EFFECTS OF UV‐B RADIATION ON THE D1 PROTEIN REPAIR CYCLE OF NATURAL PHYTOPLANKTON COMMUNITIES FROM THREE LATITUDES (CANADA,BRAZIL, AND ARGENTINA)1

Ultraviolet radiation effects were examined in natural phytoplankton communities from Rimouski (Canada), Ubatuba (Brazil), and Ushuaia (Argentina). Outdoor pump‐mixed mesocosms were submitted to ambient solar radiation (NUVB) and ambient with additional UV‐B radiation (UVBR) from lamps (HUVB), corresponding to a local 60% ozone depletion scenario. At all sites, neither algal biomass nor dark‐adapted F_v/F_m were significantly affected by additional UVBR, suggesting the presence of active UV protection or repair mechanisms. To examine the role of D1 protein turnover, essential for PSII repair, short‐term surface incubations were performed in the presence or absence of lincomycin, a chloroplast protein synthesis inhibitor. Effects on PSII were determined using chl a in vivo fluorescence, whereas the D1 protein was detected immunochemically. In the absence of D1 repair, D1 pools and F_v/F_m decreased to a similar extent under both light treatments. In the presence of D1 repair, D1 pools suffered faster net degradation under HUVB compared with NUVB, whereas F_v/F_m was maintained for both light treatments, suggesting that HUVB exposure in field populations had more effect on D1 synthesis and PSII repair than on D1 degradation. The fewer undamaged reaction centers remaining in phytoplankton under HUVB were able to maintain F_v/F_m or actually recovered during the dark acclimation before F_v/F_m measurements. The D1 pools suffered faster net degradation at the tropical site where high irradiance drove faster D1 degradation and high water temperature enabled fast enzymatic activities. This study shows the crucial role of dynamic changes in D1 turnover in the photobiology of natural planktonic communities across a range of latitudes.     

Different photochemical responses of phytoplankters from the large shallow Taihu Lake of subtropical China in relation to light and mixing

The maximum quantum yield of photosystem II was estimated from variable chlorophyll a fluorescence in samples of phytoplankton collected from the Taihu Lake in China to determine the responses of different phytoplankters to irradiance and vertical mixing. Meteorological and environmental variables were also monitored synchronously. The maximum quantum yield of three phytoplankton groups: cyanobacteria, chlorophytes, and diatoms/dinoflagellates, showed a similar diurnal change pattern. F _v/F _m decreased with a significant depth-dependent variation as irradiance increased during the morning and increased as irradiance declined in the afternoon. Furthermore, the rates of F _v/F _m depression were dependent upon the photon flux density, whereas the rates of recovery of F _v/F _m were dependent upon the historical photon density. Moreover, photoinhibition affected the instantaneous growth rates of phytoplankton. Although at noon cyanobacteria had a higher photoinhibition value (up to 41%) than chlorophytes (32%) and diatoms/dinoflagellates (34%) at the surface, no significant difference in diurnal growth rates among the three phytoplankton groups were observed indicating that cyanobacteria could photoacclimate better than chlorophytes and diatoms/dinoflagellates. In addition, cyanobacteria had a higher nonphotochemical quenching value than chlorophytes and diatoms/dinoflagellates at the surface at noon, which indicated that cyanobacteria were better at dissipating excess energy. The ratios of enclosed bottle samples F _v/F _m to free lake samples F _v/F _m showed different responses for the three phytoplankton groups to irradiance and vertical mixing when wind speed was approximately constant at about 3.0 m s⁻¹. When wind speed was lower than 3.0 m s⁻¹, cyanobacteria accumulated mainly at the surface and 0.3 m, because of their positive buoyancy, where diurnal growth rates of phytoplankton were relatively higher than those at 0.6 m and 0.9 m. Chlorophytes were homogenized completely by vertical mixing, while diatoms/dinoflagellates avoided active high irradiance by moving downward at noon, and then upward again when irradiance decreased. These results explain the dominance of cyanobacteria in Taihu Lake. Handling editor: L. Naselli-Flores     

DIFFERENTIAL RESPONSES OF ANABAENA SP. PCC 7120 (CYANOPHYCEAE) CULTURED IN NITROGEN‐DEFICIENT AND NITROGEN‐ENRICHED MEDIA TO ULTRAVIOLET‐B RADIATION1

Stratospheric ozone depletion increases the amount of ultraviolet‐B radiation (UVBR) (280–320 nm) reaching the surface of the earth, potentially affecting phytoplankton. In this work, Anabaena sp. PCC 7120, a typically nitrogen (N)‐fixing filamentous bloom‐forming cyanobacterium in freshwater, was individually cultured in N‐deficient and N‐enriched media for long‐term acclimation before being subjected to ultraviolet‐B (UVB) exposure experiments. Results suggested that the extent of breakage in the filaments induced by UVBR increases with increasing intensity of UVB stress. In general, except for the 0.1 W · m^?2 treatment, which showed a mild increase, UVB exposure inhibits photosynthesis as evidenced by the decrease in the chl fluorescence parameters maximum photochemical efficiency of PSII (F_v/F_m) and maximum relative electron transport rate. Complementary chromatic acclimation was also observed in Anabaena under different intensities of UVB stress. Increased total carbohydrate and soluble protein may provide some protection for the culture against damaging UVB exposure. In addition, N‐deficient cultures with higher recovery capacity showed overcompensatory growth under low UVB (0.1 W · m^?2) exposure during the recovery period. Significantly increased (～830%) ATPase activity may provide enough energy to repair the damage caused by exposure to UVB.     

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.     

SYNTHESIS OF MYCOSPORINE-LIKE AMINO ACIDS IN CHONDRUS CRISPUS (FLORIDEOPHYCEAE) AND THE CONSEQUENCES FOR SENSITIVITY TO ULTRAVIOLET B RADIATION

The induction and protective role of the UV-absorbing compounds known as mycosporine-like amino acids (MAAs) were examined in sublittoral Chondrus crispus Stackh. transplanted for 2 weeks in the spring and summer to shallow water under three irradiance conditions: PAR (photosynthetically active radiation; 400–700 nm), PAR + UVA (PAR + 320– 400 nm), PAR + UVA + UVB (PAR + UVA + 280– 320 nm). Sublittoral thalli collected around Helgoland, North Sea, Germany, from 6 m below the mean low water of spring tides contained less than 0.1 mg·g⁻¹ dry weight (DW) total MAAs, whereas eulittoral samples contained over 1 mg·g⁻¹ DW. Transplantation to shallow water led to the immediate synthesis of three MAAs in the following temporal order: shinorine (λ_max 334 nm), asterina (λ_max 330 nm), and palythine (λ_max 320 nm), with the shinorine content peaking and then declining after 2 days (exposure to 100 mol photons·m⁻²). Maximum total MAA content (2 mg·g⁻¹ DW) also occurred after 2 days of induction, exceeding the content normally found in eulittoral samples. Furthermore, the relative proportion of the different MAAs at this time was different than that in eulittoral samples. After 2 days the total content declined to the eulittoral value, with palythine as the principal MAA. Similar data were obtained for all treatments, indicating that MAA synthesis in C. crispus was induced by PAR and not especially stimulated by UV radiation. The ability of photosystem II (PSII) to resist damage by UVB was tested periodically during the acclimation period by exposing samples to a defined UVB dose in the lab. Changes in chlorophyll fluorescence (F_v/F_m and effective quantum yield, φ_II) indicated that PSII function was inhibited during the initial stage of acclimation but gradually improved with time. No difference among screening treatments was detected except in spring for the samples acclimating to PAR + UVA + UVB. In this treatment F_v/F_m and φ_II were significantly lower than in the other treatments. During the first week of each experiment, growth rates were also significantly reduced by UVB. The reductions occurred despite maximum MAA content, indicating an incomplete protection of photosynthetic and growth-related processes.     

Underestimate of PS2 efficiency in the field due to high leaf temperature resulting from leaf clipping and its amendment

Chlorophyll fluorescence parameter F_v/F_m, an indicator of the maximum efficiency of PS2, is routinely measured in the field with plant leaves darkened by leaf clips. I found that on a sunny day of subtropical summer, the F_v/F_m ratio was often underestimated because of a large F₀ value resulted from a high leaf temperature caused by clipping the leaf under high irradiance, especially for long (e.g. 20 min) duration. This phenomenon may overestimate the down-regulation of PS2 efficiency under high irradiance. When leaf temperature was lower than 40 °C, the F₀ level of rice leaves under clipping remained practically unchanged. However, F₀ increased drastically with leaf temperature rising over 40 °C. In most measurements, no significant difference in F_m was found between rice leaves dark-adapted by leaf clips for 10 min and for 20 min. Therefore, shading leaf clips to prevent a drastic increase of leaf temperature, using F₀ measured immediately after the leaf being darkened to calculate F_v/F_m, as well as shortening the duration of leaf clipping are useful means to avoid an underestimate of F_v/F_m.     

An inter-basin comparison of nutrient limitation and the irradiance response of pulse-amplitude modulated (PAM) fluorescence in Lake Erie phytoplankton

This research examined the application of the maximum quantum efficiency (F _v/F _m) and relative electron transport rate versus irradiance curves (rETR) as a rapid, sensitive assessment of Lake Erie phytoplankton nutrient status. I evaluated the potential benefits of the variable fluorescence parameters by comparing these parameters with chemical and physiological nutrient status assays. I tested the hypothesis that F _v/F _m and rETR curves could diagnose nutrient status in natural lake phytoplankton and be capable of discriminating which inorganic nutrient is limited temporally and spatially. F _v/F _m was on average highest in the more eutrophic west basin (WB) and lowest in the more oligotrophic central basin (CB). According to the chemical and physiological indicators, P deficiency was most severe in the CB during summer stratification and N deficiency was strongest in the WB during isothermal conditions. Like F _v/F _m, rETR at light saturation (rETR_max) and the initial slope of the rETR versus irradiance curve (α) decreased as the severity of N and P deficiency increased. Amendment with N or P stimulated increased F _v/F _m, rETR_max, and α in N- and P-limited samples, respectively, and abolished the photoinhibition apparent in rETR curves of nutrient-limited samples. These results supported the view that the N and P deficiency assays, and corresponding variations of variable fluorescence parameters, were valid indicators of widely variable N and P deficiency in the phytoplankton, and could be used to provide a promising tool in determining phytoplankton nutrient status. Contrary to my hopes, it did not appear that rETR–irradiance curves could discriminate between N and P deficiency. Identification of the most limiting nutrient still demanded additional information beyond the variable fluorescence measurements.     

Photoinhibition of colonial and unicellular <Emphasis Type="Italic">Microcystis</Emphasis> cells in a summer bloom in Lake Taihu

To evaluate the photoinhibition of colonial and unicellular cells of Microcystis aeruginosa under natural conditions, the maximum and effective quantum yields of photosystem II were measured from variable chlorophyll a fluorescence in samples from Lake Taihu during a summer bloom from June 19 to 21, 2006. Diurnal changes in the photoinhibition of Microcystis cells incubated immediately below the surface in clear bottles for 30 min and in situ samples under natural conditions were measured. At solar noon during the three days, the mean values of maximum quantum yield (F _v/F _m) and effective quantum yield (ΔF/F _m′) for unicellular cells (F _v/F _m = 0.15, ΔF/F _m′ = 0.10) were lower than those for colonial cells (F _v/F _m = 0.25, ΔF/F _m′ = 0.15). For in situ samples, the values of F _v/F _m and ΔF/F _m′ for colonial cells at solar noon on the three days (F _v/F _m 0.30, 0.25, 0.29; ΔF/F _m′ 0.24, 0.21, 0.22) were also higher than those of unicellular cells (F _v/F _m 0.26, 0.18, 0.25; ΔF/F _m′ 0.15, 0.11, 0.14). The results indicate that colony formation has a protective effect on Microcystis cells by reducing the occurrence of photoinhibition under high light intensities.     

Multi‐wavelength Pulse Amplitude Modulated fluorometry (Phyto‐PAM) reveals differential effects of ultraviolet radiation on the photosynthetic physiology of phytoplankton pigment groups

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Physiological state of phytoplankton communities in the Southwest Atlantic sector of the Southern Ocean, as measured by fast repetition rate fluorometry

The majority of the Southern Ocean is a high-nutrient low-chlorophyll (HNLC) ecosystem. Localized increases in chlorophyll concentration measured in the wake of bathymetric features near South Georgia demonstrate variations in the factors governing the HNLC condition. We explore the possibility that the contrast between these areas of high-chlorophyll and surrounding HNLC areas is associated with variations in phytoplankton photophysiology. Total dissolvable iron concentrations, phytoplankton photophysiology and community structure were investigated in late April 2003 on a transect along the North Scotia Ridge (53–54°S) between the Falkland Islands and South Georgia (58–33°W). Total dissolvable iron concentrations suggested a benthic source of iron near South Georgia. Bulk community measurements of dark-adapted photochemical quantum efficiency (F _v/F _m) exhibited a sharp increase to the east of 46°W coincident with a decrease in the functional absorption cross-section (σ_PSII). Phytoplankton populations east of 46°W thus displayed no physiological symptoms of iron or nitrate stress. Contrasting low F _v/F _m west of 46°W could not be explained by variations in the macronutrients nitrate and silicic acid and may be the result of taxon specific variability in photophysiology or iron stress. We hypothesize that increased F _v/F _m resulted from local relief from iron-stress near South Georgia, east of Aurora Bank, an area previously speculated to be a “pulse point” source of iron. Our measurements provide one of the first direct physiological confirmations that iron stress is alleviated in phytoplankton populations near South Georgia.     

FLUORESCENCE-BASED MAXIMAL QUANTUM YIELD FOR PSII AS A DIAGNOSTIC OF NUTRIENT STRESS

In biological oceanography, it has been widely accepted that the maximum quantum yield of photosynthesis is influenced by nutrient stress. A closely related parameter, the maximum quantum yield for stable charge separation of PSII, (φ _PSII )_m, can be estimated by measuring the increase in fluorescence yield from dark-adapted minimal fluorescence (F_o) to maximal fluorescence (F_m) associated with the closing of photosynthetic reaction centers with saturating light or with a photosynthetic inhibitor such as 3′-(3,4-dichlorophenyl)-1′,1′-dimethyl urea (DCMU). The ratio F_v/F_m (= (F_m− F_o)/F_m) is thus used as a diagnostic of nutrient stress. Published results indicate that F_v/F_m is depressed for nutrient-stressed phytoplankton, both during nutrient starvation (unbalanced growth) and acclimated nutrient limitation (steady-state or balanced growth). In contrast to published results, fluorescence measurements from our laboratory indicate that F_v/F_m is high and insensitive to nutrient limitation for cultures in steady state under a wide range of relative growth rates and irradiance levels. This discrepancy between results could be attributed to differences in measurement systems or to differences in growth conditions. To resolve the uncertainty about F_v/F_m as a diagnostic of nutrient stress, we grew the neritic diatom Thalassiosira pseudonana (Hustedt) Hasle et Heimdal under nutrient-replete and nutrient-stressed conditions, using replicate semicontinuous, batch, and continuous cultures. F_v/F_m was determined using a conventional fluorometer and DCMU and with a pulse amplitude modulated (PAM) fluorometer. Reduction of excitation irradiance in the conventional fluorometer eliminated overestimation of F_o in the DCMU methodology for cultures grown at lower light levels, and for a large range of growth conditions there was a strong correlation between the measurements of F_v/F_m with DCMU and PAM (r² = 0.77, n = 460). Consistent with the literature, nutrient-replete cultures showed consistently high F_v/F_m (∼0.65), independent of growth irradiance. Under nutrient-starved (batch culture and perturbed steady state) conditions, F_v/F_m was significantly correlated to time without the limiting nutrient and to nutrient-limited growth rate before starvation. In contrast to published results, our continuous culture experiments showed that F_v/F_m was not a good measure of nutrient limitation under balanced growth conditions and remained constant (∼0.65) and independent of nutrient-limited growth rate under different irradiance levels. Because variable fluorescence can only be used as a diagnostic for nutrient-starved unbalanced growth conditions, a robust measure of nutrient stressed oceanic waters is still required.     

Recovery from winter depression of photosynthesis in pine and spruce

Summary Recovery from winter depression of photosynthesis was studied in Pinus sylvestris, Pinus conforta and Picea abies by means of chlorophyll fluorescence and gas exchange measurements. During the winter 1986–1987 the fluorescence yield was low and no variable fluorescence was detectable before the end of March. In the field recovery of variable fluorescence/maximum fluorescence (F_v/F_m) during spring was slow for all three species studied. The temperature dependence of recovery was confirmed from measurements of the potential rate of recovery of F_v/F_m at different temperatures in the laboratory. At 20° C, F_v/F_m increased from 0.1 to 0.8 within 3 days. Recovery of F_v/F_m was paralleled by an increase in apparent photon yield. No significant differences could be demonstrated between the studied tree species in potential rate of recovery in the laboratory or in actual recovery in the field.     

Responses of estuarine intertidal microphytobenthic algal assemblages to enhanced ultraviolet B radiation

As a result of ozone depletion, ground doses of ultraviolet B (UVB) radiation in the mid latitudes of the Northern Hemisphere have increased since the 1980s, and current predictions indicate no possible alleviation until at least post 2020. Mudflats and sandflats are important coastal-zone habitats, and support extensive biofilms of benthic microalgae (microphytobenthos). In intertidal situations, these assemblages are exposed to high levels of UVB radiation during periods of tidal exposure. Exposure of intertidal biofilms dominated by epipelic (mud-inhabiting) diatoms to 0, 0.18 or 0.35 W m⁻² UVB radiation for between 4 and 10 days resulted in no significant decreases in the maximum PSII quantum efficiency (F_v/F_m) throughout diel exposure periods. Although the quantum efficiency of electron transport (F_q′/F_m′) showed significant reductions early in some experiments, the major response was an increase in F_q′/F_m′ in UVB exposed biofilms. This increase in F_q′/F_m′ was suggestive of a protective vertical migration down into the sediment. Single-cell and whole biofilm fluorescence imaging demonstrated, for the first time, that motile diatoms are able to detect UVB radiation independently of UVA or photosynthetically active radiation (PAR) and migrate rapidly down (within 15 min) into the sediments to avoid it. This behavioural acclimation mechanism appears to prevent significant accumulation of UVB induced damage to the algae. UVB exposure had no significant effect of biofilm photosynthesis (measured by ¹⁴C carbon fixation), but did alter organic carbon allocation patterns, with significantly less new carbon allocated to intracellular storage (glucan) and extracellular colloidal carbohydrate fractions. Significant reductions in the sediment standing stocks of chlorophyll a (Chl a), colloidal carbohydrates, extracellular polymeric substances (EPS) were seen after 7 days of UVB exposure. This study showed that marine intertidal benthic diatoms use a behavioural strategy to avoid exposure to UVB and that this response is effective as a short-term protection mechanism against UVB damage. However, altered carbon allocation patterns feed forward over time into changes in biofilm biomass and sediment carbohydrate dynamics. This suggests that continual long-term exposure to UVB may impact on sediment carbon cycling and trophic interactions and on the stabilization of sediments by microalgal biofilms through their production of extracellular carbohydrates.     

Evaluating the Superoxide Dismutase Activity and Chlorophyll Fluorescence in Picea Abies Leaves Growing at Different Altitudes

Superoxide dismutase (SOD) activity and parameters of chlorophyll fluorescence, the ratio of maximal to variable fluorescence (F_v/F_m), maximal fluorescence (F_m), and minimal fluorescence (F₀) were determined on Picea abies growing at different altitudes. The decreases of F_v/F_m and F_m, in comparison to samples from the lower stands (control), were found on trees from the highest stands. The decrease of fluorescence parameters was reversible, at least partly, after keeping branches for some days in the laboratory. F_v/F_m measured in spring when trees were partially covered with snow revealed greater degree of photoinactivation in branches collected from above the snow in comparison to those from below the snow. In samples collected from above snow also slower recovery from stress was observed. Two main SOD isoforms were determined in needles of P. abies, and classified as CuZnSODs. The activity of both SOD isoforms was increasing with the altitude, thus indicating the highest level of oxidative stress at the timberline zone. This revised version was published online in August 2006 with corrections to the Cover Date.     

Solar UV-B radiation, associated with ozone depletion, inhibits the Antarctic terrestrial microalga, Stichococcus bacillaris

This study shows that increased UV-B arising from stratospheric ozone depletion over Antarctica reduced cell viability and the maximum quantum yield of photochemistry (F _v/F _m) in a unicellular terrestrial microalga, Stichococcus bacillaris. In the UV waveband, rates of F _v/F _m decline increased with decreasing wavelength. Photosynthetically active radiation (PAR) also reduced F _v/F _m, though less than UV radiation. Further experiments under different ozone column thicknesses showed a significantly greater decline in cell viability and F _v/F _m under ozone depletion compared with non-depleted conditions. The inhibitory effects of ambient solar radiation suggest that S. bacillaris is unlikely to inhabit soil surfaces, but colonises shaded areas beneath soil surface particles. During periods of ozone depletion, increases in the ratio of UV-B:PAR may reduce the thickness of the sub-surface zone where light conditions are suitable for colonisation by this alga.     

GROWTH AND PHOTOPROTECTION IN THREE DINOFLAGELLATES (INCLUDING TWO STRAINS OF ALEXANDRIUM TAMARENSE) AND ONE DIATOM EXPOSED TO FOUR WEEKS OF NATURAL AND ENHANCED ULTRAVIOLET‐B RADIATION1

Long‐term growth response to natural solar radiation with enhanced ultraviolet‐B (UVB) exposure was examined in two species of dinoflagellates [Alexandrium tamarense (M. Lebour) Balech, At, and Heterocapsa triquetra (Ehrenb.) F. Stein, Ht], including two strains of A. tamarense, one from Spain and another from UK, and one diatom species (Thalassiosira pseudonana Hasle et Heimdal). We examined whether variable photoprotection (mycosporine‐like amino acids [MAAs] and xanthophyll‐cycle pigments) affected photosynthetic performance, phytoplankton light absorption, and growth. Growth rate was significantly reduced under enhanced UVB for the UK strain of At and for Ht (both grew very little) as well as for the diatom (that maintained high growth rates), but there was no effect for the Spanish strain of At. MAA concentration was high in the dinoflagellates, but undetectable in the diatom, which instead used the xanthophyll cycle for photoprotection. The highest cell concentrations of MAAs and photoprotective pigments were observed in the UK strain of At, along with lowest growth rates and F_v/F_m, indicating high stress levels. In contrast, the Spanish strain showed progressive acclimation to the experimental conditions, with no significant difference in growth between treatments. Increase in total MAAs followed linearly the cumulative UVB of the preceding day, and both total and primary MAAs were maintained at higher constitutive levels in this strain. Acclimation to enhanced UVB in the diatom resulted in an increase in PSII activity and reduction in nonphotochemical quenching, indicating an increased resistance to photoinhibition after a few weeks. All four species showed increased phytoplankton light absorption under enhanced UVB. Large intrastrain differences suggest a need to consider more closely intraspecific variability in UV studies.     

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     

Physiological responses of phytoplankton communities in the Irish Sea to simulated upwelling

Understanding the dynamics of upwelling systems, especially the interactions between nutrients and light, has benefited from the application of models of varying complexity. Validation of such models using unialgal cultures or field observations has often proven difficult, but short-term incubations of contained natural assemblages and use of instantaneous physiological indicators offer an alternative approach. In May and June 1996, phytoplankton communities deep in the euphotic zone were sampled from nearly identical physical environments. Replicate samples (20 l volume) were incubated on deck at 50% surface irradiance with either no nutrient additions (Controls) or additions of 20 μM nitrate (Enrichments). Over 24 h, variable fluorescence (F _v:F _m), nitrate reductase activity (NR), nutrients, chlorophyll a and particulate C and N were monitored. Initial chlorophyll a (～3 μg l^?1), phosphate (～0.2 μM), nitrate (～1.5 μM) and silicate (～3 μM) were similar in both months. Changes in NR and F _v:F _m indicated clear physiological responses to changes in irradiance and added nitrate that differed between months. In May, Controls and Enrichments responded in the same way. F _v:F _m stayed constant (0.5), chlorophyll a increased slightly, and NR activity increased markedly in all samples. In contrast, in June, treatments responded quite differently. F _v:F _m was near the theoretical maximum (0.7–0.8) initially and remained constant in Enrichments, but fell sharply in Controls. Declines in controls were also seen for chlorophyll a, and NR activity. Thus, the addition of 20 μM nitrate had a significant effect even though ambient levels of nitrate (>1 μM) should not have been limiting. Small (<20 μm) flagellates predominated in the May samples, but in June large and chain-forming centric diatoms constituted a significant proportion of the phytoplankton community. We conclude that the response of a phytoplankton community to environmental changes can depend on factors that are poorly represented by bulk measurements of chlorophyll, nutrients and particulate elements.     

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.     

Ambient levels of UV-B in Hawaii combined with nutrient deficiency decrease photosynthesis in near-isogenic maize lines varying in leaf flavonoids: Flavonoids decrease photoinhibition in plants exposed to UV-B

Near-isogenic lines of maize varying in their genes for flavonoid biosynthesis were utilized to examine the effects of foliar flavonoids and nutrient deficiency on maximum net photosynthetic rate (P _N) and chlorophyll (Chl) fluorescence (F_v/F_m) in response to ultraviolet-B (UV-B) radiation. Plants with deficient (30 to 70 % lower N, K, Mn, Fe, and Zn) and sufficient nutrients were exposed to four irradiation regimes: (1) no UV-B with solar photosynthetically active radiation (PAR), (2) two day shift to ambient artificial UV-B, 8.2–9.5 kJ m⁻² d⁻¹ (21–25 mmol m⁻² d⁻¹); (3) continuous ambient artificial UV-B; (4) continuous solar UV-B in Hawaii 12–18 kJ m⁻² d⁻¹ (32–47 mmol m⁻² d⁻¹). The natural ratio of UVB: PAR (0.25–0.40) was maintained in the UV-B treatments. In the adequately fertilized plants, lines b and lc had higher contents of flavonoids and anthocyanins than did lines hi27 and dta. UV-B induced the accumulation of foliar flavonoids in lines hi27 and b, but not in the low flavonoid line dta or in the high flavonoid line lc. In plants grown on deficient relative to adequate nutrients, flavonoid and anthocyanin contents decreased by 30–40 and 40–50 %, respectively, and Chl a and Chl b contents decreased by 30 and 70 %, respectively. The UV-B treatments did not significantly affect P _N and F_v/F_m in plants grown on sufficient nutrients, except in the low flavonoid lines dta and hi27 in which P _N and F_v/F_m decreased by ∼15 %. P _N, F_v/F_m, and stomatal conductance decreased markedly (20–30 %) in all lines exposed to UV-B when grown on low nutrients. The decrease in F_v/F_m was 10 % less in higher flavonoid lines b and lc. The photosynthetic apparatus of maize readily tolerated ambient UV-B in the tropics when plants were adequately fertilized. In contrast, ambient UV-B combined with nutrient deficiency significantly reduced photosynthesis in this C₄ plant. Nutrient deficiency increased the susceptibility of maize to UV-B-induced photoinhibition in part by decreasing the contents of photoprotective compounds.