Effect of common bean seedling pretreatment with chlorocholine chloride on photosystem II tolerance to UV-B radiation,phytohormone content,and hydrogen peroxide content

In true leaves of common bean (Phaseolus vulgaris L., cv. Berbukskaya) 11-day-old seedlings, pretreatment with 1.6 mM chlorocholine chloride (CCC) resulted in the improved photosystem II (PSII) tolerance to UV-B radiation, an increase in the contents of cytokinins, ABA, and H₂O₂, but a decrease in the content of gibberellins. It is suggested that development of increased PSII tolerance to UV-B under the influence of CCC is partially related to the increase in the contents of ABA, cytokinins, and UV-absorbing pigments in the leaves of pretreated plants.     

Growth under UV-B radiation increases tolerance to high-light stress in pea and bean plants

Pea (Pisum sativum L.) and bean (Phaseolus vulgaris L.) plants were exposed to enhanced levels of UV-B radiation in a growth chamber. Leaf discs of UV-B treated and control plants were exposed to high-light (HL) stress (PAR: 1200 mol m^–2 s^–1) to study whether pre-treatment with UV-B affected the photoprotective mechanisms of the plants against photoinhibition. At regular time intervals leaf discs were taken to perform chlorophyll a fluorescence and oxygen evolution measurements to assess damage to the photosystems. Also, after 1 h of HL treatment the concentration of xanthophyll cycle pigments was determined. A significantly slower decline of maximum quantum efficiency of PSII (F _v/F _m), together with a slower decline of oxygen evolution during HL stress was observed in leaf discs of UV-B treated plants compared to controls in both plant species. This indicated an increased tolerance to HL stress in UV-B treated plants. The total pool of xanthophyll cycle pigments was increased in UV-B treated pea plants compared to controls, but in bean no significant differences were found between treatments. However, in bean plants thiol concentrations were significantly enhanced by UV-B treatment, and UV-absorbing compounds increased in both species, indicating a higher antioxidant capacity. An increased leaf thickness, together with increases in antioxidant capacity could have contributed to the higher protection against photoinhibition in UV-B treated plants.     

Effects of supplemental UV-B radiation on primary photosynthetic carboxylating enzymes and soluble proteins in leaves of C3 and C4 crop plants

Effects of increased UV-B radiation on activities of primary photosynthetic carboxylating enzymes and on contents of soluble proteins were studied in soybean (Glycine max [L.] Merr. cv. Bragg), pea (Pisum sativum L. cv. Little Marvel), tomato (Lycopersicon esculentum L. cv. Rutgers), and sweet corn (Zea mays L. cv. Golden Cross Bantam). The purpose was to evaluate the responses of agronomic crops to increases in solar UV-B radiation. Plants were grown and exposed under greenhouse conditions for 6 h daily to supplemental UV-B radiation which was provided by Westinghouse FS-40 fluorescent sun lamps filtered with 0.127-mm film of cellulose acetate (UV-B treated) or Mylar S (Mylar control). Three UV-B levels were tested: 1.09 (treatment T₁), 1.36 (treatment T₂), and 1.83 (treatment T₃) UV-B_seu where 1 UV-B_seu equals 16.0 mW-m² weighted by EXP-[(λ-265)/21]². These UV-B levels corresponded to 6%,21%, and 36%, respectively, of decrease in stratospheric ozone content, based on the interpolations of UV-B irradiances at a solar elevation angle of 60°. Leaves of plants of soybean, pea, and tomato exposed to UV-B radiation were generally low in RuBP carboxylase activity. On a fresh weight basis, all three UV-B radiation levels significantly reduced the enzyme activity in soybean and pea, whereas tomato plants showed significant reduction in RuBP carboxylase activity only when exposed to 1.83 and 1.36 UV-B_seu. An apparent decrease in soluble proteins was observed in leaf extracts of soybean and pea plants exposed to 1.36 and 1.83 UV-B_seu whereas higher amounts of proteins were detected in leaves of tomato plants grown under UV-B radiation. Leaves of sweet corn plants grown under Mylar control were low in PEP carboxylase activity and proteins as compared with those of control plants receiving no supplemental UV and UV-B treatment. Activities of PEP carboxylase in crode extracts from leaves of sweet corn were significantly suppressed under 1.36 and 1.83 UV-B_seu as compared with the no UV control. Some stimulation of PEP carboxylase activity was observed in corn plants exposed to 1.09 UV-B_seu.     

Ultraviolet-B radiation causes shade-type ultrastructural changes in Brassica napus

Cell and chloroplast structural changes in palisade cells from mature leaves of Brassica napus L. cv. Paroll were quantified following exposure of plants to enhanced ultraviolet-B (280–320 nm; 13 kJ m^?2 day^?1 biologically effective UV-B) radiation at two different levels of photosynthetically active radiation (PAR, 400–700 nm; 200 and 700 μmol m^?2 s^?1). Short-term changes in leaf ultrastructure after 30 min and longer term changes after one day and one week were analyzed using stereological techniques incorporating light and electron microscopy and mathematical reconstruction of a mean cell for each sample. Ultraviolet-B together with either relatively high or low PAR resulted in cell structural changes resembling those typical of plants under shade conditions, with the most marked response occurring after 30 min of UV-B radiation. The ultrastructural changes at the cellular level were generally similar in both the relatively high and low PAR plus UV-B radiation treatments. The surface areas of all three thylakoid types, the appressed, non-appressed and margin thylakoids increased in the palisade tissue under supplemental UV-B irradiation. Although the appressed and non-appressed thylakoids increased in surface area, they did not increase equally, leaving open the possibility that the two thylakoid types have independent regulatory systems or different sensitivity to UV-B radiation. Increased thylakoid packing (mm² thylakoid membrane per mm² leaf surface) in UV-B-exposed plants may increase the statistical probability of photon interception. An increased level of UV-absorbing pigments after one week of supplemental UV-B radiation did not prevent or significantly ameliorate UV effects. Our data supported the assumption that UV-B radiation may have a regulatory role besides damaging effects and that an increased UV-B environment will likely increase this regulatory influence of UV-B radiation.     

Evaluation of the role of damage to photosystem II in the inhibition of CO2 assimilation in pea leaves on exposure to UV-B radiation

Mature pea (Pisum sativum L., cv. Meteor) leaves were exposed to two levels of UV-B radiation, with and without supplementary UV-C radiation, during 15 h photoperiods. Simultaneous measurements of CO₂ assimilation and modulated chlorophyll fluorescence parameters demonstrated that irradiation with UV-B resulted in decreases in CO₂ assimilation that are not accompanied by decreases in the maximum quantum efficiency of photosystem II (PSII) primary photochemistry. Increased exposure to UV-B resulted in a further loss of CO₂ assimilation and decreases in the maximum quantum efficiency of PSII primary photochemistry, which were accompanied by a loss of the capacity of thylakoids isolated from the leaves to bind atrazine, thus demonstrating that photodamage to PSII reaction centres had occurred. Addition of UV-C to the UV-B treatments increased markedly the rate of inhibition of photosynthesis, but the relationships between CO₂ assimilation and PSII characteristics remained the same, indicating that UV-B and UV-C inhibit leaf photosynthesis by a similar mechanism. It is concluded that PSII is not the primary target site involved in the onset of the inhibition of photosynthesis in pea leaves induced by irradiation with UV-B.     

Response of photosynthesis to high light and drought for Arabidopsis thaliana grown under a UV-B enhanced light regime

Arabidopsis thaliana grown in a light regime that included ultraviolet-B (UV-B) radiation (6 kJ m⁻² d⁻¹) had similar light-saturated photosynthetic rates but up to 50% lower stomatal conductance rates, as compared to plants grown without UV-B radiation. Growth responses of Arabidopsis to UV-B radiation included lower leaf area (25%) and biomass (10%) and higher UV-B absorbing compounds (30%) and chlorophyll content (52%). Lower stomatal conductance rates for plants grown with UV-B radiation were, in part, due to lower stomatal density on the adaxial surface. Plants grown with UV-B radiation had more capacity to down regulate photochemical efficiency of photosystem II (PSII) as shown by up to 25% lower φ_PSII and 30% higher non-photochemical quenching of chlorophyll fluorescence under saturating light. These contributed to a smaller reduction in the maximum photochemical efficiency of PSII (F _v/F _m), greater dark-recovery of F _v/F _m, and higher light-saturated carbon assimilation and stomatal conductance and transpiration rates after a four-hour high light treatment for plants grown with UV-B radiation. Plants grown with UV-B were more tolerant to a 12 day drought treatment than plants grown without UV-B as indicated by two times higher photosynthetic rates and 12% higher relative water content. UV-B-grown plants also had three times higher proline content. Higher tolerance to drought stress for Arabidopsis plants grown under UV-B radiation may be attributed to both increased proline content and decreased stomatal conductance. Growth of Arabidopsis in a UV-B-enhanced light regime increased tolerance to high light exposure and drought stress.     

Effects of Enhanced Ultraviolet-B (280–320 nm) Radiation on Growth and Photosynthetic Activities in Aquatic Fern Azolla Microphylla Kaulf

The effects of enhanced UV-B radiation on growth and photosynthetic activities were investigated in fronds of the aquatic fern Azolla microphylla Kaulf. The fronds were exposed to UV-B radiation intermittently once in 3 d during 12 d. Biomass and relative growth rate of UV-B treated Azolla plants and the heterocyst frequency of the UV-B treated symbiont decreased resulting in an increase in doubling time over the control. The doubling time was 3.08 d for control and 3.35 d for UV-B irradiated plants. Chl and carotenoid contents per unit fresh mass and photosystem 2 (PS2) activity also decreased under UV-B treatment. Measurements of photosynthetic activity in terms of fluorescence kinetics and PS2 mediated O₂ evolution showed that the aquatic fern Azolla is sensitive to UV-B damage.     

Comparison of responses of bean, pea and rape plants to UV-B radiation in darkness and in light

Effect of UV-B radiation on leaves of bean, pea and rape plants was studied. UV-B radiation (11.2 kJ·m⁻²) induced more distinct reduction of the primary photosynthesis activity when applied in darkness than the same UV-B dose, extended in time, and applied with photosynthetic active radiation (PAR). The pea plants were more susceptible to UV-B in darkness, but in the presence of PAR their tolerance was higher. The CO₂ fixation in the bean and rape plants, exposed to UV-B was decreased, but for the pea plants it remained unchanged. The UV-B irradiation caused an increase in the content of ultraviolet-absorbing pigments. Additionally, the bean plants grown at UV-B increased the thickness of leaves, described as SLW.     

Combined effects of elevated UV-B radiation and the addition of selenium on common (<Emphasis Type="Italic">Fagopyrum esculentum</Emphasis> Moench) and tartary [<Emphasis Type="Italic">Fagopyrum tataricum</Emphasis> (L.) Gaertn.] buckwheat

The combined effects of UV-B irradiation and foliar treatment with selenium on two buckwheat species, common (Fagopyrum esculentum Moench) and tartary [Fagopyrum tataricum (L.) Gaertn.] buckwheat, that underwent different intensity of breeding, were examined. Plants grown outdoors under three levels of UV-B radiation were studied for 9 weeks, from sowing to ripening. At week 7 they were sprayed with solution containing 1 g(Se) m⁻³ that presumably mitigates UV-B stress. Morphological, physiological, and biochemical parameters of the plants were monitored. Elevated UV-B radiation, corresponding to a 17 % reduction of the ozone layer, induced synthesis of UV absorbing compounds. In both buckwheat species it also caused a reduction in amounts of chlorophyll a during the time of intensive growth, an effect, which was increased in tartary buckwheat in the presence of selenium. The respiratory potential, measured as terminal electron transport system activity, was lower in plants subjected to enhanced UV-B radiation during the time of intensive growth. The effective quantum yield of photosystem 2 was also reduced due to UV-B radiation in both buckwheat species and was mitigated by the addition of Se. Se treatment also mitigated the stunting effect of UV-B radiation and the lowering of biomass in common buckwheat.     

Pre-Treatment of Bean Seedlings with Choline Compounds Increases the Resistance of Photosynthetic Apparatus to UV-B Radiation and Elevated Temperatures

Bean (Phaseolus vulgaris L. cv. Berbukskaya) seedlings were pre-treated with choline compounds, 19 mM 2-ethyltrimethylammonium chloride (Ch) or 1.6 mM 2-chloroethyltrimethylammonium chloride (CCh), during 24 h, then after 6 d the excised primary leaves were exposed to UV-B and high temperature stress. Chlorophyll (Chl) fluorescence, delayed light emission, accumulation of photosynthetic pigments, contents of thiobarbituric acid reactive substances, and activities of the active oxygen detoxifying enzymes (superoxide dismutase, ascorbate peroxidase, and glutathione reductase) were examined. Pre-treatment of plants with Ch or CCh enhanced the resistance of photosystem 2 (PS2) photochemistry to UV-B and heat injuries. The higher stress resistance can be explained by the increased activity of the detoxifying enzymes. The increased content of UV-B-absorbing pigments may also contribute to the enhanced resistance of choline-treated plants to UV-B radiation.     

Uptake hydrogenase in Rhizobium and nodule leghemoglobin in cow pea miscellany hosts

Two effective strains of green gram rhizobia S24 (slow growing and Hup⁺) and M11 (fast growing and Hup^-) were tested for leghemoglobin production in nodules and effectivity on six species of cow pea miscellany hosts. Both strains nodulate green gram [Vigna radiata (L.) (Wilczek)], black gram [Vigna mungo (L.) (Hepper)], cow pea [Vigna unguiqulata (L.)], moth bean [Vigna aconitifolia (Jacq.) (Marechel)], Cluster bean [Cyamopsis tetragonoloba (L.) (Taub.)] and pigeon pea [Cajanus cajan (L.)]. In all these hosts, nodules formed by strain M11 contained 1.5 to 2 times more leghemoglobin than the nodules formed by strain S24. Gel electrophoresis of nodule contents of different host species showed a high concentration of a fast-moving ferricoxy leghemoglobin in the nodules of plants inoculated with strain M11 as compared to that of strain S24. Strain M11, however, was relatively less effective than strain S24 on black gram, cow pea and moth bean and was at par with the later on green gram, cluster bean and pigeon pea. Hydrogen recycling ability of the strain S24 was observed in nodules of all the host species. The effective functioning of strain S24 at low levels of leghemoglobin suggests an involvement of recycling hydrogenase in maintaining an appropriate oxidation-reduction potential in nodules.Abbreviations Lb Leghemoglobin - Cvr cultivar     

Short-term UV-B radiation effects on morphology,physiological traits and accumulation of bioactive compounds in Prunella vulgaris L.

Prunella vulgaris plants (full-bloom stage) were subjected to short-term (15 days) UV-B radiation in a growth chamber. UV-B radiation was effective at enhancing morphological and biomass characteristics and decreasing chlorophyll contents of P. vulgaris. Treatment of P. vulgaris with artificial UV-B radiation significantly increased peroxidase (POD), superoxide dismutase (SOD) and glutathione (GSH) activities compared to the control treatment. UV-B radiation significantly increased the levels of hydrogen peroxide (H₂O₂), malondialdehyde (MDA) and proline in leaves of P. vulgaris compared to those of control plants. In addition, the contents of total flavonoids, rosmarinic acid, caffeic acid and hyperoside significantly increased under UV-B radiation. The total phenolic levels also increased under UV-B treatment. These results demonstrated that short-term UV-B radiation can enhance production of secondary metabolites in P. vulgaris, resulting in increased spica yield compared to that of control plants.     

Time sequence of the damage to the acceptor and donor sides of photosystem II by UV-B radiation as evaluated by chlorophyll a fluorescence

The effects of ultraviolet-B (UV-B) radiation on photosystem II (PS II) were studied in leaves of Chenopodium album. After the treatment with UV-B the damage was estimated using chlorophyll a fluorescence techniques. Measurements of modulated fluorescence using a pulse amplitude modulated fluorometer revealed that the efficiency of photosystem II decreased both with increasing time of UV-B radiation and with increasing intensity of the UV-B. Fluorescence induction rise curves were analyzed using a mechanistic model of energy trapping. It appears that the damage by UV-B radiation occurs first at the acceptor side of photosystem II, and only later at the donor side.     

Effect of Cd and UV-B radiation on polypeptide composition and photosystem activities ofVigna unguiculata chloroplasts

Rates of whole chain and photosystem 2 activities in chloroplasts isolated fromVigna unguiculata L. seedlings grown under ultraviolet-B (UV-B) enhanced radiation were less affected by 3, 6 and 9 mM CdCl₂ for 60 min at 0 °C in the dark than the rates in chloroplasts from control plants grown under normal irradiation. The results are in agreement with changes in contents of chloroplast 55, 47, 43, 33, 29, 27–25, 23 and 17 kDa polypeptides that were significantly lowered at 3, 6 and 9 mM CdCl₂ only in chloroplasts from control plants. On the other hand, in the simultaneous treatment of chloroplast isolated from control plants the UV-B supported the inhibitory effect of all applied concentrations of CdCl₂. The photosystem 1 activity was only marginally affected in the all experimental variants.     

Influence of UV-B radiation on photosynthetic activity and chlorophyll fluorescence kinetics in Norway spruce [Picea abies (L.) Karst.] seedlings

Norway spruce [Picea abies (L.) Karst.] seedlings were grown in greenhouses with two supplemental levels of ultraviolet-B (UV-B) radiation. Photochemical efficiency of photosystem II and vitality index were determined monthly. At the end of the experiment, growth, chlorophyll content and photosynthetic rates were measured. The data indicate that low temperature in winter affected light dependent processes in experimental plants including control, while the rise of ambient temperatures, moderate this effect. The synergistic effects of UV-B radiation and low temperatures could only be observed in the second winter period. Measurements of net photosynthetic activity in the second winter period showed significant differences between treated and untreated plants.     

UV-B induced stress responses in three rice cultivars

UV-B responses of three rice (Oryza sativa L.) cultivars (Sasanishiki, Norin 1 and Surjamkhi) with different photolyase activity were investigated. Carbon dioxide assimilation data support that Sasanishiki was less sensitive to UV-B than Norin 1 and Surjamkhi. UV-B radiation sharply decreased the content of Rubisco protein in Surjamkhi and has no effect in Sasanishiki. The photochemical activities of photosystem (PS) 1 and PS 2 was slightly affected by UV-B treatment. The content of H₂O₂ and the activities of antioxidant enzymes, catalase (CAT), peroxides (POX) and superoxide dismutase (SOD) were enhanced after UV-B treatment. The activities of CAT and POX isoenzymes in Sasanishiki were more enhanced by UV-B radiation than those in Norin 1 and Surjamkhi.     

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.     

Some effects of enhanced UV-B irradiation on the growth and composition of plants

Barley (Hordeum vulgare), corn (Zea mays), bean (Phaseolus vulgaris), and radish (Raphanus sativus) seedlings were continuously irradiated under a lighting device for 5–10 d at an increased ultraviolet (UV)-B fluence rate. In their growth parameters, composition, and leaf surface, these four species responded differently to the increased UV-B exposure. Bean seedlings suffered the most serious effects, radish and barley less, and corn was hardly influenced at all. In all plant species, the fresh weight, the leaf area, the amounts of chlorophylls, carotenoids and the galactolipids of the chloroplasts were reduced. The lipid content of the corn and bean seedlings also diminished. But all the irradiated plants showed a rise in their protein content compared to the control plants. The content of flavonoids increased in barley and radish seedlings by about 50%. The effects on growth parameters and composition were more extensive with increasing UV-B fluence rates, at least as shown in the case of barley seedlings. The fresh weights fell proportionally with the chlorophylls and carotenoids. In contrast, the flavonoid content of barley leaves rose parallel to the increasing UV-B fluence rates and reached 180% of the value in the control plants with the highest UV-B fluence rate. Scorching appeared regularly in the form of bronze leaf discoloration at the highest UV-B fluence rates. Scanning electron micrographs of the leaf surface of UV-B irradiated plants showed deformed epidermal structures.Abbreviations MGDG monogalactosyldiglyceride - DGDG digalactosyldiglyceride - SL sulfoquinovosyldiglyceride - PG phosphatidylglycerol - PC phosphatidylcholine - PE phosphatidylethanolamine - PI phosphatidylinositol - LA leaf are - FW fresh weight - DW dry weight - SEM scanning electron microscopy - C total carotenoids - Chl total chlorophyll     

Temperature is the key to altitudinal variation of phenolics in Arnica montana L. cv. ARBO

Plants in alpine habitats are exposed to many environmental stresses, in particular temperature and radiation extremes. Recent field experiments on Arnica montana L. cv. ARBO indicated pronounced altitudinal variation in plant phenolics. Ortho-diphenolics increased with altitude compared to other phenolic compounds, resulting in an increase in antioxidative capacity of the tissues involved. Factors causing these variations were investigated by climate chamber (CC) experiments focusing on temperature and ultraviolet (UV)-B radiation. Plants of A. montana L. cv. ARBO were grown in CCs under realistic climatic and radiation regimes. Key factors temperature and UV-B radiation were altered between different groups of plants. Subsequently, flowering heads were analyzed by HPLC for their contents of flavonoids and caffeic acid derivatives. Surprisingly, increased UV-B radiation did not trigger any change in phenolic metabolites in Arnica. In contrast, a pronounced increase in the ratio of B-ring ortho-diphenolic (quercetin) compared to B-ring monophenolic (kaempferol) flavonols resulted from a decrease in temperature by 5°C in the applied climate regime. In conclusion, enhanced UV-B radiation is probably not the key factor triggering shifts in the phenolic composition in Arnica grown at higher altitudes but rather temperature, which decreases with altitude.