Effects of UV-B radiation on photosynthetic reactions in Rhizophora apiculata

Influences of UV-B radiation on Rhizophora apiculata were studied in terms of chlorophylls, their presence in protein complexes of the chloroplast, PS I and PS II photochemical activities, in vitro absorption spectrum of the chloroplast, in vivo leaf fluorescence and UV absorbing compounds. The seedlings were exposed to the various levels of UV-B radiations, equivalent to 0 (control), 10, 20, 30 and 40% stratospheric ozone depletion of the study area. The low doses of UV-B (10 and 20%) increased the reaction centre chlorophyll (10 and 8%) and activities of PS-I (98 and 39%) and PS-II (77 and 38%) respectively; whereas, 30 and 40% UV-B treatments decreased the reaction centre chlorophylls by 11 and 33% and PS II activity by 0 and 20%; while PS I activity did not show any inhibitory effect. Chloroplasts isolated from control and 10% UV-B treated plants exhibited the same level of absorption at 676 nm. In vivo leaf fluorescence was found to be diminished with UV-B radiation and at the 10% UV-B, variable fluorescence was promoted significantly by 10%. The content of UV-absorbing compounds was progressively enhanced with doses of UV-B radiation along with higher absorption at 276 and 330 nm.     

Photosynthetic UV-B Response of Beech (Fagus sylvatica L.) Saplings

Cloned saplings of beech (7-y-old) were exposed to enhanced UV-B irradiation (+25 %) continuously over three growing seasons (1999–2001). Analysis of CO₂ assimilation, variable chlorophyll (Chl) a fluorescence, and pigment composition was performed in late summer of the third growing season to evaluate the influence of long-term elevated UV-B irradiation. This influence was responsible for the stimulation of the net assimilation rate (P _N) over a range of irradiances. The increase in P _N was partially connected to increase of the area leaf mass, and thus to the increased leaf thickness. Even a higher degree of UV-B induced stimulation was observed at the level of photosystem 2 (PS2) photochemistry as judged from the irradiance response of electron transport rate and photochemical quenching of Chl a. The remarkably low irradiance-induced non-photochemical quenching of maximum Chl a fluorescence (NPQ) in the UV-B plants over the entire range of applied irradiances was attributed both to the reduced demand on non-radiative dissipation processes and to the considerably reduced contribution of the quenching localised in the inactivated PS2 reaction centres. Neither the content of Chls and total carotenoids expressed per leaf area nor the contents of lutein, neoxanthin, and the pool of xanthophyll cycle pigments (VAZ) were affected under the elevated UV-B. However, the contributions of antheraxanthin (A) and zeaxanthin (Z) to the entire VAZ pool in the dark-adapted UV-B treated plants were 1.61 and 2.14 times higher than in control leaves. Surprisingly, the retained A+Z in UV-B treated plants was not accompanied with long-term down-regulation of the PS2 photochemical efficiency, but it facilitated the non-radiative dissipation of excitation energy within light-harvesting complexes (LHC) of PS2. Thus, in the beech leaves the accumulation of A+Z, induced by other factors than excess irradiance itself, supports the resistance of PS2 against combined effects of high irradiance and elevated UV-B.     

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.     

Photosynthetic responses to solar UV-A and UV-B radiation in low-and high-altitude populations of <Emphasis Type="Italic">Hippophae rhamnoides</Emphasis>

Two contrasting sea buckthorn (Hippophae rhamnoides L.) populations from the low (LA) and high (HA) altitudinal regions were employed to evaluate the plant physiological responses to solar UV-A radiation and near-ambient UV-B radiation (UV-B+A) under the sheltered frames with different solar ultraviolet radiation transmittance. LA-population was more responsive to solar UV-A. Some modification caused by UV-A only existed in LA-population, such as significant reduction of leaf size, relative water content, and chlorophyll (Chl) b content as well as δ¹³C elevation, coupled with larger increase of contents of total carotenoids (Cars). This higher responsiveness might be an effective pre-acclimation strategy adapting for concomitant solar UV-B stress. Near-ambient UV-B+A radiation caused significant reduction of leaf size and Chl content as well as slight down-regulation of photosystem 2 activity that paralleled with higher heat dissipation, while photosynthetic rate was modestly but significantly increased. The higher photosynthesis under near-ambient UV-B+A radiation could be related to pronounced increase of leaf thickness and effective physiological modification, like the increase of leaf protective pigments (Cars and UV-absorbing compound), constant high photochemical capacity, and improved water economy.     

Changes in photosynthetic apparatus during dark incubation of detached leaves from control and ultraviolet-B treatedVigna seedlings

Changes in various components of photosynthetic apparatus during the 4 d dark incubation at 25°C of detached control and ultraviolet-B (UV-B) treatedVigna unguiculata L. leaves were examined. The photosynthetic apparatus was more degraded in younger control seedlings and for a longer time UV-B treated seedlings than in the older or for a shorter time UV-B treated seedlings. This was shown by determining the losses in chlorophyll (Chl) and protein contents, variable fluorescence yield, photosystem (PS) 2, PS1 and ribulose-1,5-bisphosphate carboxylase (RuBPC) activities, and photosynthetic¹⁴CO₂ fixation. In contrast, the Car/Chl ratio increased during the dark incubation due to less expressed degradation of Car.     

Chilling stress and chilling tolerance of sweet potato as sensed by chlorophyll fluorescence

We studied changes in the chlorophyll (Chl) fluorescence components in chilling-stressed sweet potato (Ipomoea batatas L. Lam) cv. Tainung 57 (TN57, chilling-tolerant) and cv. Tainung 66 (TN66, chilling-susceptible). Plants under 12-h photoperiod and 400 μmol m⁻² s⁻¹ irradiance at 24/20 °C (day/night) were treated by a 5-d chilling period at 7/7 °C. Compared to TN66, TN57 exhibited a significantly greater basic Chl fluorescence (F₀), maximum fluorescence (F_m), maximum fluorescence yield during actinic irradiation (F_m′ ), and the quantum efficiency of electron transport through photosystem 2, PS2 (Φ_PS2). Chilling stress resulted in decrease in the potential efficiency of PS2 (F_v/F_m), Φ_PS2, non-photochemical fluorescence quenching (NPQ), non-photochemical quenching (q_N), and the occurrence of chilling injury in TN66. Chilling increased the likelihood of photoinhibition, characterized by a decline in the Chl fluorescence of both cultivars, and photoinhibition during low temperature stress generally occurred more rapidly in TN66.     

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     

Enhanced photosystem 2 thermostability during leaf growth of Elm (Ulmus pumila) seedlings

We examined photosynthetic activities and thermostability of photosystem 2 (PS2) in leaves of elm (Ulmus pumila) seedlings from initiation to full expansion. During leaf development, net photosynthetic rate (P _N) increased gradually and reached the maximum when leaves were fully developed. In parallel with the increase of P _N, chlorophyll (Chl) content was significantly elevated. Chl a fluorescence measurements showed that the maximum quantum yield of PS2 (ϕ_PS2), the efficiency a trapped exciton, moved an electron into the electron transport chain further than Q_A ⁻ (Ψ_o), and the quantum yield of electron transport beyond Q_A (ϕ_Eo) increased gradually. These results were independently confirmed by our low irradiance experiments. When subjected to progressive heat stress, the young leaves exhibited considerably lower ϕ_PS2 and higher minimal fluorescence (F₀) than the mature leaves, revealing the highly sensitive nature of PS2 under heat in the newly initiating leaves. Further analysis showed that PS2 structure in the newly initiating leaves was strongly altered under heat, as evidenced by the increased fluorescence signals at the position of the K step. We therefore demonstrated an inhibition in the oxygen-evolving complex (OEC) in the young leaves. This resulted in decrease in amount of the functional PS2 reaction centres and relative increase in the PS2 reaction centres with inhibited electron transport at the acceptor side under heat. We suggest that the enhanced thermostability of PS2 during leaf development is associated with improved OEC stability.     

Comparison of Aerial and Submerged Leaves in Two Amphibious Species,Myosotis scorpioides and Ranunculus trichophyllus

Both amphibious species, Myosotis scorpioides and Ranunculus trichophyllus, thrive in a stressful environment (alternated flooding and drying), which is variable regarding water and radiation regimes. Plants from the field and plants grown under controlled water table maintained at 40 cm were analysed for content of chlorophyll (Chl) and UV-B screening compounds, and the efficiencies of PS2 and electron transport systems. We detected no significant differences in contents of Chl and UV-B screening compounds between submerged and aerial leaves. The measurements of respiratory potential and photochemical efficiency revealed the presence of permanent stress in M. sporpioides in the natural environment. Differences in physiological responses of submerged and aerial leaves indicated that the terrestrial environment was more favourable for M. scorpioides than for R. trichophyllus. Characteristics of both species suggested that R. trichophyllus might be a phylogenetically older aquatic plant than M. scorpioides.     

Changes in photosynthetic apparatus during dark incubation of detached leaves from control and ultraviolet-B treatedVigna seedlings

Changes in various components of photosynthetic apparatus during the 4 d dark incubation at 25°C of detached control and ultraviolet-B (UV-B) treatedVigna unguiculata L. leaves were examined. The photosynthetic apparatus was more degraded in younger control seedlings and for a longer time UV-B treated seedlings than in the older or for a shorter time UV-B treated seedlings. This was shown by determining the losses in chlorophyll (Chl) and protein contents, variable fluorescence yield, photosystem (PS) 2, PS1 and ribulose-1,5-bisphosphate carboxylase (RuBPC) activities, and photosynthetic¹⁴CO₂ fixation. In contrast, the Car/Chl ratio increased during the dark incubation due to less expressed degradation of Car.     

Carotenoid composition and photochemical activity of four sandy grassland species

The photosynthetic pigments and photochemical efficiency of photosystem 2 (PS2) were studied in four constitutive species (Achillea millefolium L., Festuca pseudovina Hack. ex Wiesb., Potentilla arenaria Borkh., and Thymus degenianus Lyka) of a semiarid grassland in South-eastern Hungary. Every species displayed typical sun-adapted traits and substantial plasticity in the composition and functioning of the photosynthetic apparatus. The contents of chlorophylls (Chls) and carotenoids (Cars) on a dry matter basis declined from May to July, however, the amount of total Cars on a Chl basis increased. This increase was the largest in Potentilla (48 %) and the smallest in Achillea (14 %). The pool of xanthophylls (VAZ) was between 25 % and 45 % of the total Car content and was larger in July than in May. The content of β-carotene increased by July, but lutein content did not change significantly. The Chl fluorescence ratio F_v/F_m was reduced by 3–10 % at noon, reflecting the down-regulation of PS2 in the period of high irradiance and high temperature. The occurrence of minimal values of ΔF/F_m’ showed close correlation to the de-epoxidation rate of violaxanthin. Hence in natural habitats these species developed a considerable capacity to dissipate excess excitation energy in the summer period in their photosynthetic apparatus through the xanthophyll cycle pool and a related photoprotective mechanism, when the photochemical utilization of photon energy was down-regulated.     

Damages of photosynthetic apparatus inAnacystis nidulans by ultraviolet-B radiation

Changes in photosystem (PS) 2 activity were measured inAnacystis nidulans cells exposed to UV-B irradiation. The decrease in PS 2 activity was biphasic in cells exposed at the surface (0 cm) and at 2 cm depth in the water column, while gradual in those exposed at 10 and 30 cm depth. Addition of supplemental “white light” reduced the extent of UV-B damage. Decrease in photosynthetic activity was primarily due to the loss of energy transfer from phycobilisome to chlorophyll as the former in cyanobacteria acts as the primary light-harvesting complex. This was supported by the absorption and fluorescence excitation and emission spectral studies. All these changes were proportionally reduced by the thickness of the water column that reduced UV-B irradiance. This work was supported in part by a grant from the Ministerio de Education y Ciencia (ref SB 94-AM086772) to N. N.     

Developmental phase-dependent photosynthetic responses to ultraviolet-B radiation: damage, defence, and adaptation of primary leaves of wheat seedlings

Alterations in photosynthetic capacity of primary leaves of wheat seedlings in response to ultraviolet-B (UV-B; 280–320 nm; 60 μmol m⁻² s⁻¹) exposure alone and in combination with photosynthetically active radiation (PAR; 400–800 nm; 200 μmol m⁻² s⁻¹) during different phases of leaf growth and development were assessed. UV-B exposure resulted in a phase-dependent differential loss in photosynthetic pigments, photochemical potential, photosystem 2 (PS2) quantum yield, and in vivo O₂ evolution. UV-B exposure induced maximum damage to the photosynthetic apparatus during senescence phase of development. The damages were partially alleviated when UV-B exposure was accompanied by PAR. UV-B induced an enhancement in accumulation of flavonoids during all phases of development while it caused a decline in anthocyanin content during senescence. The differential changes in these parameters demonstrated the adaptation ability of leaves to UV-B stress during all phases of development and the ability was modified in UV-B+ PAR exposed samples.     

Photosynthesis in leaves of <Emphasis Type="Italic">Nicotiana tabacum</Emphasis> L. infected with tobacco mosaic virus

In tobacco leaves inoculated with tobacco mosaic virus (TMV), changes in chlorophyll (Chl) and carotenoid contents, parameters of slow Chl fluorescence kinetics, i.e. the maximum quantum yield of photosystem (PS2) photochemistry F_v/F_m, the effective quantum yield of photochemical energy conversion in PS2 Φ₂, ratio of quantum yields of photochemical and concurrent non-photochemical processes in PS2 F_v/F₀, non-photochemical quenching (NPQ), and photochemical activities of isolated chloroplasts from systemically infected tobacco leaves were investigated. We compared two successive stages of infection, the first in the stage of vein clearing at 9^th day post inoculation (dpi) and the second at 22^nd dpi when two different regions, i.e. light- (LGI) or dark-green (DGI) islands in the infected leaf were apparent and symptoms were fully developed. These two different regions were measured separately. The Chl and carotenoid contents in infected leaves decreased with a progression of infection and were lowest in LGI in the second stage. Also the ratio of Chl a/b declined in similar manner. The maximum quantum yield of PS2 photochemistry F_v/F_m, was decreased in the following order: first stage, DGI, and LGI. The same is true for the ratio F_v/F₀. The decrease of Φ₂ in infected leaves declined as compared to their controls. On the contrary, NPQ increased in infected leaves, the highest value was found in the first infection stage. Photochemical activities of the whole electron transport chain in isolated chloroplasts dramatically declined with the progression of symptoms, the lowest value was in LGI. Similarly, but to a lesser extent, the activity of PS2 in isolated chloroplasts decreased in infected leaves. Generally, the most marked impairment of the photosynthetic apparatus was manifested in the LGI of infected leaves.     

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     

Effects of Season,Needle Age,and Elevated Atmospheric CO2 on Chlorophyll Fluorescence Parameters and Needle Nitrogen Concentration in Scots Pine (Pinus sylvestris)

Six-year-old Scots pine (Pinus sylvestris L.) seedlings were grown in open top chambers (OTCs) at ambient (AC) or elevated (ambient + 400 µmol mol^–1; EC) CO₂ concentration for three years (1996–1998). Chlorophyll (Chl) a fluorescence of current and one-year-old needles was measured in the field at two-weekly intervals in the period July–October 1998. In addition, Chl, carbon (C), and nitrogen (N) concentrations in both needle age classes were determined monthly during the same period. Chl fluorescence parameters were not significantly affected by EC, suggesting there was no response of the light reactions and the photochemical efficiency of photosystem 2. Chl concentrations were not significantly different but a reduced N concentration was observed in needles of EC treatment. Significant differences between needle age classes were observed for all parameters, but were most apparent under EC and toward the end of the growing season, possibly due to an acclimation process. As a result, significant interactions between CO₂ treatment, needle age class, and season were found. This study emphasizes the importance of repeated measures including different leaf/needle age classes to assess the photosynthetic response of trees under EC.     

Effect of low doses of UV-A and UV-B radiation on photosynthetic activities in<Emphasis Type="Italic">Phaseolus mungo</Emphasis> L.

The effect of low doses of UV-A (320–400 nm) and UV-B (280–320 nm) radiation on photosynthetic activities inPhaseolus mungo L. was investigated under field condition. Supplementation of UV-A enhanced the synthesis of chlorophyll and carotenoids than the UV-B supplemented plants. Significant increase was seen in the concentration of UV-B absorbing compounds of UV-B treated plants. Increase of PS 2 activity in UV-A treated plants was seen. Changes in photosynthetic activity were measured in terms of PS 2 mediated O₂ evolution and Chl a fluorescence.     

Modifications in Photosynthetic Pigments and Chlorophyll Fluorescence in 20-Year-Old Pine Trees after a Four-Year Exposure to Carbon Dioxide and Temperature Elevation

Changes in pigment composition and chlorophyll (Chl) fluorescence parameters were studied in 20 year-old Scots pine (Pinus sylvestris L.) trees grown in environment-controlled chambers and subjected to ambient conditions (CON), doubled ambient CO₂ concentration (EC), elevated temperature (ambient +2−6 °C, ET), or a combination of EC and ET (ECT) for four years. EC did not significantly alter the optimal photochemical efficiency of photosystem 2 (PS2; F_v/F_m), or Chl a+b content during the main growth season (days 150–240) but it reduced F_v/F_m and the Chl a+b content and increased the ratio of total carotenoids to Chl a+b during the ‘off season’. By contrast, ET significantly enhanced the efficiency of PS2 in terms of increases in F_v/F_m and Chl a+b content throughout the year, but with more pronounced enhancement in the ‘off season’. The reduction in F_v/F_m during autumn could be associated with the CO₂-induced earlier yellowing of the leaves, whereas the temperature-stimulated increase in the photochemical efficiency of PS2 during the ‘off season’ could be attributed to the maintenance of a high sink capacity. The pigment and fluorescence responses in the case of ECT showed a similar pattern to that for ET, implying the importance of the temperature factor in future climate changes in the boreal zone. This revised version was published online in August 2006 with corrections to the Cover Date.     

Growth and nitrogen utilization in seedlings of mountain birch (Betula pubescens ssp. tortuosa ) as affected by ultraviolet radiation (UV-A and UV-B) under laboratory and outdoor conditions

 Growth patterns and nitrogen economy were studied in pot-grown seedlings of mountain birch subjected to different ultraviolet radiation under both laboratory and outdoor conditions at Abisko in northern Sweden. In the laboratory, nutrient supply, temperature, humidity, ultraviolet radiation-A (UV-A, 320–400 nm) and B (UV-B, 280–320 nm) were controlled, while photosynthetically active radiation (PAR, 400–700 nm) and photoperiod varied naturally. Under outdoor conditions nutrient supply was controlled, and the irradiation treatments were ambient and above-ambient UV-B using additional fluorescent lamps. Mountain birch nitrogen economy was affected by increased ultraviolet radiation, as reflected by a changed relationship between plant growth and plant nitrogen both in the laboratory and outdoors. In the laboratory enhanced UV-A decreased leaf area per unit plant biomass (leaf area ratio) but increased biomass productivity, both per unit leaf area (leaf area productivity) and per unit leaf nitrogen (leaf nitrogen productivity). Low levels of UV-B affected growth patterns and nitrogen economy in a similar way to enhanced UV-A. High levels of UV-B clearly decreased relative growth rate and nitrogen productivity, as leaf area ratio, leaf area productivity and leaf nitrogen productivity were all decreased. Under outdoor conditions above-ambient levels of UV-B did not alter growth or biomass allocation traits of the seedlings, whilst nitrogen productivity was increased. Mountain birch seedlings originating from different mother trees varied significantly in their responses to different ultraviolet radiation. Received: 10 April 1997 / Accepted: 19 September 1997     

Induction of heat shock-like proteins in Vigna sinensis seedlings growing under ultraviolet-B (280-320 nm) enhanced radiation

The effect of ultraviolet-B (UV-B) enhanced fluorescent radiation on protein profile and protein synthesis has been investigated in Vigna sinensis L. cv. Walp seedlings growing at various temperatures. In seedlings growing at 30°C, UV-B radiation decreased the level of several proteins as seen in Coomassie brilliant blue stained gel. However, fluorography of the same gel indicates induction of three sets of proteins in the range of 70. 53 and 16 k Da. Such induction under UV-B enhanced radiation resembled that found after heat shock treatments. In seedlings at 10 and 20°C, induction of such proteins varied both qualitatively and quantitatively. At 40°C. UV-B enhanced radiation caused a cumulative effect with temperature. Strong induction of specific proteins by UV-B radiation in seedlings growing under normal temperature indicates a possible protective role.