UVB radiation may ameliorate photoinhibition in specific shallow-water tropical marine macrophytes

Anthropogenic stratospheric ozone depletion causes an increase in UVB radiation impinging on the earth's surface, which is a threat to plants not adapted to higher UVB irradiances. Investigations were carried out among tropical marine macrophytes, Turbinaria turbinata, Sargassum polyceratium var. ovatum, Padina sanctae-crucis, Lobophora variegate, Dictyota spec., Halimeda discoidea, Udotea flabellum, Thalassia testudinum and Syringodium filiforme collected from 0.3 to 26 m depths at the Belizean barrier reef, where ultraviolet radiation (UV)-irradiances are naturally high. Photoinhibition was induced under full solar, UV (UVA + UVB), and UVA only-depleted radiation conditions. Photosynthetic activity during high radiation stress and during recovery in reduced solar radiation was determined in vivo by measuring fluorescence changes using a PAM fluorometer device. Generally, UV caused an additional decrease of photosynthetic performance during high light stress which varies according to species, depth of growth and UV penetration at the site of collection; an observation in concordance with the conventional harmful UV-radiation effects on phototrophs. When solar radiation was reduced by 50%, significant photosynthetic recovery was observed. However, some shallow water species which are adapted to high UV were observed to recover less under treatment with depleted solar UVB radiation. Our result supports earlier reports that UVB causes not only negative effects on photosynthesis, but may also facilitate or induce recovery processes in aquatic macrophytes acclimated to high solar radiation which grow at the upper shoreline. Among the eulittoral macroalgae, e.g. Dictyota spec., P. sanctae-crucis, and H. discoidea and the seagrass T. testudinum, initiation of photosynthetic recovery processes in the presence of low irradiance of short UV-wavelengths may present an ecophysiological advantage compared to macrophytes which initiate photosynthetic recovery process during low light or in the absence of UV.     

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.     

Effect of solar ultraviolet radiation on survival of krill larvae and copepods in Antarctic Ocean

The effect of solar ultraviolet radiation on the survival rate of Antarctic zooplankton was examined in February–March in 2002. We investigated survival rate of calyptopis larvae of Euphausia superba and late copepodite stages (IV and V) of large dominant calanoid species, Calanoides acutus and Calanus propinquus reared in quartz jars with three different radiation regimes (total radiation, exclusion of UVB, exclusion of UVA and UVB) and a dark control. The survival rates of the krill larvae decreased after 3 days from start of the experiment, being below 50% at 4 days in the treatments with total radiation and exclusion of UVB, although most individuals could survive until the end of the experiments in the treatments with exclusion of both UVA and UVB and dark control. The calanoid juveniles showed almost same pattern of survival curves as the krill larvae did, but survived slightly longer. Although >10% of surface UVA radiation at 340 and 380 nm penetrated down to 30 m, both C. acutus and C. propinquus were mostly distributed above 20 m. Surface swarm of the krill larvae can be often recognized in the previous studies. These results suggest that not only solar UVB but also UVA radiation potentially lower the survival rate of Antarctic zooplankton at depth less than 20 m.     

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)     

Temperature dependence of UV radiation effects in Arctic and temperate isolates of three red macrophytes

The temperature dependence of UV effects was studied for Arctic and temperate isolates of the red macrophytes Palmaria palmata, Coccotylus truncatus and Phycodrys rubens. The effects of daily repeated artificial ultraviolet B and A radiation (UVBR: 280–320?nm, UVAR: 320–400?nm) treatments were examined for all isolates at 6, 12 and 18?°C by measuring growth, optimal quantum yield of PSII (F_v/F_m) and cyclobutane-pyrimidine dimer (CPD) accumulation. Furthermore, possible ecotypic differences in UV sensitivity between Arctic and temperate isolates were evaluated. Large species-specific differences in UV sensitivity were observed for all parameters: the lower subtidal species C. truncatus and P. rubens were highly sensitive to the UV treatments, whereas P. palmata, which predominantly occurs in the upper subtidal zone, was not affected by these treatments. Only minor differences were found between Arctic and temperate isolates, suggesting that no differences in UV sensitivity have evolved in these species. Relative growth rates were temperature-dependent, whereas species-specific UV effects on growth rates were relatively independent of temperature. In contrast, the species-specific decrease in F_v/F_m and its subsequent recovery were temperature-dependent in all species. UV effects on F_v/F_m were lower at 12 and 18?°C compared with 6?°C. In addition, UV effects on F_v/F_m decreased in the course of the experiment at all temperatures, indicating acclimation to the UV treatments. CPDs accumulated during the experiment in both isolates of P. rubens, whereas CPD concentrations remained low for the other two species. CPD accumulation appeared to be independent of temperature. The results suggest that summer temperatures occurring in temperate regions facilitate repair of UV-induced damage and acclimation to UV radiation in these algae compared with Arctic temperatures. Because the differences in UV effects on F_v/F_m, growth and CPD accumulation were relatively small over a broad range of temperatures, it was concluded that the influence of temperature on UV effects is small in these species.     

Sensitivity of Laminariales zoospores from Helgoland (North Sea) to ultraviolet and photosynthetically active radiation: implications for depth distribution and seasonal reproduction

Depth distribution of kelp species in Helgoland (North Sea) is characterized by occurrence of Laminaria digitata in the upper sublittoral, whereas L. saccharina and L. hyperborea dominate the mid and lower sublittoral region. Laminaria digitata is fertile in summer whereas both other species are fertile in autumn/winter. To determine the light sensitivity of the propagules, zoospores of L. digitata, L. saccharina and L. hyperborea were exposed in the laboratory to different exposure times of photosynthetically active radiation (PAR; 400–700 nm), PAR + UVA radiation (UVAR; 320–400 nm) and PAR + UVAR + UVB radiation (UVBR; 280–320 nm). Optimum quantum yield of PSII and DNA damage were measured after exposure. Subsequently, recovery of photosynthetic efficiency and DNA damage repair, as well as germination rate were measured after 2 and 3 d cultivation in dim white light. Photosynthetic efficiency of all species was photoinhibited already at 20 µmol photons m⁻² s⁻¹ PAR, whereas UV radiation (UVR) had a significant additional effect on photoinhibition. Recovery of the PSII function was observed in all species but not in spores exposed to irradiation longer than 4 h of PAR + UVA + UVB and 8 h of PAR + UVA. The amount of UVB-induced DNA damage measured as cyclobutane–pyrimidine dimers (CPDs) increased with exposure time and highest damage was detected in the spores of lower subtidal L. hyperborea relative to the other two species. Significant removal of CPDs indicating repair of DNA damage was observed in all species after 2 d in low white light especially in the spores of upper subtidal L. digitata. Therefore, efficient DNA damage repair and recovery of PSII damage contributed to the germination success but not in spores exposed to 16 h of UVBR. UV absorption of zoospore suspension in L. digitata is based both on the absorption by the zoospores itself as well as by exudates in the medium. In contrast, the absorption of the zoospore suspension in L. saccharina and L. hyperborea is based predominantly on the absorption by the exudates in the medium. This study indicates that UVR sensitivity of zoospores is related to the seasonal zoospore production as well as the vertical distribution pattern of the large sporophytes.     

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.     

UV‐radiation effects on photosynthesis and photoprotection in gametophytic and sporophytic stages of the freshwater red alga Kumanoa ambigua (Rhodophyta,Batrachospermales)

The aim of this study was to analyze the photosynthetic performance of gametophytic and sporophytic (‘Chantransia’) stages of Kumanoa ambigua in culture under UV radiation. We hypothesized that both life history stages of K. ambigua would exhibit different photosynthetic responses to UVR exposure. Experiments were performed under three conditions: (i) photosynthetically active radiation (PAR) only (400–700 nm), P control; (ii) PAR + UVA (320–700 nm), PA treatment; and (iii) PAR + UVA + UVB (280–700 nm), PAB treatment. The photosynthetic parameters were measured as in vivo chlorophyll a fluorescence. Differences were found between life stages, observing higher values of NPQ and effective quantum yields (ΔF/F_m′) under UVA and PAR in gametophytes compared to sporophytes. One type of mycosporine‐like amino acid (MAA) was detected in the gametophyte in all treatments, but not in the ‘Chantransia’ stage. The increased photosynthetic performance for some parameters and the presence of MAA in gametophyte suggest that it is less sensitive to UV radiation, particularly UVA, in comparison to sporophyte under culture conditions. This approach is relevant for a better understanding of the adaptation and physiological acclimation of freshwater Rhodophyta to varying light climates in terms of global changes.     

Ultraviolet-B Wavelengths Regulate Changes in UV Absorption of Cleaner Fish Labroides dimidiatus Mucus

High-energy wavelengths in the ultraviolet-B (UVB, 280-315 nm) and the UVA (315-400-nm) portion of the spectrum are harmful to terrestrial and aquatic organisms. Interestingly, UVA is also involved in the repair of UV induced damage. Organisms living in shallow coral reef environments possess UV absorbing compounds, such as mycosporine-like amino acids, to protect them from UV radiation. While it has been demonstrated that exposure to UV (280-400 nm) affects the UV absorbance of fish mucus, whether the effects of UV exposure vary between UVB and UVA wavelengths is not known. Therefore, we investigated whether the UVB, UVA, or photosynthetically active radiation (PAR, 400-700 nm) portions of the spectrum affected the UV absorbance of epithelial mucus and Fulton’s body condition index of the cleaner fish Labroides dimidiatus. We also compared field-measured UV absorbance with laboratory based high-performance liquid chromatography measurements of mycosporine-like amino acid concentrations. After 1 week, we found that the UV absorbance of epithelial mucus was higher in the UVB+UVA+PAR treatment compared with the UVA+PAR and PAR only treatments; after 2 and 3 weeks, however, differences between treatments were not detected. After 3 weeks, Fulton’s body condition index was lower for fish in the UVB+UVA+PAR compared with PAR and UVA+PAR treatments; furthermore, all experimentally treated fish had a lower Fulton’s body condition index than did freshly caught fish. Finally, we found a decrease with depth in the UV absorbance of mucus of wild-caught fish. This study suggests that the increase in UV absorbance of fish mucus in response to increased overall UV levels is a function of the UVB portion of the spectrum. This has important implications for the ability of cleaner fish and other fishes to adjust their mucus UV protection in response to variations in environmental UV exposure.     

Effects of solar radiation and solar radiation deprived of UV-B and total UV on photosynthetic oxygen production and pulse amplitude modulated fluorescence in the brown alga Padina pavonia

Abstract: The effects of solar radiation on photosynthetic oxygen production and pulse amplitude modulated (PAM) fluorescence were measured in the marine brown macroalga Padina pavonia harvested from different depths from the Greek coast near Korinth. In fluence rate-response curves the light compensation point for photosynthetic oxygen production increased and the saturation level decreased with increasing exposure time to solar radiation. Cutting off the UV-B wavelength range (280–315 nm) from solar radiation reduced the inhibition of photosynthesis, and the organisms were less affected when all of the UV radiation was filtered out. Algae collected from 7 m depth were much more prone to photoinhibition than those harvested from rock pools exposed to unfiltered solar radiation. During continuous exposure to solar radiation, rock pool algae showed photoinhibition after longer periods of time than specimens from 7 m or from dark adapted habitats. When subjected to unfiltered solar radiation the ratio of the variable fluorescence to the maximal fluorescence     (F_v = F_m− F_o) rapidly declined with increasing exposure time. However, again algae from 7 m depth were more prone to photoinhibition than rock pool algae. The differences between the two ecological strains were less obvious when UV-B or total UV was removed from solar radiation. Only in the latter case a complete recovery was observed after 2 h while, when exposed to unifiltered sunlight, only the rock pool algae recovered completely within that time.     

Immobile and tough versus mobile and weak: effects of ultraviolet B radiation on eggs and larvae of Manduca sexta

Although indirect effects of solar ultraviolet (UV) radiation on insects are well known (e.g. UV radiation can modify plant chemistry), direct effects of solar radiation on insects have received little attention. Radiation in the UVB range (300–320 nm) is damaging because it is absorbed directly by proteins and DNA. UVB should be toughest on immobile or small life stages, such as eggs or early larval instars. In the present study, the effects of UVB radiation on eggs and larvae of the tobacco hornworm Manduca sexta L. (Lepidoptera: Sphingidae) are examined. The present study aimed to address: what natural levels of UV do they experience; how does UVB affect the performance of eggs; and how does it affect the performance of larvae? In addition, do M. sexta larvae use behaviour to avoid UVB exposure and, consequently, are they physiologically less robust to UVB? In these experiments, eggs and late larval instars of M. sexta are found to be robust to natural levels of UV radiation. By contrast, young larvae are not only more susceptible to damage from UVB, but also they use behavioural means to avoid it. The strategy of using behaviour may relax selection pressures on morphological and physiological mechanisms for preventing (or recovering from) damage by environmental UV radiation.     

Bipyrimidine photoproducts rather than oxidative lesions are the main type of DNA damage involved in the genotoxic effect of solar UVA radiation

Exposure to solar UV radiation gives rise to mutations that may lead to skin cancer. UVA (320-340 nm) constitutes the large majority of solar UV radiation but is less effective than UVB (290-320 nm) at damaging DNA. Although UVA has been implicated in photocarcinogenesis, its contribution to sunlight mutagenesis has not been elucidated, and DNA damage produced by UVA remains poorly characterized. We employed HPLC-MS/MS and alkaline agarose gel electrophoresis in conjunction with the use of specific DNA repair proteins to determine the distribution of the various classes and types of DNA lesions, including bipyrimidine photoproducts, in Chinese hamster ovary cells exposed to pure UVA radiation, as well as UVB and simulated sunlight (lambda > 295 nm) for comparison. At UVA doses compatible with human exposure, oxidative DNA lesions are not the major type of damage induced by UVA. Indeed, single-strand breaks, oxidized pyrimidines, oxidized purines (essentially 8-oxo-7,8-dihydroguanine), and cyclobutane pyrimidine dimers (CPDs) are formed in a 1:1:3:10 ratio. In addition, we demonstrate that, in contrast to UVB and sunlight, UVA generates CPDs with a large predominance of TT CPDs, which strongly suggests that they are formed via a photosensitized triplet energy transfer. Moreover, UVA induces neither (6-4) photoproducts nor their Dewar isomers via direct absorption. We also show that UVA photons contained in sunlight, rather than UVB, are implicated in the photoisomerization of (6-4) photoproducts, a quickly repaired damage, into poorly repaired and highly mutagenic Dewar photoproducts. Altogether, our data shed new light on the deleterious effect of UVA.     

Comutagenesis of sodium arsenite with ultraviolet radiation in Chinese hamster V79 cells

Summary Solar ultraviolet radiation has been associated with the induction of skin cancer. Recent studies have indicated that near-ultraviolet, especially UVB, is mutagenic. Exposure to trivalent inorganic arsenic compounds has also been associated with increased skin cancer prevalence. Trivalent arsenic compounds are not mutagenicper se, but are comutagenic with a number of cancer agents. Here, we test the hypothesis that arsenite enhances skin cancer via its comutagenic action with solar ultraviolet radiation. Irradiation of Chinese hamster V79 cells with UVA (360 nm), UVB (310 nm) and UVC (254 nm) caused a fluence-dependent increase in mutations at thehprt locus. On an energy basis, UVC was the most mutagenic and UVA the least. However, when expressed as a function of toxicity, UVB was more mutagenic than UVC. Nontoxic concentrations of arsenite increased the toxicity of UVA, UVB and UVC. Arsenite acted as a comutagen at the three wavelengths; however, higher concentrations of arsenite were required to produce a significant (P < 0.05) comutagenic response with UVB. The increased mutagenicity of UVB and UVA by arsenite may play a role in arsenite-related skin cancers.     

IMPACTS OF SOLAR UV RADIATION ON THE PHOTOSYNTHESIS,GROWTH, AND UV‐ABSORBING COMPOUNDS IN GRACILARIA LEMANEIFORMIS (RHODOPHYTA) GROWN AT DIFFERENT NITRATE CONCENTRATIONS1

Solar ultraviolet radiation (UVR, 280–400 nm) is known to affect macroalgal physiology negatively, while nutrient availability may affect UV‐absorbing compounds (UVACs) and sensitivity to UVR. However, little is known about the interactive effects of UVR and nitrate availability on macroalgal growth and photosynthesis. We investigated the growth and photosynthesis of the red alga Gracilaria lemaneiformis (Bory) Grev. at different levels of nitrate (natural or enriched nitrate levels of 41 or 300 and 600 μM) under different solar radiation treatments with or without UVR. Nitrate‐enrichment enhanced the growth, resulted in higher concentrations of UVACs, and led to negligible photoinhibition of photosynthesis even at noon in the presence of UVR. Net photosynthesis during the noon period was severely inhibited by both ultraviolet‐A radiation (UVA) and ultraviolet‐B radiation (UVB) in the thalli grown in seawater without enriched nitrate. The absorptivity of UVACs changed in response to changes in the PAR dose when the thalli were shifted back and forth from solar radiation to indoor low light, and exposure to UVR significantly induced the synthesis of UVACs. The thalli exposed to PAR alone exhibited higher growth rates than those that received PAR + UVA or PAR + UVA + UVB at the ambient or enriched nitrate concentrations. UVR inhibited growth approximately five times as much as it inhibited photosynthesis within a range of 60–120 μg UVACs · g^?1 (fwt) when the thalli were grown under nitrate‐enriched conditions. Such differential inhibition implies that other metabolic processes are more sensitive to solar UVR than photosynthesis.     

Wavelength-dependent induction of UV-absorbing mycosporine-like amino acids in the red alga Chondrus crispus under natural solar radiation

Polychromatic response spectra for the induction of UV absorbing mycosporine-like amino acids (MAAs) were calculated after exposing small thalli of the red alga Chondrus crispus under various cut-off filters to natural solar radiation on the North Sea island Helgoland, Germany. The laboratory-grown specimens typically contain only traces of palythine and synthesise five different MAAs rapidly and in high concentrations after being transplanted into shallow water. The resulting qualitative and quantitative patterns of MAA induction differed markedly with respect to spectral distribution. Furthermore, the wavebands effective for MAA induction vary within the MAA. UV-B radiation had a negative effect on the accumulation of the major MAAs shinorine (λ_max=334 nm) and palythine (λ_max=320 nm), while short wavelength UV-A exhibits the highest quantum efficiency on their synthesis. In contrast, the synthesis of asterina-330 (λ_max=330 nm), palythinol (λ_max=332 nm) and palythene (λ_max=360 nm) was mainly induced by UV-B radiation. Whether the synthesis of shinorine and palythine is induced by a photoreceptor with an absorption maximum in the short wavelength UV-A and whether a second photoreceptor absorbing UV-B radiation is responsible for the induction of asterina-330, palythinol and palythene remains to be studied.Our results show that C. crispus has a high capacity to adapt flexibly the qualitative and quantitative MAA concentration to the prevailing spectral distribution of irradiance. On one hand, this is regarded as an important aspect with respect to the acclimation of algae to increasing UV-B irradiance in the context of ongoing depletion of stratospheric ozone. On the other hand, the experiment demonstrates that UV-A irradiance is more important for the induction of the major MAAs shinorine and palythine than UV-B.     

Biological weighting functions for DNA damage in sea urchin embryos exposed to ultraviolet radiation

Laboratory experiments examining the effects of ultraviolet radiation (UVR, 290-400 nm) on DNA damage were carried out using the embryos of three species of sea urchins from different habitats; Strongylocentrotus droebachiensis from the Gulf of Maine, Sterechinus neumayeri from the Antarctic, and Evechinus chloroticus from New Zealand. All three species exhibited significant amounts of accumulated DNA damage, measured as cyclobutane pyrimidine dimers (CPD) photoproducts, when exposed to UVR in the laboratory. Biological weighting functions (BWFs) revealed that S. neumayeri has significantly higher sensitivity to UVR-induced DNA damage across most of the UVR spectrum compared to the other two species, and all species were observed to have weightings in the ultraviolet-A (UVA, 320-400 nm) portion of the spectrum. The increased sensitivity to ultraviolet-B (290-320 nm) and UVA in S. neumayeri is correlated with the lowest concentration of UVR absorbing compounds observed in the embryos of the three species of urchin used in this study. Sea urchin embryos and larvae in the respective habitats of the species tested are known to occur within 5 m of the surface of the ocean where both UVB and UVA wavelengths occur. Solar irradiances of UVR at a depth of 5 m, weighted using the urchin DNA damage BWFs, show that E. chloroticus receives the greatest amount of biologically effective UVR despite having the lowest wavelength dependent weightings for DNA damage when compared to the other two species.     

Effects of solar UV stress on chlorophyll fluorescence kinetics of intertidal macroalgae from southern Spain: a case study in Gelidium species

Photoinhibition and recovery kinetics after short exposure to solar radiation following three different irradiance treatments of irradiances (PAR, PAR+UVA and PAR+UVA+UVB) was assessed in two intertidal species of the genus Gelidium, Gelidium sesquipedale and G. latifolium, collected from Tarifa (southern Spain) using in vivo chlorophyll fluorescence (PAM fluorometry). After 3 h UV radiation exposure, optimal quantum efficiency (Fv/Fm) in G. sesquipedale decreased between 25 and 35% relative to the control. Under PAR alone, values decreased to 60%. In G. latifolium, photoinhibition did not exceed 40%. Similar results were found for the effective quantum yield (ΔF/Fm′), however, no marked differences in relation to light treatments were seen. When plants were shaded for recovery from stress, only in G. latifolium a significant increase in photosynthesis was observed (between 80 and 100% of control). In contrast, photosynthesis of G. sesquipedale suffered a chronic photoinhibition or photodamage under the three light irradiances. Full solar radiation (PAR+UVA+UVB) affected also the electron transport rate in both species. Here, initial slopes of electron transport vs. irradiance curves decreased up to 60% of controls. Although the recovery kinetic under PAR+UVA+UVB conditions was delayed in G. latifolium, after 24 h recovery this species reached significantly higher than G. sesquipedale. PAR impaired electron trasport only in G. sesquipedale. Overall, both species are characterized by different capacity to tolerate enhanced solar radiation. G. latifolium is a sun adapted plant, well suited to intertidal light conditions, whereas G. sesquipedale, growing at shaded sites in the intertidal zone, is more vulnerable to enhanced UV radiation. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of UVB radiation on freshwater autotrophic and heterotrophic picoplankton in a subalpine lake

The effects of UVB radiation on the activity of heterotrophic (HPP) and autotrophic (APP) picoplankton (0.2-2 m) and of autotrophic assemblages >2 m has been measured and compared. Under natural UVB irradiance in a large, deep, oligotrophic subalpine lake (Lago Maggiore, 4555N) with mean dissolved organic carbon (DOC) concentrations of 1 mg Cl^-1, the microorganisms of the two size fractions were not significantly photoinhibited in their autotrophic and heterotrophic activities. The vertical attenuation coefficient (Kd) for irradiance at 305 nm ranged from 1.45 to 1.67 during spring and summer. The mixing layer extended to a greater depth than the layer affected by UVB radiation (z1% < zmix), thus enabling the microorganisms living there to photoadapt. As the assimilation numbers of APP and nanoplankton were higher at 2 m depth than near the surface, we suspected that the influence of longer wavelength (UVA, photosynthetically active radiation) could be stronger than UVB in affecting the photosynthetic efficiency of natural populations. The artificial increase in UVB irradiance had a higher detrimental effect on HPP due to their smaller size, less protection and indirect effects through autotrophic cell inhibition. Picocyanobacteria were percentually more affected by UVB than nanoplankton during April due to the presence of diatoms, which are more resistant than other algal groups to solar UVB irradiance. Furthermore, picocyanobacteria had lower assimilation numbers with respect to larger phytoplankton in the quartz tubes during stratification.      

Assessment of oxidative stress in the planktonic diatom Thalassiosira pseudonana in response to UVA and UVB radiation

Induction of oxidative stress by UVA and UVB in the diatom Thalassiosirapseudonana was experimentally studied. Cells, pre-grown in alight-limited continuous culture, were incubated for 4 h at175 µmol m^-2s^-1photosynthetically active radiation, withoptional supplementary UVA at an unweighted dose rate of 0.70W m^-2, or 2.79 W m^-2UVA plus 0.45 W m^-2UVB (unweighted). A fluorescence-basedmeasure of photosynthetic efficiency (F_v/F_m) decreased from0.69 to 0.58 in the presence of UVB, whereas UVA caused a minordecrease of F_v/F_m. Quantitative analysis of confocal imagesshowed a minor increase of active oxygen production associatedwith supplemental UVA alone, and a 100% increase with additionalUVB. Cellular malondialdehyde, an indicator of lipid peroxidationby active oxygen, almost doubled under UVA and increased three-foldwith additional UVB. Activities of superoxide dismutase (scavengingactive oxygen) and glutathione reductase (reducing GSSG to GSH)increased in response to UVB exposure, whereas ascorbate peroxidaseactivities did not. UVB caused a minor decrease in the glutathioneratio GSH : (GSH + 0.5GSSG), which indicates a moderate oxidativestress.     

Differential responses of tetrasporophytes and gametophytes of Mazzaella laminarioides (Gigartinales,Rhodophyta) under solar UV radiation

The effects of solar UV radiation on mycosporine‐like amino acids (MAAs), growth, photosynthetic pigments (Chl a, phycobiliproteins), soluble proteins (SP), and C and N content of Mazzaella laminarioides tetrasporophytes and gametophytes were investigated. Apical segments of tetrasporophytes and gametophytes were exposed to solar radiation under three treatments (PAR [P], PAR+UVA [PA], and PAR+UVA+UVB [PAB]) during 18 d in spring 2009, Punta Arenas, Chile. Samples were taken after 2, 6, 12, and 18 d of solar radiation exposure. Most of the parameters assessed on M. laminarioides were significantly influenced by the radiation treatment, and both gametophytes and tetrasporophytes seemed to respond differently when exposed to high UV radiation. The two main effects promoted by UV radiation were: (i) higher synthesis of MAAs in gametophytes than tetrasporophytes at 2 d, and (ii) a decrease in phycoerythrin, phycocyanin, and SPs, but an increase in MAA content in tetrasporophytes at 6 and 12 d of culture. Despite some changes that were observed in biochemical parameters in both tetrasporophytes and gametophytes of M. laminarioides when exposed to UVB radiation, these changes did not promote deleterious effects that might interfere with the growth in the long term (18 d). The tolerance and resistance of M. laminarioides to higher UV irradiance were expected, as this intertidal species is exposed to variation in solar radiation, especially during low tide.