ANALYSIS OF PORPHYRA RBC L DEMONSTRATES MULTIPLE MIGRATIONS OCCURRED BETWEEN THE NORTH ATLANTIC AND NORTH PACIFIC.

Ground level ultraviolet-B (UV-B; 290–320 nm) fluxes in Antarctica have been increasing due to stratospheric ozone depletion. Although mat-forming cyanobacteria are major component of freshwater algal biomass in Antarctica, little is known about their response to increasing ultraviolet radiation (UVR). The present study evaluated the sensitivity to UVR of two strains of mat-forming cyanobacteria with different cell size, Phormidium murrayi (6.0 x 3.2 μm) and Schizothrix calcicola (2.2 x 2.3 μm). Cyanobacterial photosynthesis was measured under different UV spectral quality and quantity achieved by polychromatic filters with different cutoff wavelengths and neutral density screens. The productivity and irradiance data were used to generate biological weighting functions (BWF) for the assessment of UV inhibition on photosynthesis. The kinetics of UV inhibition, as determined by PAM fluorometry, differed between the two species so that inhibition of P. murrayi and S. calcicola were modeled based on UV-irradiance and cumulative exposure, respectively. After a one hour exposure, BWF's did not differ between the two isolates of cyanobacteria despite their differences in cell size. To evaluate the negative impact of increased UV-B exposure due to ozone depletion on cyanobacteria, the BWF's were applied to two solar spectra obtained from McMurdo Station, one on a day when the ozone hole was prominent (O₃ = 170 Dobson units; DU = 10-3 cm O₃), and the other on a day with high ozone concentration (O₃ = 328 DU). The decrease in ozone level would reduce productivity by 3–8%. Seasonal variation of UVR has a bigger impact on cyanobacterial productivity than ozone depletion.     

TEMPERATURE DEPENDENCE OF UV RADIATION EFFECTS ON ANTARCTIC CYANOBACTERIA

The mat-forming cyanobacterium Phormidium murrayi West and West isolated from a meltwater pond on the McMurdo Ice Shelf was grown in unialgal batch cultures to evaluate the temperature dependence of ultraviolet radiation (UVR) effects on pigment composition, growth rate, and photosynthetic characteristics. Chlorophyll a concentrations per unit biomass were generally reduced in cells grown under UVR (low UV-A plus UV-B). In vivo absorbance spectra showed that the carotenoid/chlorophyll a ratio increased as a function of photosynthetically available radiation (PAR) and UVR exposure and varied inversely with temperature. Ultraviolet inhibition of growth (percentage reduction of μ_max at each temperature) increased linearly with decreasing temperature, consistent with the hypothesis that net inhibition represents the balance between temperature-independent photochemical damage and temperature-dependent biosynthetic repair. There was no significant effect of UVR on photosynthesis over the first hour of exposure, but significant UV inhibition was observed after 5 days. Unlike growth, however, there was no apparent effect of temperature on the magnitude of UV inhibition of photosynthesis. These results imply that assays of UVR effects on photosynthesis are not an accurate guide to growth responses and that low ambient temperatures can have a major influence on the UV sensitivity of polar organisms. In a set of assays at 20° C (preacclimation under 300 μmol photons·m^?2·s^?1 and 20° C), growth was strongly depressed by UVR over the first day of exposure but then gradually increased over the subsequent 4 days, approaching the growth rates in the minus UVR control. This evidence of acquired tolerance indicates that the damaging effects of UVR will be most severe in environments where there is a mismatch between the timescale of change in exposure and the timescale of UV acclimation.     

EFFECTS OF UV RADIATION ON A CHLOROPHYTE ALGA (SCENEDESMUS SP.) ISOLATED FROM THE FUMAROLE FIELDS OF MT. EREBUS,ANTARCTICA1

Acclimation to UV radiation (UVR) was examined in a unicellular chlorophyte isolated from fumarole fields adjacent to Mt. Erebus, Antarctica. Long‐term exposure to UVR (14 days) initially reduced the efficiency of photosynthetic energy conversion measured as the dark adapted quantum yield of PSII fluorescence (F_v/F_m) when compared with cultures not exposed to UVR. However, the UVR exposed cultures recovered to pre‐UVR exposure levels of photosynthetic efficiency by the end of the experimental period. Additionally, neither growth rates nor cell size were significantly affected by exposure to UVR. In contrast, the maximum rate of carbon fixation on a chl a basis was significantly reduced at the end of the experiment and was accompanied by a significant decrease in chl a concentrations. These results suggest a transient effect of UVR on primary photochemistry consistent with damage to PSII, possibly at the D1 protein, with more chronic effects on carbon fixation that did affect maximum photosynthetic capacity. Ultrastructural and molecular (18s rDNA) data show that this isolate from the fumoral fields of Mt. Erebus, Antarctica is a member of the genus Scenedesmus.     

UV‐radiation and elevated temperatures induce formation of reactive oxygen species in gametophytes of cold‐temperate/Arctic kelps (Laminariales,Phaeophyceae)

Enhanced UV‐radiation (UVR) through stratospheric ozone depletion and global warming are crucial stressors to marine macroalgae. Damages may arise through formation of reactive oxygen species (ROS) in gametophytes of ecologically important kelps, brown algae of the order Laminariales, Such stress‐induced damages may have a negative impact on their fitness and further impact their following life stages. In our study, gametophytes of three kelp species Alaria esculenta (L.) Grev., Laminaria digitata (Huds.) Lamour., Saccharina latissima (L.) Lane, Mayes, Druehl, Saunders from the Arctic, and of L. hyperborea (Gunnerus) Foslie from the North Sea were exposed to photosynthetically active radiation, UV‐A, and UV‐B radiation and four temperatures (2–18°C). ROS are formed predominantly in the peripheral cytoplasm and in chloroplasts especially after exposure to UVR. Superoxide (O₂*^‐) is additionally formed in small, globular cytoplasmic structures, possibly mitochondria. In the surrounding medium O₂*^‐‐concentration increased markedly at elevated temperatures and under UV stress in some cases. Ultrastructural damage was negligible pointing to a high stress tolerance of this developmental stage. Our data indicate that stress tolerant gametophytes of three Arctic kelp species should sustain their crucial function as seed bank for kelp populations even under prospective rising environmental perturbations.     

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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Photosynthetic performance,DNA damage and repair in gametes of the endemic Antarctic brown alga Ascoseira mirabilis exposed to ultraviolet radiation

Abstract Stress physiology on the reproductive cells of Antarctic macroalgae remained unstudied. Ascoseira mirabilis is endemic to the Antarctic region, an isolated ecosystem exposed to extreme environmental conditions. Moreover, stratospheric ozone depletion leads to increasing ultraviolet radiation (280–400 nm) at the earth's surface, thus it is necessary to investigate the capacity of reproductive cells to cope with different UV irradiances. This study is aimed to investigate the impact of exposure to different spectral irradiance on the photosynthetic performance, DNA damage and gamete morphology of the A. mirabilis. Gametangia, gametes and zygotes of the upper sublittoral brown alga A. mirabilis were exposed to photosynthetically active radiation (PAR = P; 400–700 nm), P + UV‐A radiation (UV‐A, 320–400 nm) and P + UV‐A + UV‐B radiation (UV‐B, 280–320 nm). Rapid photosynthesis versus irradiance curves of freshly released propagules were measured. Photosynthetic efficiencies and DNA damage (in terms of cyclobutane pyrimidine dimers) were determined after 1, 2, 4 and 8 h exposure as well as after 2 days of recovery in dim white light. Saturation irradiance (I_k) in freshly released propagules was 52 μmol photons m⁻² s⁻¹. Exposure for 1 h under 22 μmol photons m⁻² s⁻¹ of PAR significantly reduced the optimum quantum yield (F_v/F_m), suggesting that propagules are low light adapted. Furthermore, UVR significantly contributed to the photoinhibition of photosynthesis. Increasing dose as a function of exposure time additionally exacerbated the effects of different light treatments. The amount of DNA damage increased with the UV‐B dose but an efficient repair mechanism was observed in gametes pre‐exposed to a dose lower than 5.8 × 10³ J m⁻² of UV‐B. The results of this study demonstrate the negative impact of UV‐B radiation. However, gametes of A. mirabilis are capable of photosynthetic recovery and DNA repair when the stress factor is removed. This capacity was observed to be dependent on the fitness of the parental sporophyte.     

Response of two Antarctic bryophytes to stratospheric ozone depletion

We report a study which measured changes to the radiative environment arising from stratospheric O₃ depletion at Rothera Point on the western Antarctic Peninsula (67°S, 68°W) and subsequent associations between these changes and the pigmentation and maximum quantum yield of photochemistry (F_v/F_m) of two Antarctic bryophytes, the liverwort Cephaloziella varians and the moss Sanionia uncinata. We found a strong relationship between O₃ column depth and the ratio of UV‐B to PAR irradiance (F_UV‐B/F_PAR) recorded at ground level. Weaker, but significant, associations were also found between O₃ column depth and noon irradiances and daily doses of unweighted and biologically effective UV‐B radiation received at ground level. Regression analyses indicated that F_UV‐B/F_PAR and daily dose of unweighted UV‐B were best predictors for concentrations of total carotenoids and UV‐B screening pigments extracted from bryophyte tissues. Concentrations of these pigments were loosely but significantly positively associated with O₃‐dependent irradiance parameters. HPLC analyses of carotenoids also suggested that both species increased the synthesis of neoxanthin during periods of O₃ depletion. Violaxanthin, lutein, zeaxanthin and b,bββ‐carotene concentrations were also apparently influenced by O₃ reduction, but not consistently across both bryophyte species. Concentrations of chlorophylls a and b were apparently unaffected by O₃ depletion. No direct associations between F_v/F_m and O₃‐dependent irradiance parameters were found. However stepwise multiple regression analyses suggested that the production of UV‐B screening pigments conferred protection from elevated F_UV‐B/F_PAR on F_v/F_m in both species and that carotenoids conferred protection on F_v/F_m in Sanionia. Our data suggest that changes to the radiative environment associated with stratospheric O₃ depletion influence the pigmentation of two Antarctic bryophytes, but that F_v/F_m is unaffected, at least in part because of rapid synthesis of protective pigments.     

Effects of Solar UV Radiation on Morphology and Photosynthesis of Filamentous Cyanobacterium Arthrospira platensis

To study the impact of solar UV radiation (UVR) (280 to 400 nm) on the filamentous cyanobacterium Arthrospira (Spirulina) platensis, we examined the morphological changes and photosynthetic performance using an indoor-grown strain (which had not been exposed to sunlight for decades) and an outdoor-grown strain (which had been grown under sunlight for decades) while they were cultured with three solar radiation treatments: PAB (photosynthetically active radiation [PAR] plus UVR; 280 to 700 nm), PA (PAR plus UV-A; 320 to 700 nm), and P (PAR only; 400 to 700 nm). Solar UVR broke the spiral filaments of A. platensis exposed to full solar radiation in short-term low-cell-density cultures. This breakage was observed after 2 h for the indoor strain but after 4 to 6 h for the outdoor strain. Filament breakage also occurred in the cultures exposed to PAR alone; however, the extent of breakage was less than that observed for filaments exposed to full solar radiation. The spiral filaments broke and compressed when high-cell-density cultures were exposed to full solar radiation during long-term experiments. When UV-B was screened off, the filaments initially broke, but they elongated and became loosely arranged later (i.e., there were fewer spirals per unit of filament length). When UVR was filtered out, the spiral structure hardly broke or became looser. Photosynthetic O₂ evolution in the presence of UVR was significantly suppressed in the indoor strain compared to the outdoor strain. UVR-induced inhibition increased with exposure time, and it was significantly lower in the outdoor strain. The concentration of UV-absorbing compounds was low in both strains, and there was no significant change in the amount regardless of the radiation treatment, suggesting that these compounds were not effectively used as protection against solar UVR. Self-shading, on the other hand, produced by compression of the spirals over adaptive time scales, seems to play an important role in protecting this species against deleterious UVR. Our findings suggest that the increase in UV-B irradiance due to ozone depletion not only might affect photosynthesis but also might alter the morphological development of filamentous cyanobacteria during acclimation or over adaptive time scales.     

UV‐A/BLUE LIGHT–INDUCED REACTIVATION OF SPORE GERMINATION IN UV‐B IRRADIATED ULVA PERTUSA (CHLOROPHYTA)1

Recent reduction in the ozone shield due to manufactured chlorofluorocarbons raised considerable interest in the ecological and physiological consequences of UV‐B radiation (λ=280–315 nm) in macroalgae. However, early life stages of macroalgae have received little attention in regard to their UV‐B sensitivity and UV‐B defensive mechanisms. Germination of UV‐B irradiated spores of the intertidal green alga Ulva pertusa Kjellman was significantly lower than in unexposed controls, and the degree of reduction correlated with the UV doses. After exposure to moderate levels of UV‐B irradiation, subsequent exposure to visible light caused differential germination in an irradiance‐ and wavelength‐dependent manner. Significantly higher germination was found at higher photon irradiances and in blue light compared with white and red light. The action spectrum for photoreactivation of germination in UV‐B irradiated U. pertusa spores shows a major peak at 435 nm with a smaller but significant peak at 385 nm. When exposed to December sunlight, the germination percentage of U. pertusa spores exposed to 1 h of solar radiation reached 100% regardless of the irradiation treatment conditions. After a 2‐h exposure to sunlight, however, there was complete inhibition of germination in PAR+UV‐A+UV‐B in contrast to 100% germination in PAR or PAR+UV‐A. In addition to mat‐forming characteristics that would act as a selective UV‐B filter for settled spores under the parental canopy, light‐driven repair of germination after UV‐B exposure could explain successful continuation of U. pertusa spore germination in intertidal settings possibly affected by intense solar UV‐B radiation.     

产毒与不产毒铜绿微囊藻对模拟酸雨及紫外辐射的生理响应

为了比较研究酸雨与紫外辐射对淡水水体常见藻华蓝藻的生理学影响,选取铜绿微囊藻(Microcystis aeruginosa)产毒(FACHB-905)与不产毒(FACHB-469)株系作为实验材料,通过人工模拟酸雨,研究了不同p H处理后2藻株的光合生理变化以及对紫外辐射的敏感性的异同。实验设置3个p H梯度,p H7.10为对照组(正常培养基培养的藻体),两模拟酸雨处理组(p H5.65和p H4.50);两种辐射处理,可见光处理(PAR)以及全波长辐射处理(PAB)。研究结果表明,905藻株细胞粒径在各p H处理下都要显著高于469藻株,模拟酸雨处理显著降低了两藻株细胞的平均粒径及体积,但叶绿素含量显著提高;酸雨处理同时也引起细胞死亡率的增加,表现为藻体有效光化学效率显著降低,生长速率显著受到抑制,低p H下呈负增长,且这种抑制程度在469下更为显著。高的可见光以及紫外辐射处理,使两株系有效光化学效率随p H的降低而呈降低趋势,其中469藻株降低至更低的水平,且高光辐射以及紫外诱导的抑制率要显著高于905藻体,这可能与469藻株较低的光保护色素有关(较低的类胡萝卜素以及紫外吸收物质)。在未来全球变化背景下,不同种类的浮游植物对环境变化的响应及适应能力不同,可改变水体的群落结构和种群丰度,铜绿微囊藻905较469较强的耐受酸雨以及紫外辐射的能力,可能会使该株系在竞争力上占据优势。     

BIOLOGICAL WEIGHTING FUNCTIONS FOR UV INHIBITION OF PHOTOSYNTHESIS IN THE KELP LAMINARIA HYPERBOREA (PHAEOPHYCEAE)1

Different wavelengths of sunlight either drive or inhibit macroalgal production. Ultraviolet radiation (UVR) effectively disrupts photosynthesis, but since UVR is rapidly absorbed in coastal waters, macroalgal photoinhibition and tolerance to UVR depend on the depth of attachment and acclimation state of the individual. The inhibition response to UVR is quantified with a biological weighting function (BWF), a spectrum of empirically derived weights that link irradiance at a specific wavelength to overall biological effect. We determined BWFs for shallow (0 m, mean low water [MLW]) and deep (10 m) Laminaria hyperborea (Gunnerus) Foslie collected off the island of Finnøy, Norway. For each replicate sporophyte, we concurrently measured both O₂ evolution and ¹³C uptake in 48 different light treatments, which varied in UV spectral composition and irradiance. The relative shape of the kelp BWF was most similar to that of a land plant, and the absolute spectral weightings and sensitivity were typically less than phytoplankton, particularly in the ultraviolet radiation A (UVA) region. Differences in BWFs between O₂ and ¹³C photosynthesis and between shallow (high light) and deep (low light) kelp were also most significant in the UVA. Because of its greater contribution to total incident irradiance, UVA was more important to daily loss of production in kelp than ultraviolet radiation B (UVB). Photosynthetic quotient (PQ) also decreased with increased UVR stress, and the magnitude of PQ decline was greater in deepwater kelp. Significantly, BWFs assist in the comparison of biological responses to experimental light sources versus in situ sunlight and are critical to quantifying kelp production in a changing irradiance environment.     

Ultraviolet radiation,ozone depletion,and marine photosynthesis

Concerns about stratospheric ozone depletion have stimulated interest in the effects of UVB radiation (280–320 nm) on marine phytoplankton. Research has shown that phytoplankton photosynthesis can be severely inhibited by surface irradiance and that much of the effect is due to UV radiation. Quantitative generalization of these results requires a biological weighting function (BWF) to quantify UV exposure appropriately. Different methods have been employed to infer the general shape of the BWF for photoinhibition in natural phytoplankton, and recently, detailed BWFs have been determined for phytoplankton cultures and natural samples. Results show that although UVB photons are more damaging than UVA (320–400 nm), the greater fluxes of UVA in the ocean cause more UV inhibition. Models can be used to analyze the sensitivity of water column productivity to UVB and ozone depletion. Assumptions about linearity and time-dependence strongly influence the extrapolation of results. Laboratory measurements suggest that UV inhibition can reach a steady-state consistent with a balance between damage and recovery processes, leading to a non-linear relationship between weighted fluence rate and inhibition. More testing for natural phytoplankton is required, however. The relationship between photoinhibition of photosynthesis and decreases in growth rate is poorly understood, so long-term effects of ozone depletion are hard to predict. However, the wide variety of sensitivities between species suggests that some changes in species composition are likely. Predicted effects of ozone depletion on marine photosynthesis cannot be equated to changes in carbon flux between the atmosphere and ocean. Nonetheless, properly designed studies on the effects of UVB can help identify which physiological and ecological processes are most likely to dominate the responses of marine ecosystems to ozone depletion.Abbreviations BWF biological weighting function - BWF/P-I photosynthesis versus photosynthetically available irradiance as influenced by biologically-weighted UV - Chl chlorophyll a - DOM dissolved organic matter - E _PAR irradiance in energy units (PAR) - E _s saturation parameter for PAR in the BWF/P-I model - E _inh ^* biologically-weighted dimensionless fluence rate for photoinhibition of photosynthesis by UV and PAR - biological weighting coefficient - % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGafqyTduMbae% baaaa!37AC!\[\bar \varepsilon \]_PAR biological weighting coefficient for damage to photosynthesis by E _PAR - k() diffuse attenuation coefficient for wavelength - MAAs mycosporine-like amino acids - PAR photosynthetically available radiation - P ^B rate of photosynthesis normalized to Chl - P _s ^B maximum attainable rate of photosynthesis in the absence of photoinhibition - UVA ultraviolet A (320–400 nm) - UVB ultraviolet B (280–320 nm)     

Inhibition by ultraviolet and photosynthetically available radiation lowers model estimates of depth‐integrated picophytoplankton photosynthesis: global predictions for Prochlorococcus and Synechococcus

Phytoplankton photosynthesis is often inhibited by ultraviolet (UV) and intense photosynthetically available radiation (PAR), but the effects on ocean productivity have received little consideration aside from polar areas subject to periodic enhanced UV‐B due to depletion of stratospheric ozone. A more comprehensive assessment is important for understanding the contribution of phytoplankton production to the global carbon budget, present and future. Here, we consider responses in the temperate and tropical mid‐ocean regions typically dominated by picophytoplankton including the prokaryotic lineages, Prochlorococcus and Synechococcus. Spectral models of photosynthetic response for each lineage were constructed using model strains cultured at different growth irradiances and temperatures. In the model, inhibition becomes more severe once exposure exceeds a threshold (E_max) related to repair capacity. Model parameters are presented for Prochlorococcus adding to those previously presented for Synechococcus. The models were applied to estimate midday, water column photosynthesis based on an atmospheric model of spectral radiation, satellite‐derived spectral water transparency and temperature. Based on a global survey of inhibitory exposure severity, a full‐latitude section of the mid‐Pacific and near‐equatorial region of the east Pacific were identified as representative regions for prediction of responses over the entire water column. Comparing predictions integrated over the water column including versus excluding inhibition, production was 7–28% lower due to inhibition depending on strain and site conditions. Inhibition was consistently greater for Prochlorococcus compared to two strains of Synechococcus. Considering only the surface mixed layer, production was inhibited 7–73%. On average, including inhibition lowered estimates of midday productivity around 20% for the modeled region of the Pacific with UV accounting for two‐thirds of the reduction. In contrast, most other productivity models either ignore inhibition or only include PAR inhibition. Incorporation of E_max model responses into an existing spectral model of depth‐integrated, daily production will enable efficient global predictions of picophytoplankton productivity including inhibition.     

Responses of Antarctic Iridaea cordata (Rhodophyta) tetraspores exposed to ultraviolet radiation

In Antarctica ozone depletion is highest during spring, coinciding with the reproduction of many seaweed species. Propagules are the life-stage of an alga most susceptible to environmental perturbations. Therefore, fertile thalli of Iridaea cordata (Turner) Bory (Rhodophyta) were collected in the eulittoral of King George Island (Antarctica) to examine spore susceptibility to ultraviolet radiation (UVR). In the laboratory, freshly released tetraspores were exposed to photosynthetically active radiation (PAR) (400–700 nm), PAR+UV-A (320–700 nm) or PAR+UV-A+UV-B (280–700 nm). Photosynthetic efficiency was measured during 1–8 h of exposure and after 48 h of recovery. Additionally, mycosporine-like amino acids (MAAs) and DNA damage were determined. Saturating irradiance of photosynthesis of freshly released tetraspores was 57 µmol photons m⁻² s⁻¹. Exposure to increasing fluence of PAR reduced photosynthetic efficiency. UVR further decreased the photosynthetic efficiencies of the tetraspores but spores were able to recover completely after UVR exposure and 2 days post-cultivation under low PAR. DNA damage was minimal and lesions were effectively repaired under photoreactivating light. Concentrations of the MAAs shinorine and palythine were higher in tetraspores treated with UVR than in spores only exposed to PAR. Generally, the tetraspores show a good UV tolerance. This flexible response of the tetraspores of this species to changing radiation conditions enables the alga to grow along a considerable depth gradient from the sublittoral to the eulittoral where they can be exposed to enhanced UVBR under conditions of stratospheric ozone depletion.     

Ultraviolet radiation and consumer effects on a field-grown intertidal macroalgal assemblage in Antarctica

Ultraviolet radiation (UVR) research on marine macroalgae has hithero focussed on physiological effects at the organism level, while little is known on the impact of UV radiation on macroalgal assemblages and even less on interactive effects with other community drivers, e.g. consumers. Field experiments on macrobenthos are scarce, particularly in the Antarctic region. Therefore, the effects of UVR and consumers (mainly limpets were excluded) on early successional stages of a hard bottom macroalgal community on King George Island, Antarctica, were studied. In a two‐factorial design experimental units [(1) ambient radiation, 280–700 nm; (2) ambient minus UVB, 320–700 nm and (3) ambient minus UVR, 400–700 nm vs. consumer–no consumer] were installed between November 2004 and March 2005 (n= 4 plus controls). Dry mass, species richness, diversity and composition of macroalgal assemblages developing on ceramic tiles were followed. Consumers significantly suppressed green algal recruits and total algal biomass but increased macroalgal richness and diversity. Both UVA and UVB radiation negatively affected macroalgal succession. UVR decreased the density of Monostroma hariotii germlings in the first 10 weeks of the experiment, whereas the density of red algal recruits was significantly depressed by UVR at the end of the study. After 106 days macroalgal diversity was significantly higher in UV depleted than in UV‐exposed assemblages. Furthermore, species richness was significantly lower in the UV treatments and species composition differed significantly between the UV‐depleted and the UV‐exposed treatment. Marine macroalgae are very important primary producers in coastal ecosystems, serving as food for herbivores and as habitat for many organisms. Both, UVR and consumers significantly shape macroalgal succession in the Antarctic intertidal. Consumers, particularly limpets can mediate negative effects of ambient UVR on richness and diversity till a certain level. UVB radiation in general and an increase of this short wavelength due to stratospheric ozone depletion in particular may have the potential to affect the zonation, composition and diversity of Antarctic intertidal seaweeds altering trophic interactions in this system.     

Combined effects of ocean acidification and solar UV radiation on photosynthesis,growth, pigmentation and calcification of the coralline alga Corallina sessilis (Rhodophyta)

Previous studies have shown that increasing atmospheric CO₂ concentrations affect calcification in some planktonic and macroalgal calcifiers due to the changed carbonate chemistry of seawater. However, little is known regarding how calcifying algae respond to solar UV radiation (UVR, UVA+UVB, 280–400 nm). UVR may act synergistically, antagonistically or independently with ocean acidification (high CO₂/low pH of seawater) to affect their calcification processes. We cultured the articulated coralline alga Corallina sessilis Yendo at 380 ppmv (low) and 1000 ppmv (high) CO₂ levels while exposing the alga to solar radiation treatments with or without UVR. The presence of UVR inhibited the growth, photosynthetic O₂ evolution and calcification rates by13%, 6% and 3% in the low and by 47%, 20% and 8% in the high CO₂ concentrations, respectively, reflecting a synergistic effect of CO₂ enrichment with UVR. UVR induced significant decline of pH in the CO₂‐enriched cultures. The contents of key photosynthetic pigments, chlorophyll a and phycobiliproteins decreased, while UV‐absorptivity increased under the high pCO₂/low pH condition. Nevertheless, UV‐induced inhibition of photosynthesis increased when the ratio of particulate inorganic carbon/particulate organic carbon decreased under the influence of CO₂‐acidified seawater, suggesting that the calcified layer played a UV‐protective role. Both UVA and UVB negatively impacted photosynthesis and calcification, but the inhibition caused by UVB was about 2.5–2.6 times that caused by UVA. The results imply that coralline algae suffer from more damage caused by UVB as they calcify less and less with progressing ocean acidification.     

Multivariate analysis of intraspecific responses to UV-B radiation in white clover (Trifolium repens L.)

White clover (Trifolium repens L.) is experiencing increased levels of ultraviolet‐B (UV‐B) radiation in temperate pastures due to the depletion of the stratospheric ozone layer. Based on 17 morphological, morphogenetic and physiological attributes, this study analysed the consequences of enhanced UV‐B on 26 white clover populations using principal components analysis (PCA). After 18 d of exposure to 13·3 kJ m ^{? 2} d ^{? 1} UV‐B in controlled environments, UV‐B significantly decreased above‐ground and below‐ground plant growth attributes, epidermal cell surface area and maximum quantum efficiency of photosystem II photochemistry (F_v/F_m). Aspects of cell division and cell expansion both were negatively affected by UV‐B. Stomatal density, specific leaf mass, root‐to‐shoot ratio and levels of UV‐B‐absorbing compounds increased in response to UV‐B. In the multivariate analysis, the main dimension of UV‐B sensitivity was characterized by changes in plant growth attributes. Alterations in partitioning within and between plant organs constituted a secondary tier of UV‐B responsiveness. Plant characteristics related to UV‐B tolerance included lower growth rate, smaller epidermal cell surface area and higher UV‐B‐induced levels of UV‐B‐absorbing compounds. The results suggest overall UV‐B tolerance for slower‐growing populations from less productive habitats with higher natural UV‐B irradiance.     

SHORT‐TERM AND LONG‐TERM EFFECTS OF TEMPERATURE ON PHOTOSYNTHESIS IN THE DIATOM THALASSIOSIRA PSEUDONANA UNDER UVR EXPOSURES1

Temperature is expected to modify the effects of ultraviolet radiation (UVR) on photosynthesis by affecting the rate of repair. We studied the effect of short‐term (1 h) and long‐term (days) acclimation to temperature on UVR photoinhibition in the diatom Thalassiosira pseudonana Hasle et Heimdal. Photosynthesis was measured during 1 h exposures to varying irradiances of PAR and UVR + PAR at 15, 20, and 25°C, the latter corresponding to the upper temperature limit for optimal growth in T. pseudonana. The exposures allowed the estimation of photosynthesis–irradiance (P–E) curves and biological weighting functions (BWFs) for photoinhibition. For the growth conditions used, temperature did not affect photosynthesis under PAR. However, photoinhibition by UVR was highly affected by temperature. For cultures preacclimated to 20°C, the extent of UVR photoinhibition increased with decreasing temperature, from 63% inhibition of PAR‐only photosynthesis at 25°C to 71% at 20°C and 85% at 15°C. These effects were slightly modified after several days of acclimation: UVR photoinhibition increased from 63% to 75% at 25°C and decreased from 85% to 80% at 15°C. Time courses of photochemical efficiency (Φ_PSII) under UVR + PAR were also fitted to a model of UVR photoinhibition, allowing the estimation of the rates of damage (k) and repair (r). The r/k values obtained for each temperature treatment verified the responses observed with the BWF (R² = 0.94). The results demonstrated the relevance of temperature in determining primary productivity under UVR exposures. However, the results suggested that temperature and UVR interact mainly over short (hours) rather than long (days) timescales.     

RESPONSE OF LAMINARIA SACCHARINA (PHAEOPHYTA) GROWTH AND PHOTOSYNTHESIS TO SIMULTANEOUS ULTRAVIOLET RADIATION AND NITROGEN LIMITATION1

The brown macroalga Laminaria saccharina (L.) J. V. Lamour. was grown in large outdoor tanks at 50% ambient solar radiation for 3–4 weeks in July and August of 2000, 2001, and 2002, in either ambient or nitrogen (N)–enriched seawater and in either ambient light [PAR + ultraviolet radiation (UVR)] or ambient light minus UVR. Growth, N‐content, photosynthetic pigments, and RUBISCO content increased in N‐enriched seawater, indicating N‐limitation. UVR inhibited growth, but this inhibition was ameliorated by N‐enrichment. The response of growth to UVR could not be explained by changes in respiration and photosynthesis. Gross light‐saturated photosynthesis (P_max) remained unaffected by UVR but was significantly higher under N‐enrichment, as was dark respiration (R_d). UVR had no effect on pigments or N content. However, RUBISCO contents were low in the presence of UVR, reflecting the overall change in soluble cellular protein. Overall, our data indicate that the response to UVR in L. saccharina depends on other environmental factors, such as N, and these effects need to be considered when evaluating the response of macroalgae to increased UVR.     

COMBINED EFFECTS OF ULTRAVIOLET RADIATION AND TEMPERATURE ON MORPHOLOGY,PHOTOSYNTHESIS, AND DNA OF ARTHROSPIRA (SPIRULINA) PLATENSIS (CYANOPHYTA)1

Natural levels of solar UVR were shown to break and alter the spiral structure of Arthrospira (Spirulina) platensis (Nordst.) Gomont during winter. However, this phenomenon was not observed during summer at temperatures of ～30°C. Since little has been documented on the interactive effects of solar UV radiation (UVR; 280–400 nm) and temperature on cyanobacteria, the morphology, photosynthesis, and DNA damage of A. platensis were examined using two radiation treatments (PAR [400–700 nm] and PAB [PAR + UV‐A + UV‐B: 280–700]), three temperatures (15, 22, and 30°C), and three biomass concentrations (100, 160, and 240 mg dwt [dry weight] · L^?1). UVR caused a breakage of the spiral structure at 15°C and 22°C, but not at 30°C. High PAR levels also induced a significant breakage at 15°C and 22°C, but only at low biomass densities, and to lesser extent when compared with the PAB treatment. A. platensis was able to alter its spiral structure by increasing helix tightness at the highest temperature tested. The photochemical efficiency was depressed to undetectable levels at 15°C but was relatively high at 30°C even under the treatment with UVR in 8 h. At 30°C, UVR led to 93%–97% less DNA damage when compared with 15°C after 8 h of exposure. UV‐absorbing compounds were determined as negligible at all light and temperature combinations. The possible mechanisms for the temperature‐dependent effects of UVR on this organism are discussed in this paper.