Comparative study of the action of ultraviolet-C and ultraviolet- B radiation on photosynthetic electron transport

The effect of ultraviolet-C (UV-C, mainly 254 nm radiation) and ultraviolet-B (UV-B, 290-320 nm) radiation on the photosynthetic electron transport reactions has been investigated. The rates of Hill activity mediated by ferricyanide and dichlorodimethoxy-p-benzoquinone (DCDMQ) were differently sensitive to UV-C but equally inhibited by UV-B. Replacement of water with diphenylcarbazide was ineffective in restoring the activity of dichlorophenol indophenol (DCPIP) Hill reaction in UV-B treated chloroplasts, but had significant effect in UV-C treated chloroplasts.
Photobleaching of carotenoids in the presence of carbonyl cyanide-m-chlorophenyl-hydrazone, an indicator of the photochemical reaction associated with the reaction centre of photosystem II, was suppressed and is paralleled by the changes in Hill activity only in UV-B-treated chloroplasts. Carotenoid photobleaching occurred even in UV-C treated chloroplasts showing no measurable Hill activity. UV-C and UV-B irradiation diminished variable fluorescence. With UV-B treated, but not with UV-C treated chloroplasts, an increase in the fluorescence yield was observed upon the addition of 3-(3,4-dichIorophenyl)-l,l-dimethylurea (DCMU) and/or Na dithionite.
Photosystem I activity was found to be unaffected by both UV-C and UV-B radiation at the fluences tested. Kinetics of P700 photooxidation and dark reversal in UV treated chloroplasts indicate that only the electron flow from photosystem II to photosystem I is impaired. It is concluded that while UV-B radiation inactivates specifically the photosystem II reaction centre, UV-C radiation acts at plastoquinone, the quencher Q, and the water oxidizing enzyme system.     

Net Photosynthesis, Electron Transport Capacity, and Ultrastructure of Pisum sativum L. Exposed to Ultraviolet-B Radiation

Pisum sativum L. was exposed to ultraviolet-B (UV-B) radiation (280-315 nm) in greenhouse and controlled environment chambers to examine the effect of this radiation on photosynthetic processes. Net photosynthetic rates of intact leaves were reduced by UV-B irradiation. Stable leaf diffusion resistances indicated that the impairment of photosynthesis did not involve the simple limitation of CO₂ diffusion into the leaf. Dark respiration rates were increased by previous exposure to this radiation. Electron transport capacity as indicated by methylviologen reduction was also sensitive to UV-B irradiation. The ability of ascorbate-reduced 2,6-dichlorophenolindophenol to restore much of the electron transport capacity of the UV-B-irradiated plant material suggested that inhibition by this radiation was more closely associated with photosystem II than with photosystem I. Electron micrographs indicated structural damage to chloroplasts as well as other organelles. Plant tissue irradiated for only 15 minutes exhibited dilation of thylakoid membranes of the chloroplast in some cells. Some reduction in Hill reaction activity was also evidenced in these plant materials which had been irradiated for periods as short as 15 minutes.     

On the possible site of inhibition of photosynthetic electron transport by ultraviolet-B (UV-B) radiation

In Amaranthus chloroplasts that are exposed to ultraviolet-B (UV-B) radiation, the electron flow from water to dichlorophenol indophenol (DCPIP) was inhibited, but the electron flow from reduced DCPIP to methyl viologen remains unaffected. Diphenylcarbazide was ineffective in restoring the activity of DCPIP Hill reaction in UV-B irradiated chloroplasts. Electron flow from water to ferricyanide or dichloro-dimethoxy- p -benzoquinone was inhibited to a degree similar to that of the DCPIP Hill reaction.
The rate of carotenoid photobleaching in the presence of carbonyl cyanide- m -chlorophenylhydrazone, an indicator of the photochemical reaction near the vicinity of reaction centre of photosystem II, was suppressed and paralleled with the inhibition of the DCPIP Hill reaction.
In the UV-B treated chloroplasts, the variable part of the fluorescence transient was diminished. Though the fluorescence yield was lowered by the UV-B radiation, addition of 3-(3,4-dichlorophenyl)-l, l-dimethylurea (DCMU) and/or sodium dithionite increased the emission markedly. With the increase in the dosage of UV-B irradiation, the time required to reach the steady state fluorescence level became longer in the absence of DCMU and shorter in the presence of DCMU. The kinetics of 520 nm absorbance change was markedly unaltered by the UV-B irradiation but its dark decay was prolonged. It is concluded that UV-B irradiation inactivates the photosystem II reaction centre.     

Evidence for the ultraviolet-B (280–320 nm) radiation induced structural reorganization and damage of photosystem II polypeptides in isolated chloroplasts

Direct evidence for the possible loss of photosystem II (PS II) activity in chloroplasts of Vigna sinensis L. cv. Walp after ultraviolet-B (UV-B, 280–320 nm) radiation treatment was provided by polyacrylamide gel electrophoretic analysis of PS II polypeptides. A 30 min UV-B treatment of chloroplasts caused a 50% loss of PS II activity. The artificial electron donor. Mn²⁺ failed to restore UV-B radiation induced loss of PS II activity, while diphenyl carbazide (DPC) and NH₂OH only partially restored activity. Such a loss in PS II activity was found to be primarily due to a loss of 23 and 33 kDa extrinsic polypeptides. UV-B treatment induced the synthesis of a few polypeptides and a 29 kDa light-harvesting chlorophyll protein.     

Internal filters: Prospects for UV-acclimation in higher plants

Wavelength-selective absorption of solar radiation within plant leaves allows penetration of visible radiation (400–700 nm) to the chloroplasts, while removing much of the damaging ultraviolet-B (UV-B, 280–320 nm) radiation. Flavonoids are important in this wavelength-selective absorption. Induction of flavonoid synthesis by solar radiation, and specifically by UV-B radiation, is discussed as this relates to the potential acclimation of plants to enhanced solar UV-B radiation that would result from stratospheric ozone reduction.     

Time-resolved spectroscopic fluorescence imaging, transient absorption and vibrational spectroscopy of intact and photo-inhibited photosynthetic tissue.

Fluorescence, absorption and vibrational spectroscopic techniques were used to study spinach at the photosystem II (PS II), chloroplast and cellular levels and to determine the effects and mechanisms of ultraviolet-B (UV-B) photoinhibition of these structures. Two-photon fluorescence spectroscopic imaging of intact chloroplasts shows significant spatial variations in the component fluorescence spectra in the range 640-740 nm, indicating that the type and distribution of chlorophylls vary markedly with position in the chloroplast. The chlorophyll distributions and excitonic behaviour in chloroplasts and whole plant tissue were studied using picosecond time-gated fluorescence imaging, which also showed UV-induced kinetic changes that clearly indicate that UV-B induces both structural and excitonic uncoupling of chlorophylls within the light-harvesting complexes. Transient absorption measurements and low-frequency infrared and Raman spectroscopy show that the predominant sites of UV-B damage in PS II are at the oxygen-evolving centre (OEC) itself, as well as at specific locations near the OEC-binding sites.     

Inhibition of violaxanthin deepoxidation by ultraviolet-B radiation in isolated chloroplasts and intact leaves

The effect of pretreatment with ultraviolet-B (UV-B) light (280-320 nanometers) on the enzymatic conversion of the diepoxyxanthophyll violaxanthin to the epoxy-free zeaxanthin occurring in thylakoid membranes was investigated. When isolated chloroplasts of pea (Pisum sativum) were exposed to UV-B, a biologically effective fluence of 7000 joules per square meter caused about 50% inhibition of the activity of the violaxanthin deepoxidase, measured as the first order rate constant of the absorbance change at 505 nanometers. The dose requirement for the inhibition of the deepoxidase in intact leaves, however, was about 2 orders of magnitude higher. The inhibition of the rate constant was observed for both the dark deepoxidation at pH 5, and for the light-driven deepoxidation induced by the lumen acidification due to electron transport from H₂O to methylviologen or due to a photosystem I partial reaction with duroquinol as the electron donor. The availability of violaxanthin was not directly affected by UV-B radiation, as shown for UV-B-treated chloroplasts by the final extent of the 505 nanometer change measured in the dark at pH 5 or by the partial photosystem I reaction. A significant decrease in the violaxanthin availability was observed when lumen acidification was caused by electron transport from H₂O to methylviologen. That effect was probably caused by the wellknown UV-B inhibition of photosystem II with a subsequent decreased ability to reduce the plastoquinone pool, the redox state of which is believed to regulate the final amount of converted violaxanthin.     

Resistance of reaction centers from Rhodobacter sphaeroides against UV-B radiation. Effects on protein structure and electron transport

Inhibition of electron transport and damage to the protein subunits by ultraviolet-B (UV-B, 280–320 nm) radiation have been studied in isolated reaction centers of the non-sulfur purple bacterium Rhodobacter sphaeroides R26. UV-B irradiation results in the inhibition of charge separation as detected by the loss of the initial amplitude of absorbance change at 430 nm reflecting the formation of the P⁺(Q_AQ_B)^– state. In addition to this effect, the charge recombination accelerates and the damping of the semiquinone oscillation increases in the UV-B irradiated reaction centers. A further effect of UV-B is a 2 fold increase in the half- inhibitory concentration of o-phenanthroline. Some damage to the protein subunits of the RC is also observed as a consequence of UV-B irradiation. This effect is manifested as loss of the L, M and H subunits on Coomassie stained gels, but not accompanied with specific degradation products. The damaging effects of UV-B radiation enhanced in reaction centers where the quinone was semireduced (Q_B ^–) during UV-B irradiation, but decreased in reaction centers which lacked quinone at the Q_B binding site. In comparison with Photosystem II of green plant photosynthesis, the bacterial reaction center shows about 40 times lower sensitivity to UV-B radiation concerning the activity loss and 10 times lower sensitivity concerning the extent of reaction center protein damage. It is concluded that the main effect of UV-B radiation in the purple bacterial reaction center occurs at the Q_AQ_B quinone acceptor complex by decreasing the binding affinity of Q_B and shifting the electron equilibration from Q_AQ_B ^– to Q_A ^–Q_B. The inhibitory effect is likely to be caused by modification of the protein environment around the Q_B binding pocket and mediated by the semiquinone form of Q_B. The UV-resistance of the bacterial reaction center compared to Photosystem II indicates that either the Q_AQ_B acceptor complex, which is present in both types of reaction centers with similar structure and function, is much less susceptible to UV damage in purple bacteria, or, more likely, that Photosystem II contains UV-B targets which are more sensitive than its quinone complex.Abbreviations Bchl bacteriochlorophyll - P Bchl dimer - Q_A primary quinone electron acceptor - Q_B secondary quinone electron acceptor - RC reaction center - UV-B ultraviolet-B     

The Regulation of Exogenous Jasmonic Acid on UV-B Stress Tolerance in Wheat

Enhanced ultraviolet-B radiation (UV-B, 280?C320?nm) is recognized as one of the environmental stress factors that cannot be neglected. Jasmonic acid (JA) is an important signaling molecule in a plant??s defense against biotic and abiotic stresses. To determine the role of exogenous JA in the resistance of wheat to stress from UV-B radiation, wheat seedlings were exposed to 0.9?kJ?m^?2?h^?1 UV-B radiation for 12?h after pretreatment with 1 and 2.5?mM JA, and the activity of antioxidant enzymes, the level of malondialdehyde (MDA), the content of UV-B absorbing compounds, photosynthetic pigments, and proline and chlorophyll fluorescence parameters were measured. The results of two-way ANOVA illustrated that the activities of superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT), MDA level, anthocyanin and carotenoid (Car) content, and almost all chlorophyll fluorescence parameters were significantly affected by UV-B, JA, and UV-B?×?JA (P?<?0.05) [the maximal efficiency of photosystem II photochemistry (F _v/F _m) was not affected significantly by UV-B radiation]. Duncan??s multiple-range tests demonstrated that UV-B stress induced a significant reduction in plant photosystem II (PSII) function and SOD activity and an increased level of membrane lipid peroxidation, indicative of the deleterious effect of UV-B radiation on wheat. JA pretreatment obviously mitigated the detrimental effect of UV-B on PSII function by increasing F _v/F _m, reaction centers?? excitation energy capture efficiency (F _v??/F _m??), effective photosystem II quantum yield (??PSII), and photosynthetic electron transport rate (ETR), and by decreasing nonphotochemical quenching (NPQ) of wheat seedlings. Moreover, the activity of SOD and the content of proline and anthocyanin were provoked by exogenous JA. However, the MDA level was increased and Car content was decreased by exogenous JA with or without the presence of supplementary UV-B, whereas the contents of chlorophyll and flavonoids and related phenolics were not affected by exogenous JA. Meanwhile, exogenous JA resulted in a decrease of CAT and POD activities under UV-B radiation stress. These results partly confirm the hypothesis that exogenous JA could counteract the negative effects of UV-B stress on wheat seedlings to some extent.     

短期增强UV-B辐射对青榨槭幼苗生理特性的影响

在中国科学院茂县生态站选择2年生青榨槭幼苗进行室外盆栽实验,以人工增强0.27 W·m^-2(7.7kJ·m^-2·d^-1)的UV-B辐射剂量,研究模拟当地平流层臭氧削减1%时近地面太阳UV-B的增强对木本植物幼苗生理生态适应性的影响.处理0 d后,移除UV-B处理装置,就地测定气体交换参数和叶绿素荧光参数,并取样测定叶绿素及UV吸收物质含量.结果表明,增强UV-B辐射显著降低了青榨槭幼苗叶片最大净光合速率P_max(对照为6.214,处理为4.42),显著增加叶片暗呼吸速率Rd(对照为0.413,处理为1.29)和光补偿点LCP(对照为21.629,处理为9.861),但对表观量子速率a(对照为0.021,处理为0.032)影响不明显;它降低青榨槭幼苗每日净光合速率和水分利用效率,以及原初光化学效率(F_v/F_m)和实际光化学量子产量(ΦPS∏).此外,增强UV-B辐射使叶片叶绿素a、b含量降低(对照为16.23,.39,处理为13.17,4.93),但对叶绿素b含量影响未达显著水平.增强UV-B处理降低了青榨槭幼苗UV吸收物质含量(对照为0.87,处理为0.79).光合指标、叶绿素荧光指标和叶绿素含量的变化有较好的一致性,表明增强UV-B对青榨槭幼苗的光合作用可能有一定的抑制作用.     

The sensitivity of Photosystem II to damage by UV-B radiation depends on the oxidation state of the water-splitting complex

The water-oxidizing complex of Photosystem II is an important target of ultraviolet-B (280-320 nm) radiation, but the mechanistic background of the UV-B induced damage is not well understood. Here we studied the UV-B sensitivity of Photosystem II in different oxidation states, called S-states of the water-oxidizing complex. Photosystem II centers of isolated spinach thylakoids were synchronized to different distributions of the S(0), S(1), S(2) and S(3) states by using packages of visible light flashes and were exposed to UV-B flashes from an excimer laser (lambda=308 nm). The loss of oxygen evolving activity showed that the extent of UV-B damage is S-state-dependent. Analysis of the data obtained from different synchronizing flash protocols indicated that the UV-sensitivity of Photosystem II is significantly higher in the S(3) and S(2) states than in the S(1) and S(0) states. The data are discussed in terms of a model where UV-B-induced inhibition of water oxidation is caused either by direct absorption within the catalytic manganese cluster or by damaging intermediates of the water oxidation process.     

Low radiance in the region of wavelengths 620-660 nm reduces the UV-B-induced damage to photosystem II in spinach leaves

The effect of monochromatic preirradiation in the region of wavelengths 550-730 nm on the parameters of variable and delayed fluorescence in primary spinach leaves irradiated with UV-B was investigated. It was shown that irradiation of leaves with UV-B at lambda(M) = 300 nm reduced photoinduced changes in variable and delayed fluorescence as well as increased the half rise-time of maximum fluorescence. Preirradiation in the region of 620-660 nm (I = 2-4 W x m(-2)) reduced partly the damage to photosystem II induced by UV-B, whereas preirradiation at lambda(m) = 550, 600, 700, and 730 nm had no significant influence. Potential receptors of red light that take part in the protection of photosystem II against UV-B are discussed.     

Epidermal transmittance and phenolic composition in leaves of atrazine-tolerant and atrazine-sensitive cultivars of Brassica napus grown under enhanced UV-B radiation

Experiments were conducted on the atrazine-tolerant mutant Stallion and the atrazine-sensitive cv. Paroll of Brassica napus L., which were grown under either visible light or with the addition of UV-B radiation (280–320 nm) for 15 days. The mutant has been shown to be sensitive to high levels of visible light as compared to the atrazine-sensitive cultivar and therefore we wished to determine plant response to UV-B radiation with respect to potential pigment changes, certain anatomical features, radiation penetration and partial photosynthesis. With regard to pigment changes, we were particularly interested in whether the compositional shift in flavonol pigments under enhanced UV-B radiation, previously suggested to favour increased antioxidant activity, is confined to the adaxial epidermis, which generally receives most UV-B radiation or whether the pigment shift is also inducible in the abaxial epidermis.As was to be expected, the penetration of UV-B radiation (310 nm) was lower in the UV-B-exposed plants, which was correlated with an increased amount of UV-screening pigments in the adaxial and abaxial epidermal layers. The main flavonoid glycosides showed the largest shift from kaempferol to quercetin as aglycone moiety in the adaxial epidermal layer. However, in the abaxial epidermal layer the hydroxycinnamic acid (HCA) derivatives and kaempferol glycosides were predominant. Penetration of 430 nm light was higher after UV-B exposure, and probably contributed to the fact that photosynthetic efficiency of photosystem II was unchanged or higher after UV-B exposure. UV-B radiation decreased leaf area in the atrazine-tolerant mutant only. Both cultivars showed an increased leaf thickness after UV-B exposure due to cell elongation mainly of the palisade tissue. This was especially evident in the mutant.     

Modification of leaf cytology and anatomy in Brassica napus grown under above ambient levels of supplemental UV-B radiation.

Plants exposed to natural solar radiation usually show acclimation responses on a daily and seasonal basis. Many of these responses are complex and modified by interactions with acclimation responses to other climatic factors. While changes in photosynthetically active radiation (PAR, 400-700 nm) are the driving force for many acclimation responses in plants, radiation outside the PAR range is also important. Recently, interest has increased in the potential role of UV-A (320-400 nm) and UV-B (280-320 nm) components of sunlight in plant developmental, physiological and daily acclimation processes. In order to explore the role of UV-B further, Brassica napus L. cv Paroll plants were grown to maturity under 13 kJ d(-1) of biologically effective ultraviolet-B radiation (UV-B(BE), 280-320 nm) plus 800 micromol photons m(-2) s(-1) photosynthetically active radiation (PAR, 400-700 nm) or PAR alone. Leaf anatomy and palisade cell structure were quantified using stereological techniques. The leaves of plants grown under UV-B radiation exhibited an increase in overall leaf width, although no change in leaf anatomy was discerned. Palisade cells in UV-B exposed leaves showed a significant decrease in chloroplast, mitochondrial, starch, and microbody volume density (Vv), while the vacuolar Vv increased compared to cells exposed to PAR only. In UV-B exposed leaves, there was an increase in the appressed and non-appressed thylakoid surface area density (Sv) within the chloroplasts. Since the relative proportion of appressed to non-appressed thylakoid surface area did not change, both thylakoid systems changed in concert with each other. Thylakoid stacks were broader and shorter in leaves subjected to UV-B. In general these responses were similar to those which occurred in plants moved from a high to low PAR environment and similar to mature plants exposed to 13 kJ d(-1) UV-B(BE) for only a short period of time. Although UV absorbing pigments increased by 21% in UV-B exposed leaves, there was no significant difference in chlorophyll a,b or carotenoid content compared to plants exposed to only PAR.     

Impact of ultraviolet radiation on cell structure, UV-absorbing compounds, photosynthesis, DNA damage, and germination in zoospores of Arctic Saccorhiza dermatodea

Stratospheric ozone depletion leads to enhanced UV-B radiation. Therefore, the capacity of reproductive cells to cope with different spectral irradiance was investigated in the laboratory. Zoospores of the upper sublittoral kelp Saccorhiza dermatodea were exposed to varying fluence of spectral irradiance consisting of photosynthetically active radiation (PAR, 400-700 nm; =P), PAR+UV-A radiation (UV-A, 320-400 nm; =PA), and PAR+UV-A+UV-B radiation (UV-B, 280-320 nm; =PAB). Structural changes, localization of phlorotannin-containing physodes, accumulation of UV-absorbing phlorotannins, and physiological responses of zoospores were measured after exposure treatments as well as after 2-6 d recovery in dim white light (8 mumol photon m(-2) s(-1)). Physodes increased in size under PAB treatment. Extrusion of phlorotannins into the medium and accumulation of physodes was induced not only under UVR treatment but also under PAR. UV-B radiation caused photodestruction indicated by a loss of pigmentation. Photosynthetic efficiency of spores was photoinhibited after 8 h exposure to 22 and 30 mumol photon m(-2) s(-1) of PAR, while supplement of UVR had a significant additional effect on photoinhibition. A relatively low recovery of photosystem II function was observed after 2 d recovery in spores exposed to 1.7 x 10(4) J m(-2) of UV-B, with a germination rate of only 49% of P treatment after 6 d recovery. The amount of UV-B-induced DNA damage measured as cyclobutane-pyrimidine dimers (CPDs) increased with the biologically effective UV-B dose (BED(DNA)). Significant removal of CPDs indicating repair of DNA damage was observed after 2 d in low white light. The protective function of phlorotannins has restricted efficiency for a single cell. Within a plume of zoospores, however, each cell can buffer each other and protect the lower layer of spores from excessive radiation. Exudation of phlorotannins into the water can also reduce the impact of UV-B radiation on UV-sensitive spores. The results of this study showed that the impact of UVR on reproductive cells can be mitigated by protective and repair mechanisms.     

Flash-induced carotenoid radical cation formation in Photosystem II

Light excitation of chloroplasts at low temperature produces absorption changes (ΔA) with a large positive peak at 990 nm and a bleaching around 480 nm. ΔA at 990 nm rises with $\text{t}{}_{\mathrm{<mtext>1</mtext><mtext>2</mtext>}}\text{= 0.6}\text{ms}$ at 20–77 K and remains largely stable. This signal is not observed when Photosystem II (PS II) photochemistry is blocked by reduction of the primary plastoquinone. It is observed also in purified PS II particles, in which case it could be shown that during a sequence of short flashes, the absorption at 990 nm rises in parallel with plastoquinone reduction measured at 320 nm. In chloroplasts the light-induced 990-nm ΔA at 77 K is increased under oxidizing conditions (addition of ferricyanide) and upon addition of 2-(3-chloro-4-trifluoromethyl)anilino-3,5-dinitrothiophene (ANT2p). At 21°C, flash excitation of chloroplasts or of PS II particles induces only a very small ΔA at 990 nm, even when this is measured with a 100-ns time resolution or when the material is preilluminated. In both materials, however, a large flash-induced ΔA takes place when various lipophilic anions are added. After a flash the signal rises in less than 100 μs and its decay varies with experimental conditions; the decay is strongly accelerated by benzidine. The difference spectrum measured in PS II particles includes a broad peak around 990 nm and a bleaching around 490 nm. These absorption changes are attributed to a carotenoid radical cation formed at the PS II reaction center. It is estimated that in the presence of lipophilic anions at room temperature, one cation can be formed by a single flash in 80% of the reaction centers. At cryogenic temperature approx. 8% of the PS II reaction centers can oxidize a carotenoid after a single flash.     

The sensitivity of Photosystem II to damage by UV-B radiation depends on the oxidation state of the water-splitting complex

The water-oxidizing complex of Photosystem II is an important target of ultraviolet-B (280-320 nm) radiation, but the mechanistic background of the UV-B induced damage is not well understood. Here we studied the UV-B sensitivity of Photosystem II in different oxidation states, called S-states of the water-oxidizing complex. Photosystem II centers of isolated spinach thylakoids were synchronized to different distributions of the S₀, S₁, S₂ and S₃ states by using packages of visible light flashes and were exposed to UV-B flashes from an excimer laser (λ = 308 nm). The loss of oxygen evolving activity showed that the extent of UV-B damage is S-state-dependent. Analysis of the data obtained from different synchronizing flash protocols indicated that the UV-sensitivity of Photosystem II is significantly higher in the S₃ and S₂ states than in the S₁ and S₀ states. The data are discussed in terms of a model where UV-B-induced inhibition of water oxidation is caused either by direct absorption within the catalytic manganese cluster or by damaging intermediates of the water oxidation process.     

Energy and electron transfer in photosystem II of a chlorophyll b-containing Synechocystis sp. PCC 6803 mutant

Using a Synechocystis sp. PCC 6803 mutant strain that lacks photosystem (PS) I and that synthesizes chlorophyll (Chl) b, a pigment that is not naturally present in the wild-type cyanobacterium, the functional consequences of incorporation of this pigment into the PS II core complex were investigated. Despite substitution of up to 75% of the Chl a in the PS II core complex by Chl b, the modified PS II centers remained essentially functional and were able to oxidize water and reduce Q(A), even upon selective excitation of Chl b at 460 nm. Time-resolved fluorescence decay measurements upon Chl excitation showed a significant reduction in the amplitude of the 60-70 ps component of fluorescence decay in open Chl b-containing PS II centers. This may indicate slower energy transfer from the PS II core antenna to the reaction center pigments or slower energy trapping. Chl b and pheophytin b were present in isolated PS II reaction centers. Pheophytin b can be reversibly photoreduced, as evidenced from the absorption bleaching at approximately 440 and 650 nm upon illumination in the presence of dithionite. Upon excitation at 685 nm, transient absorption measurements using PS II particles showed some bleaching at 650 nm together with a major decrease in absorption around 678 nm. The 650 nm bleaching that developed within approximately 10 ps after the flash and then remained virtually unchanged for up to 1 ns was attributed to formation of reduced pheophytin b and oxidized Chl b in some PS II reaction centers. Chl b-containing PS II had a lower rate of charge recombination of Q(A)(-) with the donor side and a significantly decreased yield of delayed luminescence in the presence of DCMU. Taken together, the data suggest that Chl b and pheophytin b participate in electron-transfer reactions in PS II reaction centers of Chl b-containing mutant of Synechocystis without significant impairment of PS II function.     

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.     

Sudden exposure to solar UV-B radiation reduces net CO(2) uptake and photosystem I efficiency in shade-acclimated tropical tree seedlings

Tree seedlings developing in the understory of the tropical forest have to endure short periods of high-light stress when tree-fall gaps are formed, and direct solar radiation, including substantial UV light, reaches the leaves. In experiments simulating the opening of a tree-fall gap, the response of photosynthesis in leaves of shade-acclimated seedlings (Anacardium excelsum, Virola surinamensis, and Calophyllum longifolium) to exposure to direct sunlight (for 20-50 min) was investigated in Panama (9 degrees N). To assess the effects of solar UV-B radiation (280-320 nm), the sunlight was filtered through plastic films that selectively absorbed UV-B or transmitted the complete spectrum. The results document a strong inhibition of CO(2) assimilation by sun exposure. Light-limited and light-saturated rates of photosynthetic CO(2) uptake by the leaves were affected, which apparently occurred independently of a simultaneous inhibition of potential photosystem (PS) II efficiency. The ambient UV-B light substantially contributed to these effects. The photochemical capacity of PSI, measured as absorbance change at 810 nm in saturating far-red light, was not significantly affected by sun exposure of the seedlings. However, a decrease in the efficiency of P700 photooxidation by far-red light was observed, which was strongly promoted by solar UV-B radiation. The decrease in PSI efficiency may result from enhanced charge recombination in the reaction center, which might represent an incipient inactivation of PSI, but contributes to thermal dissipation of excessive light energy and thereby to photoprotection.