Effects of UV-B radiation on photosynthesis and growth of terrestrial plants

The photosynthetic apparatus of some plant species appears to be well-protected from direct damage from UV-B radiation. Leaf optical properties of these species apparently minimizes exposure of sensitive targets to UV-B radiation. However, damage by UV-B radiation to Photosystem II and Rubisco has also been reported. Secondary effects of this damage may include reductions in photosynthetic capacity, RuBP regeneration and quantum yield. Furthermore, UV-B radiation may decrease the penetration of PAR, reduce photosynthetic and accessory pigments, impair stomatal function and alter canopy morphology, and thus indirectly retard photosynthetic carbon assimilation. Subsequently, UV-B radiation may limit productivity in many plant species. In addition to variability in sensitivity to UV-B radiation, the effects of UV-B radiation are further confounded by other environmental factors such as CO₂, temperature, light and water or nutrient availability. Therefore, we need a better understanding of the mechanisms of tolerance to UV-B radiation and of the interaction between UV-B and other environmental factors in order to adequately assess the probable consequences of a change in solar radiation.Abbreviations A_max light and CO₂ saturated rate of oxygen evolution - Ci internal CO₂ concentration - F_v/F_m ratio of variable to total fluorescence yield - PAR photosynthetically active radiation (400–700 nm) - PS II Photosystem II - _app apparent quantum yield of photosynthesis - SLW specific leaf weight - UV-B ultraviolet-B radiation between 290–320 nm     

On the relationship between the quantum yield of Photosystem II electron transport,as determined by chlorophyll fluorescence and the quantum yield of CO2-dependent O2 evolution

We tested the two empirical models of the relationship between chlorophyll fluorescence and photosynthesis, previously published by Weis E and Berry JA 1987 (Biochim Biophys Acta 894: 198–208) and Genty B et al. 1989 (Biochim Biophys Acta 990: 87–92). These were applied to data from different species representing different states of light acclimation, to species with C₃ or C₄ photosynthesis, and to wild-type and a chlorophyll b-less chlorina mutant of barley. Photosynthesis measured as CO₂-saturated O₂ evolution and modulated fluorescence were simultaneously monitored over a range of photon flux densities. The quantum yields of O₂ evolution (ØO₂) were based on absorbed photons, and the fluorescence parameters for photochemical (q_p) and non-photochemical (q_N) quenching, as well as the ratio of variable fluorescence to maximum fluorescence during steady-state illumination (F'_v/F'_m), were determined. In accordance with the Weis and Berry model, most plants studied exhibited an approximately linear relationship between ØO₂/q_p (i.e., the yield of O₂ evolution by open Photosystem II reaction centres) and q_N, except for wild-type barley that showed a non-linear relationship. In contrast to the linear relationship reported by Genty et al. for q_p×F'_v/F'_m (i.e., the quantum yield of Photosystem II electron transport) and ØCO₂, we found a non-linear relationship between q_p×F'_v/F'_m and ØO₂ for all plants, except for the chlorina mutant of barley, which showed a largely linear relationship. The curvilinearity of wild-type barley deviated somewhat from that of other species tested. The non-linear part of the relationship was confined to low, limiting photon flux densities, whereas at higher light levels the relationship was linear. Photoinhibition did not change the overall shape of the relationship between q_p×F'_v/F'_m and ØO₂ except that the maximum values of the quantum yields of Photosystem II electron transport and photosynthetic O₂ evolution decreased in proportion to the degree of photoinhibition. This implies that the quantum yield of Photosystem II electron transport under high light conditions may be similar for photoinhibited and non-inhibited plants. Based on our experimental results and theoretical analyses of photochemical and non-photochemical fluoresce quenching processes, we conclude that both models, although not universal for all plants, provide useful means for the prediction of photosynthesis from fluorescence parameters. However, we also discuss that conditions which alter one or more of the rate constants that determine the various fluorescence parameters, as well as differential light penetration in assays for oxygen evolution and fluorescence emission, may have direct effect on the relationships of the two models.Abbreviations F₀ and F'₀ fluorescence when all Photosystem II reaction centres are open in dark- and light-acclimated leaves, respectively - F_m and F'_m fluorescence when all Photosystem II reaction centres are closed in dark and light, respectively - F_v variable fluorescence equal to F_m-F₀ - F_s steady state level of fluorescence in light - F'_v and F'_m variable (F'_m-F'₀) and maximum fluorescence under steady state light conditions - HEPES N-2-hydroxyethylpiperazine-N-2-ethane-sulphonic acid - Q_A the primary, stabile quinone acceptor of Photosystem II - q_N non-photochemical quenching of fluorescence - q_p photochemical quenching of fluorescence - ØO₂ quantum yield of CO₂-saturated O₂ evolution based on absorbed photons     

Peroxidation of lipids and growth inhibition induced by UV-B irradiation

Cotyledons excised from dark-grown seedlings of cucumber (Cucumis sativus L.) were cultured in vitro under UV radiation at different wavelengths, obtained by passage of light through cut-off filters with different transmittance properties. Growth and the synthesis of chlorophyll (Chl) in cotyledons were inhibited and malondialdehyde was accumulated upon irradiation at wavelengths below 320 nm. Exogenous application of scavengers of free radicals reversed the growth inhibition induced by UV-B. Measurement of the fluorescence of Chl a suggested that electron transfer in photosystems was affected by UV-B irradiation. On the basis of these results, the involvement is postulated of active species of oxygen in damages to thylakoid membranes and the growth inhibition that are induced by UV-B irradiation.Abbreviations Chl chlorophyll - F_m maximal fluorescence (dark) - F_m maximal fluorescence (light) - F_v variable fluorescence (dark) - F_v variable fluorescence (light) - MDA malondialdehyde - O₂ Superoxide radical - PS photosystem - q_N non-photochemical quenching of fluorescence - q_P photochemical quenching of fluorescence - UV-B_BE biologically effective UV-B radiation - WL(T = 0.5) wavelength at which 50% transmittance occurs     

Effects of dehydration on the electron transport of Chlorella. An in vivo fluorescence study

Chlorella was used to study the effects of dehydration on photosynthetic activities. The use of unicellular green algae assured that the extent of dehydration was uniform throughout the whole cell population during the course of desiccation. Changes in the activities of the cells were monitored by measurements of fluorescence induction kinetics. It was found that inhibition of most of the photosynthetic activities started at a similar level of cellular water content. They included CO₂ fixation, photochemical activity of Photosystem II and electron transport through Photosystem I. The blockage of electron flow through Photosystem I was complete and the whole transition occurred within a relative short time of dehydration. On the other hand, the suppression of Photosystem II activity was incomplete and the transition took a longer time of dehydration. Upon rehydration, the inhibition of Photosystem II activity was fully reversible when samples were in the middle of the transition, but was not thereafter. The electron transport through Photosystem I was also reversible during the transition, but was only partially afterward.Abbreviations DCMU 3-(3,4-dichlorophenyl)-1,1-dimethyl urea - F_m maximum fluorescence yield - F₀ non-variable fluorescence level emitted when all PS II centers are open - F_v variable part of fluorescence - PS photosystem - Q_A primary quinone acceptor of Photosystem II     

Spectroscopy of non-photochemical and photochemical quenching of chlorophyll fluorescence in leaves; evidence for a role of the light harvesting complex of Photosystem II in the regulation of energy dissipation

Dissipation of absorbed excitation energy as heat, measured by its effect on the quenching of chlorophyll fluorescence, is induced under conditions of excess light in order to protect the photosynthetic apparatus of plants from light-dependent damage. The spectral characteristics of this quenching have been compared to that due to photochemistry in the Photosystem II reaction centre using leaves of Guzmania monostachia. This was achieved by making measurements at 77K when fluorescence emission bands from each type of chlorophyll protein complex can be distinguished. It was demonstrated that photochemistry and non-photochemical dissipation preferentially quench different emission bands and therefore occur by dissimilar mechanisms at separate sites. It was found that photochemistry was associated with a preferential quenching of emission at 688 nm whereas the spectrum for rapidly reversible non-photochemical quenching had maxima at 683 nm and 698 nm, suggesting selective quenching of the bands originating from the light harvesting complexes of Photosystem II. Further evidence that this was occurring in the light harvesting system was obtained from the fluorescence excitation spectra recorded in the quenched and relaxed states.Abbreviations pH transthylakoid pH gradient - F_o minimum level of chlorophyll fluorescence when Photosystem II reaction centres are open - F_m maximum level of fluorescence when Photosystem II reaction centres are closed - F_v variable fluorescence F_m minus F_o - F'_o F_o in any quenched state - F_m F_m in any quenched state - LHCII light harvesting complexes of Photosystem II - PSI Photosystem I - PS II Photosystem II - qN non-photochemical quenching of chlorophyll fluorescence - qE non-photochemical quenching of chlorophyll fluorescence that occurs in the presence of a pH     

Photosystem II reaction centres stay intact during low temperature photoinhibition

Photoinhibition of photosynthesis was studied in intact barley leaves at 5 and 20°C, to reveal if Photosystem II becomes predisposed to photoinhibition at low temperature by 1) creation of excessive excitation of Photosystem II or, 2) inhibition of the repair process of Photosystem II. The light and temperature dependence of the reduction state of Q_A was measured by modulated fluorescence. Photon flux densities giving 60% of Q_A in a reduced state at steady-state photosynthesis (300 mol m^–2s^–1 at 5°C and 1200 mol m^–2s^–1 at 20°C) resulted in a depression of the photochemical efficiency of Photosystem II (F_v/F_m) at both 5 and 20°C. Inhibition of F_v/F_m occurred with initially similar kinetics at the two temperatures. After 6h, F_v/F_m was inhibited by 30% and had reached steady-state at 20°C. However, at 5°C, F_v/F_m continued to decrease and after 10h, F_v/F_m was depressed to 55% of control. The light response of the reduction state of Q_A did not change during photoinhibition at 20°C, whereas after photoinhibition at 5°C, the proportion of closed reaction centres at a given photon flux density was 10–20% lower than before photoinhibition.Changes in the D1-content were measured by immunoblotting and by the atrazine binding capacity during photoinhibition at high and low temperatures, with and without the addition of chloramphenicol to block chloroplast encoded protein synthesis. At 20°C, there was a close correlation between the amount of D1-protein and the photochemical efficiency of photosystem II, both in the presence or in the absence of an active repair cycle. At 5°C, an accumulation of inactive reaction centres occurred, since the photochemical efficiency of Photosystem II was much more depressed than the loss of D1-protein. Furthermore, at 5°C the repair cycle was largely inhibited as concluded from the finding that blockage of chloroplast encoded protein synthesis did not enhance the susceptibility to photoinhibition at 5°C.It is concluded that, the kinetics of the initial decrease of F_v/F_m was determined by the reduction state of the primary electron acceptor Q_A, at both temperatures. However, the further suppression of F_v/F_m at 5°C after several hours of photoinhibition implies that the inhibited repair cycle started to have an effect in determining the photochemical efficiency of Photosystem II.Abbreviations CAP D-threochloramphenicol - F₀ and F ₀ fluorescence when all Photosystem II reaction centres are open in dark- and light-acclimated leaves, respectively - F_m and F _m fluorescence when all Photosystem II reaction centres are closed in dark- and light-acclimated leaves, respectively - F_s fluorescence at steady state - Q_A the primary, stable quinone acceptor of Photosystem II - q_N non-photochemical quenching of fluorescence - q_P photochemical quenching of fluorescence     

Enhanced UV-B radiation alleviates the adverse effects of summer drought in two Mediterranean pines under field conditions

The effects of enhanced UV-B (290-320 nm) radiation on two native Mediterranean pines (Pinus pinea L., Pinus halepensis Mill.) were recorded during a one-year field study. Plants received ambient or ambient plus supplemental UV-B radiation (simulating a 15% stratospheric ozone depletion over Patras. Greece, 38.3°N. 29.1°E) and only natural precipitation, i.e. they were simultaneously exposed to other natural stresses. particularly water stress during summer. Supplemental UV-B irradiation started in early February, 1993 and up to late June, no effects were observed on growth and photochemical efficiency of photosystem II, as measured by chlorophy II fluorescence induction. Water stress during the summer was manifested in the control plants as a decline in the ratio of variable to maximum fluorescence (F_v/F_m), the apparent photon yield for oxygen evolution (φ_l) and the photosynthetic capacity at 5% CO₂ (P_m). In addition, a partial needle loss was evident. Under supplemental UV-B radiation, however, the decreases in F_v/F_m, φ_i, and P_m. as well as needle losses were significantly less. Soon after the first heavy autumn rains. photosynthetic parameters in both control and UV-B treated plants recovered to similar values. but the transient summer superiority of UV-B irradiated plants resulted in a significant increase in their dry weight measured at plant harvest. during late January. 1994. Plant height. UV-B absorbing compounds, photosynthetic pigments and relative water content measured at late spring. late summer and at plant harvest, were not significantly affected by supplemental UV-B radiation. The results indicate that enhanced UV-B radiation may be beneficial for Mediterranean pines through a partial alleviation of the adverse effects of summer drought.     

Inhibition of Photosystem II by UV-B Radiation and the Conditions for Recovery in the Liverwort Conocephalum conicum Dum.

Inhibition of photosynthesis by UV-B was investigated in the thalloid liverwort Conocephalum conicum Dum. UV-B irradiance was adjusted to a strength producing 50% inhibition of the rate of photosynthesis during 10 min of irradiation. A linear relationship of the fluorescence terms F_v/F_m of photosystem (PS) II and J_P was observed following a UV-B irradiation. This suggested that PS II was a major site of UV-B-induced damage of photosynthesis. The apparent inhibition of F_v/F_m was much smaller when electron flow to the secondary PS II acceptor Q_B was inhibited by DCMU or when F_v/F_m was measured at 77 K. Apparently, the major target of UV-B effects was electron donation to the PS II reaction center, rather than electron transfer reactions at the PS II acceptor side. The time required for repair of PS II from UV-B-induced damage was light-dependent and minimal at a flux density of 5 μE m^?2 s^?1. Low temperatures and the presence of streptomycin inhibited the repair processes of PS II, indicating that protein synthesis may be involved in the recovery of PS II. The data indicate that UV-B irradiation on bright and cool winter days may be most harmful for photosynthesis of C. conicum. A repeated irradiation of the thalli with UV-B induced tolerance of photosynthesis which was related to an accumulation of pigments with a maximum of absorption around 315 nm.     

Down regulation of photosynthesis in Artabotrys hexapetatus by high light

Using variable to maximum fluorescence (F_v/F_m) as the criterion, the down regulation of photosynthesis by high light stress was characterized in the detached leaves of Artabotrys hexapetatus. The decrease in F_v/F_m was corelated with the decrease in oxygen evolution by thylakoids isolated from high light exposed leaves. The decrease in F_v/F_m was linear with increasing time of exposure to high light. A comparison of recovery measured as F_v/F_m, in low light versus dark, revealed that the recovery in darkness was as significant as in low light. Since the relaxation of fluorescence was a rapid response after exposure to high light and the fact that the recovery occurs in total darkness, it is concluded that photoinhibition and down regulation of photosynthesis by high light are independent events.Abbreviation Fpl- initial plateau - F_m- maximum fluorescence - F_o- prompt fluorescence - F_v- variable fluorescence - PFD- photon flux density - PS I (II)- Photosystem I (II)     

Evidence that UV-B tolerance of the photosynthetic apparatus in microalgae is related to the D1-turnover mediated repair cycle in vivo

The present study was conceived to elucidate the potential importance of the D1 turnover-mediated repair mechanism in UV-B tolerance of the photosynthetic apparatus in microalgae. To this end, the lab-identified UV-B sensitive and tolerant species of Chlorophyte and Chromophyte algae was used to examine photosynthetic response to UV-B exposure in the presence vs. the absence of streptomycin, an inhibitor of chloroplast protein synthesis. Measurements of photosynthetic O₂ evolution capacity and chlorophyll fluorescence parameters (F_v/F_m, Φ_PSII) illustrated species-specific UV-B sensitivity of the photosynthetic apparatus. Addition of the inhibitor streptomycin caused significant enhancements of UV-B-caused depression of photosynthesis in UV-B tolerant species, while little effect was observed in the sensitive species. In the tolerant species, recovery from UV-B induced 20 percnt; decline in F_v/F_m reached completion within 2 hours, much faster than that in the sensitive species. Immunoblotting revealed that exposure to UV-B radiation caused substantial degradation of the D1 protein in the sensitive Heterococcus brevicellularis, which was little enhanced by addition of the inhibitor. The same UV-B exposure lead to less D1 degradation in the tolerant Scenedesmus sp., which was significantly enhanced by addition of the inhibitor. This study shows that UV-B tolerance of the photosynthetic apparatus in microalgae was associated with a strong capacity for recovery from the UV-B-induced damage and this capacity related to the D1 turnover-mediated repair cycle, and largely determined UV-B tolerance of the photosynthetic apparatus in these organisms.     

Influence of ultraviolet-B (UV-B) radiation on photosynthetic and growth characteristics in field-grown cassava (Manihot esculentum Crantz)

The effects of ultraviolet-B (UV-B between 290 and 320 nm) on photosynthesis and growth characteristics were investigated in field grown cassava (Manihot esculentum Crantz). Plants were grown at ambient and ambient plus a 5.5kJ m^?2 d^?1 supplementation of UV-B radiation for 95 d. The supplemental UV-B fluence used in this experiment simulated a 15% depletion in stratospheric ozone at the equator (0°N). Carbon dioxide exchange, oxygen evolution, and the ratio of variable to maximum fluorescence (F_v/F_m) were determined for fully expanded leaves after 64–76 d of UV-B exposure. AH plants were harvested after 95 d of UV-B exposure, assayed for chlorophyll and UV-B absorbing compounds, and separated into leaves, petioles, stems and roots. Exposure to UV-B radiation had no effect on in situ rates of photosynthesis or dark respiration. No difference in the concentration of UV-B absorbing compounds was observed between treatments. A 2-d daytime diurnal comparison of F_v to F_m ratios indicated a significant decline in F_v/F_m ratios and a subsequent increase in photoinhibition under enhanced UV-B radiation if temperature or PPF exceeded 35°C or 1800μmol m^?2 s^?1, respectively. However, UV-B effects on fluorescence kinetics appeared to be temporal since maximal photosynthetic rates as determined by oxygen evolution at saturated CO₂ and PPF remained unchanged. Although total biomass was unaltered with UV-B exposure, alterations in the growth characteristics of cassava grown with supplemental UV-B radiation are consistent with auxin destruction and reduced apical dominance. Changes in growth included an alteration of biomass partitioning with a significant increase in shoot/root ratio noted for plants receiving supplemental UV-B radiation. The increase in shoot/root ratio was due primarily to a significant decrease in root weight (–32%) with UV-B exposure. Because root production determines the harvest-able portion of cassava, UV-B radiation may still influence the yield of an important tropical agronomic species, even though photosynthesis and total dry biomass may not be directly affected.     

Responses of Strawberry (Fragaria × ananassa) to Supplemental UV-B Radiation and Selenium Under Field Conditions

In greenhouse experiments, selenium (Se) has been shown to defend plants against detrimental effects of heavy UV-B radiation stress. The aim of this study was to investigate whether this positive effect can be found in open-field conditions with enhancement of UV-B radiation. In the experiment, conducted with strawberry (Fragaria×ananassa, cultivars “Jonsok” and “Polka”) over two growing seasons, plants were exposed to UV-B radiation (including UV-A) and cultivated without Se or supplied with Se added at two levels (0.1 and 1.0 mg kg⁻¹). The plants were monitored for growth, flavonoids, chlorophyll fluorescence, net photosynthesis as well as tissue and cell structure. Photosystem II was observed to be sensitive to UV-B stress under field conditions. In the leaves, a decrease in F_v/F_m was seen at the end of the growing season, implying a cumulative effect of UV-B stress. Several parameters, especially cell and tissue structures, were affected by UV-B and UV-A treatments, which proves the need for UV-A control in outdoor UV-B supplementation studies. Addition of Se did not ameliorate the harmful effects of UV-B but the lower Se-increment level increased leaf growth. The effects of UV-B and Se differed during the two experimental years, indicating the need to repeat experiments during several growing seasons.     

Age-related differences in frost sensitivity of the photosynthetic apparatus of two Plagiomnium species

Shoots of two species of moss, Plagiomnium undulatum (Hedw.) Kop. and Plagiomnium affine (Funck) Kop., were subjected to freezing at various temperatures. After thawing, the activities of different photosynthetic reactions were determined in relation to the ages of the leaves. Analysis of the fast kinetics of chlorophyll-a fluorescence of individual leaves showed that young and old tissues were considerably less frost tolerant than mature ones. In principle, the pattern of freeze inactivation of photosynthetic reactions resembles that observed in higher plants. The decreases in the amplitude of F_v (variable fluorescence) and the ratio of F_v to F_m (maximum fluorescence) with increasing freezing stress reflect a progressive inactivation of photosystem II (PSII)-mediated electron transport, i.e. inhibition of photoreaction to photochemistry and-or electron donation to the photochemical reaction, and thus a decline in the potential photochemical efficiency of PSII. The insignificant change in the F₀ (constant fluorescence) level during progressive decline of F_v indicates that the excitation-energy transfer between antenna pigments and from those to reaction centres of PSII was little impaired by lethal freezing stress. Sugar analyses of various stem sections showed that ontogenetic variation in the frost tolerance of leaves cannot be attributed to differences in the cellular levels of sucrose, glucose and fructose.Abbreviations and Symbols DCMU 3-(3,4-dichlorophenyl)-1,1-dimethylurea - F_m maximum fluorescence - F₀ constant (initial) fluorescence - F_v variable fluorescence     

Theoretical assessment of alternative mechanisms for non-photochemical quenching of PS II fluorescence in barley leaves

The components of non-photochemical chlorophyll fluorescence quenching (qN) in barley leaves have been quantified by a combination of relaxation kinetics analysis and 77 K fluorescence measurements (Walters RG and Horton P 1991). Analysis of the behaviour of chlorophyll fluorescence parameters and oxygen evolution at low light (when only state transitions — measured as qN_t — are present) and at high light (when only photoinhibition — measured as qN_i — is increasing) showed that the parameter qN_t represents quenching processes located in the antenna and that qN_i measures quenching processes located in the reaction centre but which operate significantly only when those centres are closed. The theoretical predictions of a variety of models describing possible mechanisms for high-energy-state quenching, measured as the residual quenching, qN_e, were then tested against the experimental data for both fluorescence quenching and quantum yield of oxygen evolution. Only one model was found to agree with these data, one in which antennae exist in two states, efficient in either energy transfer or energy dissipation, and in which those photosynthetic units in a dissipative state are unable to exchange energy with non-dissipative units.Abbreviations: F_o, F_m room-temperature chlorophyll fluorescence yield with all centres open, closed - F_v variable fluorescence yield - LHC II light-harvesting chlorophyll-protein complex of PS II - PS I, PS II Photosystem I, II - P700, P680 primary donor in Photosystem I, II - Q_A primary electron acceptor of PS II - P_max maximum quantum yield of oxygen evolution - qN coefficient of non-photochemical quenching of variable fluorescence - qN_e, qN_t, qN_i coefficient of non-photochemical quenching due to high-energy-state, state transition, photoinhibition - qO coefficient of quenching of dark level fluorescence - qP coefficient of photochemical quenching of variable fluorescence - _P intrinsic quantum yield of open PS II reaction centres = _s/qP - _{PS 2} quantum yield of PS = qP × F_v/F_m - _S quantum yield of oxygen evolution = rate of oxygen evolution/light intensity     

Photosystem II function and herbicide binding sites during photoinhibition of spinach chloroplasts in-vivo and in-vitro

The time courses of some Photosystem II (PS II) parameters have been monitored during in-vivo and in-vitro photoinhibition of spinach chloroplasts, at room temperature and at 10 °C or 0 °C. Exposing leaf discs of low-light grown spinach at 25 °C to high light led to photoinhibition of chloroplasts in-vivo as manifested by a parallel decrease in the number of functional PS II centres, the variable chlorophyll fluorescence at 77K (F _v /F _m ), and the number of atrazine-binding sites. When the photoinhibitory treatment was given at 10 °C, the former two parameters declined in parallel but the loss of atrazine-binding sites occurred more slowly and to a lesser extent. During in-vitro photoinhibition of chloroplast thylakoids at 25 °C, the loss of functional PS II centres proceeded slightly more rapidly than the loss of atrazine-binding sites, and this difference in rate was further increased when the thylakoids were photoinhibited at 0 °C. During the recovery phase of leaf discs (up to 9 h) the increases in F _v /F _m preceded that of the number of functional PS II centres, while only a further decline in the number of atrazine-binding sites was observed. The recovery of variable chlorophyll fluorescence and the concentration of functional PS II centres occurred more rapidly at 25 °C than at 10 °C. These results suggest that the photoinhibition of PS II function is a relatively temperature-independent early photochemical event, whereas the changes in the concentration of herbicide-binding sites appear to be a more complex biochemical process which can occur with a delayed time course.Abbreviations BSA bovine serum albumin - Chl chlorophyll - D1 32kDa herbicide-binding polypeptide in photosystem II and product of the psbA gene - D2 34kDa polypeptide in photosystem II which is the product of the psbD gene - DCMU 3-(3,4-dichlorophenyl)-1,1-dimethylurea - DCPIP 2,6-dichlorophenolin-dophenol - F ₀, F _v , F _m chlorophyll fluorescence with reaction centres open, variable and maximum fluorescence, respectively - LDS lithium dodecyl sulfate - MES 2-(N-morpholino) ethanesulfonic acid - PSII photosystem II - Q_A, Q_B first and second quinone-type PS II acceptor, respectively     

Responses of Nine Bryophyte and One Lichen Species from Different Microhabitats to Elevated UV-B Radiation

Chlorophyll fluorescence parameters (F_v/F_m, R_Fd) of nine bryophyte and one lichen species were investigated after prolonged exposure to elevated UV-B radiation. The majority of the investigated bryophytes showed a prompt or inducible tolerance to increase UV-B irradiation. Among the investigated species high degree of UV-tolerance coincides with strong desiccation tolerance.     

UV-B response of green and etiolated barley seedlings

7-d-old etiolated and green barley seedlings (Hordeum vulgare L. cv. Alfa) were irradiated with UV-B for 30 min and then kept for 24 h in light or darkness. Chlorophyll (Chl) synthesis was inhibited by about 30 % as a result of UV-B irradiation, but there were no significant changes in photochemical activity measured by variable to maximum fluorescence ratio (F_v/F_m), quantum yield (Φ_PS2) and oxygen evolution rate. Electron transport of etiolated seedlings was similar to that of green ones, nevertheless, the Chl content was more then 2-fold lower. Ribulose-1,5-bisphosphate carboxylase/oxygenase large and small subunits were diminished as a result of UV-B irradiation in etiolated and green plants, especially in those kept in the darkness. Catalase activity decreased and total superoxide dismutase activity increased in green and etiolated plants following UV-B treatment. When benzidine was used as a substrate, an isoform located between guaiacol peroxidases 2 and 3 (guaiacol peroxidase X) appeared, which was specific for UV-B treatment. As a result of irradiation, the contents of UV-B absorbing and UV-B induced compounds increased in green seedlings but not in etiolated seedlings.     

Changes in D1-protein turnover and recovery of photosystem II activity precede accumulation of chlorophyll in plants after release from mineral stress

After seven weeks of a combined magnesium and sulphur deficiency, spinach (Spinacea oleracea L.) plants showed a substantial accumulation of inactivated photosystem II (PSII) centres as indicated by a 40% decrease of the chlorophyll (Chl) fluorescence parameter F_v/F_m (F_v being the yield of variable fluorescence and F_m the yield of maximal fluorescence when all reaction centres are closed) together with a severe loss of leaf Chl content of 75%. The responses of the photosynthetic apparatus were examined when the deficient plants were transferred back to a rich nutrient medium. During the first 24 h of the recovery phase, thylakoid protein synthesis measured as incorporation of [¹⁴C]leucine per unit of Chl increased substantially. The synthesis rate of the D1 reaction-centre polypeptide of PSII, which in the deficient plants was reduced to 50% of the non-deficient control, was stimulated eight- to ninefold. D1-protein content, which in the deficient plants was reduced to 40% of the non-deficient control, started to increase 2 d later. Thus, D1-protein degradation was also enhanced. The increased D1-protein turnover led to a rapid repair of the existing PSII centres as indicated by the rise of F_v/F_m. It was completed at day 7 of the recovery phase. At day 2 of the recovery phase, the synthesis of other thylakoid proteins such as the D2 protein, cytochrome b ₅₅₉, CP 47 and the 33-kDa polypeptide of the water-splitting system, became stimulated. This process resulted in an accumulation of new PSII centres. During the first week, formation of new PSII centres was not associated with an increase in leaf Chl content. The Chl content of the recovering leaves only started to increase when the ratio of PSII polypeptides versus LHCII (light-harvesting complex of PSII), which was substantially diminished in the deficient plants, became comparable to that of the control. The recovery process was accompanied by substantial changes in thylakoid protein phosphorylation. Their relevance to thylakoid protein turnover and stability is discussed.Abbreviations Chl chlorophyll - cyt cytochrome - F_o yield of intrinsic fluorescence when all PSII centres are open in the dark - F_m yield of maximal fluorescence when all reaction centres are closed - F_m fluorescence yield when all reaction centres are closed (after a saturating flash) under steady-state conditions - F_v yield of variable fluorescence, (difference between F_oand F_m) - F yield of variable fluorescence under steady state conditions - LHC light-harvesting complex - PQ plastoquinone - Q_A primary quinone acceptor of PSII - Q_B secondary quinone acceptor of PSII - q_P photochemical quenching - q_n non-photochemical quenching The authors like to thank Dipl. Biol. Britta Untereiser for determining the chlorophyll fluorescence quenching factors. This work was supported by grants from the Bundesminister für Forschung und Technologie, the Project Europäisches Forschungszentrum and the German Israeli Foundation in cooperation with Prof. I. Ohad, Hebrew University, Jerusalem, Israel.     

A simple model relating photoinhibitory fluorescence quenching in chloroplasts to a population of altered Photosystem II reaction centers

A model is presented describing the relationship between chlorophyll fluorescence quenching and photoinhibition of Photosystem (PS) II-dependent electron transport in chloroplasts. The model is based on the hypothesis that excess light creates a population of inhibited PS II units in the thylakoids. Those units are supposed to posses photochemically inactive reaction centers which convert excitation energy to heat and thereby quench variable fluorescence. If predominant photoinhibition of PS II and cooperativity in energy transfer between inhibited and active units are presumed, a quasi-linear correlation between PS II activity and the ratio of variable to maximum fluorescence, F_VF_M, is obtained. However, the simulation does not result in an inherent linearity of the relationship between quantum yield of PS II and F_VF_M ratio. The model is used to fit experimental data on photoinhibited isolated chloroplasts. Results are discussed in view of current hypotheses of photoinhibition.Abbreviations F_M maximum total fluorescence - F₀ initial fluorescence - F_V maximum variable fluorescence - PS Photosystem - Q_A, Q_B primary and secondary electron acceptors of Photosystem II     

The efficiency of electron transfer from QA − to the donor side of Photosystem II decreases during induction of photosynthesis: Evidences from chlorophyll fluorescence and photoacoustic techniques

The amplitudes ratio of the fast and slow phases (A_fast/A_slow) in the kinetics of the dark relaxation of variable chlorophyll fluorescence (F_V) was studied after various periods of illumination of dark-adapted primary barley leaves. Simultaneously, photosynthetic activity was monitored using the photoacoustic technique and the photochemical and non-photochemical fluorescence quenching parameters. The ratio A_fast/A_slow changed with the preceding illumination time in a two-step manner. During the first stage of photosynthetic induction (0–20 s of illumination), characterized by a drop in O₂-dependent photoacoustic signal following an initial spike and by a relatively stable small value of photochemical F_V quenching, the ratio A_fast/A_slow remained practically unaltered. During the second stage (20–60 s of illumination), when both the rate of O₂ evolution and the photochemical F_V quenching were found to be sharply developed, a marked increase in the above ratio was also observed. A linear correlation was found between the value of the photochemical quenching and the ratio A_fast/A_slow during the second phase of photosynthetic induction. It is concluded that the slow phase appearing in the kinetics of F_V dark relaxation is not due to the existence of Photosystem II reaction centres lacking the ability to reduce P700⁺ with high rates, but is instead related to the limitation of electron release from Photosystem I during the initial stage of the induction period of photosynthesis. This limitation keeps the intersystem electron carriers in the reduced state and thus increases the probability of back electron transfer from Q_A ^– to the donor side of Photosystem II.Abbreviations A_fast/A_slow the ratio of magnitudes between the fast and slow phases of dark relaxation of variable fluorescence - F_O initial level of chlorophyll fluorescence - F_V variable chlorophyll fluorescence (F-F_O) - (F_V)_S the yield of variable chlorophyll fluorescence under saturating pulse in illuminated leaves - (F_V)_M the yield of variable chlorophyll fluorescence under saturating pulse in dark-adapted leaves - PA photoacoustic - PSI Photosystem I - PS II Photosystem II - qN non-photochemical quenching - qQ photochemical quenching