wavelength (PSII) Chl to long wavelength (PSI) Chl were both greater in the cells grown in red or blue light.     

Examination of chlorophyll fluorescence in sulfur-deprived cells of <Emphasis Type="Italic">Chlamydomonas reinhardtii</Emphasis>

Pulse amplitude modulation fluorimetry was used to assess chlorophyll fluorescence parameters in Chlamydomonas reinhardtii cells during sulfur deprivation. A significant (fourfold) increase in the chlorophyll fluorescence yield (parameters F ₀ and F _m) normalized to the chlorophyll concentration was shown for deprived cells. The chlorophyll content did not change during the deprivation experiments. An analysis of nonphotochemical quenching of chlorophyll fluorescence indicated a considerable modification of the energy deactivation pathways in photosystem II (PSII) of sulfur-deprived cells. For example, starved cells exhibited a less pronounced pH-dependent quenching of excited states and a higher thermal dissipation of excess light energy in the reaction centers of PSII. It was also shown that the photosynthetic apparatus of starved cells is primarily in state 2 and that back transition to state 1 is suppressed. However, these changes cannot cause the discovered elevation of chlorophyll fluorescence intensity (F ₀ and F _m) in the cells under sulfur limitation. The observed increase in the chlorophyll fluorescence intensity under sulfur deprivation may be due to partial dissociation of peripheral light-harvesting complexes from the reaction centers of PSII or a malfunction of the dissipative cycle in PSII, involving cytochrome b ₅₅₉.     

Photoinhibition of photosystem I in <Emphasis Type="Italic">Nephrolepis falciformis</Emphasis> depends on reactive oxygen species generated in the chloroplast stroma

We studied how high light causes photoinhibition of photosystem I (PSI) in the shade-demanding fern Nephrolepis falciformis, in an attempt to understand the mechanism of PSI photoinhibition under natural field conditions. Intact leaves were treated with constant high light and fluctuating light. Detached leaves were treated with constant high light in the presence and absence of methyl viologen (MV). Chlorophyll fluorescence and P700 signal were determined to estimate photoinhibition. PSI was highly oxidized under high light before treatments. N. falciformis showed significantly stronger photoinhibition of PSI and PSII under constant high light than fluctuating light. These results suggest that high levels of P700 oxidation ratio cannot prevent PSI photoinhibition under high light in N. falciformis. Furthermore, photoinhibition of PSI in N. falciformis was largely accelerated in the presence of MV that promotes the production of superoxide anion radicals in the chloroplast stroma by accepting electrons from PSI. From these results, we propose that photoinhibition of PSI in N. falciformis is mainly caused by superoxide radicals generated in the chloroplast stroma, which is different from the mechanism of PSI photoinhibition in Arabidopsis thaliana and spinach. Here, we provide some new insights into the PSI photoinhibition under natural field conditions.     

Chlorophyll a fluorescence of typical desert plant Alhagi sparsifolia Shap. at two light levels

Alhagi sparsifolia Shap. is exposed to a high-irradiance environment as the main vegetation found in the forelands of the Taklamakan Desert. We investigated chlorophyll a fluorescence emission of A. sparsifolia seedlings grown under ambient (HL) and shade (LL) conditions. Our results indicated that the fluorescence intensity in the leaves was significantly higher for LL-grown plants than that under HL. High values of the maximum quantum yield of PSII for primary photochemistry (φP_o) and the quantum yield that an electron moves further than Q_A - (φE_o) in the plants under LL conditions suggested that the electron flow from Q_A - (primary quinone electron acceptors of PSII) to Q_B (secondary quinone acceptor of PSII) or Q_B - was enhanced at LL compared to natural HL conditions. The efficiency/probability with which an electron from the intersystem electron carriers was transferred to reduce end electron acceptors at the PSI acceptor side and the quantum yield for the reduction of end electron acceptors at the PSI acceptor side were opposite to φP_o, and φE_o. Thus, we concluded that the electron transport on the donor side of PSII was blocked under LL conditions, while acceptor side was inhibited at the HL conditions. The PSII activity of electron transport in the plants grown in shade was enhanced, while the energy transport from PSII to PSI was blocked compared to the plants grown at HL conditions. Furthermore, PSII activity under HL was seriously affected in midday, while the plants grown in shade enhanced their energy transport.     

Acclimation of Arabidopsis thaliana to the light environment: changes in photosynthetic function

Arabidopsis thaliana (L.) Heynh. cv. Landsberg erecta was grown under light regimes of differing spectral qualities, which results in differences in the stoichiometries of the two photosynthetic reaction centres. The acclimative value of these changes was investigated by assessing photosynthetic function in these plants when exposed to two spectrally distinct actinic lights. Plants grown in an environment enriched in far-red light were better able to make efficient use of non-saturating levels of actinic light enriched in long-wavelength red light. Simultaneous measurements of chlorophyll fluorescence and absorption changes at 820 nm indicated that differences between plants grown under alternative light regimes can be ascribed to imbalances in excitation of photosystems I and II (PSI, PSII). Measurements of chlorophyll fluorescence emission and excitation spectra at 77 K provided strong evidence that there was little or no difference in the composition or function of PSI or PSII between the two sets of plants, implying that changes in photosynthetic stoichiometry are primarily responsible for the observed differences in photosynthetic function.Abbreviations Chl chlorophyll - FR far-red light - HF highirradiance FR-enriched light (400 mol·m^–2·s^–1, RFR = 0.72) - HW high-irradiance white light (400 mol·m^–2 1·1 s^–1RFR = 1.40) - LHCI, LHCII light-harvesting complex of PSI, PSII - qO quenching of dark-level chlorophyll fluorescence - qN non-photochemical quenching of variable chlorophyll fluorescence - qP photochemical quenching of variable chlorophyll fluorescence - R red light - Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase We thank Dr. Sasha Ruban for assistance with the 77 K fluorescence measurements and for helpful discussions. This work was supported by Natural Environment Research Council Grant GR3/7571A.     

Effect of light quality on chloroplast-membrane organization and function in pea

The effect of light quality during plant growth of chloroplast membrane organization and function in peas (Pisum sativum L. cv. Alaska) was investigated. In plants grown under photosystem (PS) I-enriched (far-red enriched) illumination both the PSII/PSI stoichiometry and the electrontransport capacity ratios were high, about 1.9. In plants grown under PSII-enriched (far-red depleted) illumination both the PSII/PSI stoichiometry and the electron-transport capacity ratios were significantly lower, about 1.3. In agreement, steady-state electron-transport measurements under synchronous illumination of PSII and PSI demonstrated an excess of PSII in plants grown under far-red-enriched light. Sodium dodecylsulfate polyacrylamide gel electrophoretic analysis of chlorophyll-containing complexes showed greater relative amounts of the PSII reaction center chlorophyll-protein complex in plants grown under farred-enriched light. Additional changes were observed in the ratio of light-harvesting chlorophyll a/b protein to PSII reaction center chlorophyll-protein under the two different light-quality regimes. The results demonstrate the dynamic nature of chloroplast structure and support the notion that light quality is an important factor in the regulation of chloroplast membrane organization and-function.Abbreviations and symbols Chl chlorophyll - CPa PSII reaction center chlorophyll protein complex - CPI PSI chlorophyll protein complex - FR-D light depleted in far-red sensitizing primarily PSII - FR-E light enriched in far-red sensitizing primarily PSI - LHCP PSII light-harvesting chlorophyll a/b protein complex - P ₇₀₀ primary electron donor of PSI - PSI, PSII photosystems I and II, respectively - Q primary electron acceptor of PSII     

Effects of solar ultraviolet radiation on the potential efficiency of photosystem II in leaves of tropical plants

The effects of solar ultraviolet (UV)-B and UV-A radiation on the potential efficiency of photosystem II (PSII) in leaves of tropical plants were investigated in Panama (9°N). Shade-grown tree seedlings or detached sun leaves from the outer crown of mature trees were exposed for short periods (up to 75 min) to direct sunlight filtered through plastic or glass filters that absorbed either UV-B or UV-A+B radiation, or transmitted the complete solar spectrum. Persistent changes in potential PSII efficiency were monitored by means of the dark-adapted ratio of variable to maximum chlorophyll a fluorescence. In leaves of shade-grown tree seedlings, exposure to the complete solar spectrum resulted in a strong decrease in potential PSII efficiency, probably involving protein damage. A substantially smaller decline in the dark-adapted ratio of variable to maximum chlorophyll a fluorescence was observed when UV-B irradiation was excluded. The loss in PSII efficiency was further reduced by excluding both UV-B and UV-A light. The photoinactivation of PSII was reversible under shade conditions, but restoration of nearly full activity required at least 10 d. Repeated exposure to direct sunlight induced an increase in the pool size of xanthophyll cycle pigments and in the content of UV-absorbing vacuolar compounds. In sun leaves of mature trees, which contained high levels of UV-absorbing compounds, effects of UV-B on PSII efficiency were observed in several cases and varied with developmental age and acclimation state of the leaves. The results show that natural UV-B and UV-A radiation in the tropics may significantly contribute to photoinhibition of PSII during sun exposure in situ, particularly in shade leaves exposed to full sunlight.     

The identification of IsiA proteins binding chlorophyll <Emphasis Type="Italic">d</Emphasis> in the cyanobacterium <Emphasis Type="Italic">Acaryochloris marina</Emphasis>

The bioavailable iron in many aquatic ecosystems is extremely low, and limits the growth and photosynthetic activity of phytoplankton. In response to iron limitation, a group of chlorophyll-binding proteins known as iron stress-induced proteins are induced and serve as accessory light-harvesting components for photosystems under iron limitation. In the present study, we investigated physiological features of Acaryochloris marina in response to iron-deficient conditions. The growth doubling time under iron-deficient conditions was prolonged to ~3.4 days compared with 1.9 days under normal culture conditions, accompanied with dramatically decreased chlorophyll content. The isolation of chlorophyll-binding protein complexes using sucrose density gradient centrifugation shows six main green bands and three main fluorescence components of 712, 728, and 748 nm from the iron-deficient culture. The fluorescence components of 712 and 728 nm co-exist in the samples collected from iron-deficient and iron-replete cultures and are attributed to Chl d-binding accessory chlorophyll-binding antenna proteins and also from photosystem II. A new chlorophyll-binding protein complex with its main fluorescence peak at 748 nm was observed and enriched in the heaviest fraction from the samples collected from the iron-deficient culture only. Combining western blotting analysis using antibodies of CP47 (PSII), PsaC (PSI) and IsiA and proteomic analysis on an excised protein band at ~37 kDa, the heaviest fraction (?F6) isolated from iron-deficient culture contained Chl d-bound PSI–IsiA supercomplexes. The PSII-antenna supercomplexes isolated from iron-replete conditions showed two fluorescence peaks of 712 and 728 nm, which can be assigned as 6-transmembrane helix chlorophyll-binding antenna and photosystem II fluorescence, respectively, which is supported by protein analysis of the fractions (F5 and F6).     

Effect of Light Quality on the Composition, Function, and Structure of Photosynthetic Thylakoid Membranes of Asplenium australasicum (Sm.) Hook

The effect of light quality on the composition, function and structure of the thylakoid membranes, as well as on the photosynthetic rates of intact fronds from Asplenium australasicum, a shade plant, grown in blue, white, or red light of equal intensity (50 microeinsteins per square meter per second) was investigated. When compared with those isolated from plants grown in white and blue light, thylakoids from plants grown in red light have higher chlorophyll a/chlorophyll b ratios and lower amounts of light-harvesting chlorophyll a/b-protein complexes than those grown in blue light. On a chlorophyll basis, there were higher levels of PSII reaction centers, cytochrome f and coupling factor activity in thylakoids from red light-grown ferns, but lower levels of PSI reaction centers and plastoquinone. The red light-grown ferns had a higher PSII/PSI reaction center ratio of 4.1 compared to 2.1 in blue light-grown ferns, and a larger apparent PSI unit size and a lower PSII unit size. The CO₂ assimilation rates in fronds from red light-grown ferns were lower on a unit area or fresh weight basis, but higher on a chlorophyll basis, reflecting the higher levels of electron carriers and electron transport in the thylakoids.

The structure of thylakoids isolated from plants grown under the three light treatments was similar, with no significant differences in the number of thylakoids per granal stack or the ratio of appressed membrane length/nonappressed membrane length. The large freeze-fracture particles had the same size in the red-, blue-, and white-grown ferns, but there were some differences in their density. Light quality is an important factor in the regulation of the composition and function of thylakoid membranes, but the effects depend upon the plant species.

     

Evidence of the supercomplex organization of photosystem II and light-harvesting complexes in <Emphasis Type="Italic">Nannochloropsis granulata</Emphasis>

Diverse light-harvesting complexes (LHCs) have been found in photosynthetic microalgae that originated from secondary endosymbiosis involving primary red algae. However, the associations between LHCs and photosystem I (PSI) and photosystem II (PSII) in these microalgae are not fully understood. Eustigmatophyta is a red algal lineage that appears to have a unique organization in its photosynthetic machinery, consisting of only chlorophyll a and carotenoids that are atypical compared with other closely related groups. In this study, the supramolecular organization of pigment–protein complexes in the eustigmatophyte alga, Nannochloropsis granulata was investigated using Clear Native (CN) PAGE coupled with two-dimensional (2D) SDS-PAGE. Our results showed two slowly migrating green bands that corresponded to PSII supercomplexes, which consisted of reaction centers and LHCs. These green bands were also characterized as PSII complexes by their low temperature fluorescence emission spectra. The protein subunits of the PSII–LHC resolved by 2D CN/SDS-PAGE were analyzed by mass spectrometry, and four different LHC proteins were identified. Phylogenetic analysis of the identified LHC protein sequences revealed that they belonged to four different Lhc groups; (1) stress-related Lhcx proteins, (2) fucoxanthin chlorophyll a/c-binding Lhcf proteins, (3) red-shifted Chromera light-harvesting proteins (Red-CLH), and (4) Lhcr proteins, which are commonly found in organisms possessing red algal plastids. This is the first report showing evidence of a pigment–protein supercomplex consisting of PSII and LHCs, and to identify PSII-associated LHC proteins in Nannochloropsis.     

Evidence for light-independent and steeply decreasing PSII efficiency along twig depth in four tree species

Recent reports have indicated a considerably inactivated PSII in twig cortices, in spite of the low light transmittance of overlying periderms. Corresponding information for more deeply located and less illuminated tissues like xylem rays and pith are lacking. In this investigation we aimed to characterize the efficiency of PSII and its light sensitivity along twig depth, in conjunction with the prevailing light quantity and quality. To that aim, optical methods (spectral reflectance and transmittance, chlorophyll fluorescence imaging, low temperature fluorescence spectra) and photoinhibitory treatments were applied in cut twig sections of four tree species, while corresponding leaves served as controls. Compared to leaves, twig tissues displayed lower chlorophyll (Chl) levels and dark-adapted PSII efficiency, with strong decreasing gradients towards the twig center. The low PSII efficiencies in the inner stem were not an artifact due to an actinic effect of measuring beam or to an enhanced contribution of PSI fluorescence. In fact, the PSII/PSI ratios in cortices were higher and those in the xylem rays similar to that of leaves. Inner twig tissues were quite resistant to photoinhibitory treatments, tolerating irradiation levels several-fold higher than those encountered in their microenvironment. Moreover, the extent of high light tolerance was similar in naturally exposed and shaded twig sides. The results indicate an increasing, inherent and light-independent inactivation of PSII along twig depth. The findings are discussed on the basis of a recently proposed model for photosynthetic electron flow in twigs, taking into account the specific atmospheric and light microenvironment as well as the possible metabolic needs of such bulky organs.     

Light-Triggered Action Potentials and Changes in Quantum Efficiency of Photosystem II in <Emphasis Type="Italic">Anthoceros</Emphasis> Cells

Capillary microelectrodes and pulse amplitude-modulated microfluorometry were used to study light-triggered changes in cell membrane potential, chlorophyll fluorescence, and photochemical yield of PSII in chloroplasts of a hornwort Anthoceros sp. The action potential was generated by illuminating the plant sample for a few seconds. It was accompanied by a reversible decrease in quantum efficiency of PSII and by nonphotochemical quenching of fluorescence that continued as long as 10 min after the light stimulus. The presence of ammonium ions (2 mM) enhanced the amplitude and prolonged the duration of dark changes of fluorescence parameters in accordance with the reported increase in duration and amplitude of the light-triggered action potential in the presence of NH ₄ ⁺ . A rapid retardation of PSII activity within the first seconds of illumination was also evident from absorbance changes at 810 nm reflecting the redox conversions of chlorophyll P700. The PSII-dependent stage of reduction in the induction curves of P700 absorbance was strongly suppressed, and the amplitudes of signals induced by white and far-red light (717 nm) differed insignificantly. It is concluded that a short-term irradiation triggers the generation of ΔpH at the thylakoid membranes, which is accompanied by inhibition of the plasma membrane H⁺ pump and by reversible inactivation of PSII due to increased thermal dissipation of chlorophyll excitations.     

Effects of growth irradiance levels on the ratio of reaction centers in two species of marine phytoplankton

Cells of two species of single-celled marine algae, the diatom Skeletonema costatum (Greve), Cleve, and the chlorophyte Dunaliella tertiolecta Butcher, were cultured in white light of high (500-600 microeinsteins per square meter per second) and low (30 microeinsteins per square meter per second) intensity. For both algal species, cells grown at low light levels contained more chlorophyll a and had a lower ratio of chlorophyll a to chlorophylls b or c than did cells grown at high light levels. When photosynthetic unit sizes were measured on the basis of either oxygen flash yields or P₇₀₀ photooxidation, different results were obtained with the different species. In the chlorophyte, the cellular content of photosystem I (PSI) and photosystem II (PSII) reaction centers increased in tandem as chlorophyll a content increased so that photosynthetic unit sizes changed only slightly and the ratio PSI:PSII reaction centers remained constant at about 1.1. In the diatom, as the chlorophyll content of the cells increased, the number of PSI reaction centers decreased and the number of PSII reaction centers increased so that the ratio of PSI:PSII reaction centers decreased from about unity to 0.44. In neither organism did photosynthetic capacity correlate with changes in cellular content of PSI or PSII reaction centers. The results are discussed in relationship to the physical and biological significance of the photosynthetic unit concept.     

Evidence for active cyclic electron flow in twig chlorenchyma in the presence of an extremely deficient linear electron transport activity

Fast and slow chlorophyll fluorescence induction curves at high and low actinic visible light, post-illumination changes in fluorescence yield and reflectance changes at 820 nm induced by far-red light were used to characterize the state of PSII and PSI and their electron transport capabilities in chlorophyllous twig cortices of Eleagnus angustifolius L., while corresponding leaves served as controls. Twigs displayed low dark-adapted PSII photochemical efficiencies and particularly low linear electron transport rates when illuminated. In addition, their PSII population was characterized by a high proportion of inactive, non-Q_B-reducing centers and an incomplete quenching of fluorescence during the slow induction phase. It is suggested that PSII in twigs is an inefficient electron donor to PSI and/or the reductive pentose phosphate cycle. Yet, in spite of this apparent PSII deficiency, pools of intermediate electron carriers and potential PSI activity were more than sufficient to support the observed linear electron transport rates. Moreover, the rate of PSI reduction upon far-red/dark transitions and the magnitude of fluorescence yield increase upon white light/dark transitions were compatible with an efficient electron flow to PSI from stromal donors in the absence of PSII activity. We conclude that corticular chlorenchyma may be actively engaged in cyclic at the expense of a linear electron flow and discuss the possible physiological significance of this finding in conjunction with the particular microenvironmental conditions encountered within twigs.     

Fluorescence changes accompanying short-term light adaptations in photosystem I and photosystem II of the cyanobacterium <Emphasis Type="Italic">Synechocystis</Emphasis> sp. PCC 6803 and phycobiliprotein-impaired mutants: State 1/State 2 transitions and carotenoid-induced quenching of phycobilisomes

The features of the two types of short-term light-adaptations of photosynthetic apparatus, State 1/State 2 transitions, and non-photochemical fluorescence quenching of phycobilisomes (PBS) by orange carotene-protein (OCP) were compared in the cyanobacterium Synechocystis sp. PCC 6803 wild type, CK pigment mutant lacking phycocyanin, and PAL mutant totally devoid of phycobiliproteins. The permanent presence of PBS-specific peaks in the in situ action spectra of photosystem I (PSI) and photosystem II (PSII), as well as in the 77 K fluorescence excitation spectra for chlorophyll emission at 690 nm (PSII) and 725 nm (PSI) showed that PBS are constitutive antenna complexes of both photosystems. The mutant strains compensated the lack of phycobiliproteins by higher PSII content and by intensification of photosynthetic linear electron transfer. The detectable changes of energy migration from PBS to the PSI and PSII in the Synechocystis wild type and the CK mutant in State 1 and State 2 according to the fluorescence excitation spectra measurements were not registered. The constant level of fluorescence emission of PSI during State 1/State 2 transitions and simultaneous increase of chlorophyll fluorescence emission of PSII in State 1 in Synechocystis PAL mutant allowed to propose that spillover is an unlikely mechanism of state transitions. Blue–green light absorbed by OCP diminished the rout of energy from PBS to PSI while energy migration from PBS to PSII was less influenced. Therefore, the main role of OCP-induced quenching of PBS is the limitation of PSI activity and cyclic electron transport under relatively high light conditions.     

Photosynthetic Responses of Leaves to Water Stress, Expressed by Photoacoustics and Related Methods : II. The Effect of Rapid Drought on the Electron Transport and the Relative Activities of the Two Photosystems

The effect of rapid dehydration of detached tobacco leaves (Nicotiana tabacum L.) on the photochemical apparatus of photosynthesis was studied in vivo by a combination of methods: photoacoustics, chlorophyll a fluorescence, and cytochrome f difference spectroscopy. It was shown that the inhibition of gross O₂ evolution was mainly caused by inactivation of PSII: (a) The saturation curve of cytochrome-f photooxidation by farred (>710 nanometers) light was resistant to the stress, leading to the conclusion that photosystem I (PSI) was largely unaffected by the stress. (b) The extent of the chlorophyll a variable fluorescence arising from photosystem II (PSII) decreased with the progression of the stress, but was largely unaffected when the leaf was preincubated with electron donors to PSII, such as hydroxylamine. It is concluded that the drought damage to PSII occurred on the photooxidative side. Despite the extensive inhibition of PSII and the relative preservation of PSI, the apparent PSII/PSI activity balance was somewhat larger in stressed leaves than in the control, as indicated by photoacoustic measurements of Emerson enhancement. These measurements were performed continuously under conditions which favor transitions to either state 1 or 2, showing that the transition to state 2 was considerably inhibited. Simultaneous measurements of chlorophyll fluorescence induction at 680 and 730 mm at room temperature were also used to probe changes in energy distribution between PSII and PSI and indicated that the transition from a dark adapted state to state 2 was also affected in water-stressed leaves. The saturation curve of the far-red light effect in Emerson enhancement was not changed by the stress, giving another independent evidence for the drought resistance of PSI activity. This apparent preservation of the imbalance in photochemical activities in favor of PSII, despite the fact that PSII is strongly inhibited, and PSI is not, supports a previous suggestion that the electron transfer between the two photosystems is not random but that a large extent of PSII and PSI units are specifically linked.