Combined effects of solar ultraviolet radiation and nutrients addition on growth, biomass and taxonomic composition of coastal marine phytoplankton communities of Patagonia

Experiments (6-8 days) were conducted during late summer, late fall and late winter, 2003 with waters collected off Bahía Nueva, Chubut, Argentina (42.7 degrees S, 65 degrees W) to determine the combined effects of solar ultraviolet radiation (UVR, 280-400 nm) and nutrient addition on phytoplankton communities. Samples were put in UVR-transparent containers and incubated under two radiation treatments: (a) Samples exposed to full solar radiation (PAB treatment, 280-400 nm) and (b) Samples exposed only to PAR (PAR treatment, 400-700 nm). At the beginning of the experiments, nutrients (i.e., NaPO(4)H(2) and NaNO(3)) were added to one set of samples from each radiation treatment (N cultures) whereas in the other set, nutrients remained at the concentration of the seawater. Chlorophyll a, biomass, UV-absorbing compounds and taxonomic composition were recorded throughout the experiments. N cultures always had significantly higher growth rates (P<0.05) than that in non-enriched cultures. At the beginning of experiments, phytoplankton communities were generally dominated by monads and flagellates but by the end, diatoms comprised the bulk of biomass, with only one to four taxa dominating, suggesting a selection towards more tolerant/less sensitive species. Over the experimental time frame, the observed taxonomic changes were mostly due to nutrient availability, and to a lesser extent to solar UVR exposure. Overall, the results indicate that environmental conditions (i.e., light history, nutrient concentration) together with the physiological status of the cells play a very important role at the time to assess the combined effect of nutrient addition and solar radiation on coastal phytoplankton assemblages from Patagonia.     

Effects of solar ultraviolet radiation on photosynthesis of the marine red tide alga Heterosigma akashiwo (Raphidophyceae)

In order to assess the short- and long-term impacts of UV radiation (UVR, 280-400nm) on the red tide alga, Heterosigma akashiwo, we exposed the cells to three different solar radiation treatments (PAB: 280-700nm, PA: 320-700nm, P: 400-700nm) under both solar and artificial radiation. A significant decrease in the effective quantum yield (Y) during high irradiance periods (i.e., local noon) was observed, but the cells partially recovered during the evening hours. Exposure to high irradiances for 15, 30, and 60min under a solar simulator followed by the recovery (8h) under dark, 9 and 100micromolphotonsm(-2)s(-1) of PAR, highlighted the importance of the irradiance level during the recovery period. Regardless the radiation treatments, the highest recovery (both in rate and total Y) was found at a PAR irradiance of 9micromolphotonsm(-2)s(-1), while the lowest was observed at 100micromolphotonsm(-2)s(-1). In all experiments, PAR was responsible for most of the observed inhibition; nevertheless, the cells exposed only to PAR had the highest recovery in any condition, as compared to the other radiation treatments. In long-term experiments (10 days) using semi-continuous cultures, there was a significant increase of UV-absorbing compounds (UV(abc)) per cell from 1.2 to >4x10(-6)microgUV(abc)cell(-1) during the first 3-5 days of exposure to solar radiation. The highest concentration of UV(abc) was found in samples exposed in the PAB as compared to PA and P treatments. Growth rates (mu) mimic the behavior of UV-absorbing compounds, and during the first 5 days mu increased from <0.2 to ca. 0.8, and stayed relatively constant at this value during the rest of the experiment. The inhibition of the Y decreased with increasing acclimation of cells. All our data indicates that H. akashiwo is a sensitive species, but was able acclimate relatively fast (3-5 days) synthesizing UV-absorbing compounds and thus reducing any impact either on photosystem II or on growth.     

Variability of UVR effects on photosynthesis of summer phytoplankton assemblages from a tropical coastal area of the South China Sea

From June to September 2005, we carried out experiments to determine the ultraviolet radiation (UVR) -induced photoinhibition of summer phytoplankton assemblages from a coastal site of the South China Sea. Variability in taxonomic composition was determined throughout the summer, with a peak chlorophyll a (chl a approximately 20 microg chl a L(-1)) dominated by the diatom Skeletonema costatum that was detected early in the study period; the rest of the time samples were characterized by monads and flagellates, with low chl a values (1-5 chl a microg L(-1)). Surface water samples were placed in quartz tubes, inoculated with radiocarbon and exposed to solar radiation for 2-3 h to determine photosynthetic rates under three quality radiation treatments (i.e. PAB, 280-700 nm; PA, 320-700 nm and P, 400-700 nm) using different filters and under seven levels of ambient irradiance using neutral density screens (P vs E curves). UVR inhibition of samples exposed to maximum irradiance (i.e. at the surface) varied from -12.2% to 50%, while the daytime-integrated UVR-related photoinhibition in surface seawater varied from -62% to 7%. The effects of UVR on the photosynthetic parameters P(B)(max) and E(k) were also variable, but UV-B accounted for most of the observed variability. During sunny days, photosynthesis of microplankton (>20 microm) and piconanoplankton (<20 microm) were significantly inhibited by UVR (mostly by UV-B). However, during cloudy days, while piconanoplankton cells were still inhibited by UVR, microplankton cells used UVR (mostly UV-A) as the source of energy for photosynthesis, resulting in higher carbon fixation in samples exposed to UVR than the ones exposed only to photosynthetically active radiation (PAR). Our results indicate that size structure and cloudiness clearly condition the overall impact of UVR on phytoplankton photosynthesis in this tropical site of South China. In addition, model predictions for this area considering only PAR for primary production might have underestimated carbon fixation due to UVR contribution.     

Vertical migration and motility responses in three marine phytoplankton species exposed to solar radiation

Diurnal vertical migration in the water column and the impact of solar radiation on motility were investigated in three marine phytoplankton species: Tetraselmis suecica, Dunaliella salina and Gymnodinium chlorophorum. Cells were exposed to solar radiation either in ultraviolet radiation (UVR, 280-400 nm) transparent Plexiglas tubes (45 cm length, 10 cm diameter) or in quartz tubes under three radiation treatments: PAB (280-700 nm), PA (320-700 nm) and P (400-700 nm). The three species displayed different behavior after exposure to solar radiation. Tetraselmis suecica was insensitive to UVR and under high solar radiation levels, cells accumulated preferentially near the surface. Exposure experiments did not indicate any significant changes in swimming speed nor in the percentage of motile cells after 5 h of exposure. On the other hand, D. salina was sensitive to UV-B displaying a significant decrease in swimming speed and percentage of motile cells after 2-3 h of exposure. Moreover, D. salina cells migrated deep in the water column when irradiance was high. The response of G. chlorophorum was in between that of the other two species tested, with a slight (but significant) decrease in swimming speed and percentage of motile cells in all radiation treatments after 5 h of exposure. While G. chlorophorum cells were more or less homogenously distributed in the water column, a slight (but significant) avoidance response to high radiation was observed at local noon, with cells migrating deep in the water column. Our data clearly indicate that these sub-lethal effects of solar radiation are species-specific and they might have important implications for the aquatic ecosystem.     

Influence of temperature and UVR on photosynthesis and morphology of four species of cyanobacteria

During the late austral spring of 2009 we carried out experiments (4days of duration) with four cyanobacteria species, Anabaena sp., Nostoc sp., Arthrospira platensis and Microcystis sp., to assess the combined effects of temperature and solar radiation on photosynthesis performance and morphology. Two experimental temperatures (18°C and 23°C, simulating a 5°C increase under a scenario of climate change) and three radiation treatments (by using different filters/materials) were implemented: (i) P (PAR, 400-700nm), (ii) PA (PAR+UV-A, 320-700nm) and, (iii) PAB (PAR+UV-A+UV-B, 280-700nm). In general, samples under the P treatment had less decrease/higher recovery rates of effective photochemical quantum yield (Y) than those receiving UV-A or UV-A+UV-B. The effects of increased temperature were species-specific: At the end of the experiments, it was seen that increased temperature benefited photosynthetic performance of Anabaena sp. and Nostoc sp. but not of Microcystis sp. and A. platensis. Higher temperature was also associated to an increase in the chain area of Anabaena sp., and to bigger trichomes in A. platensis; however, no morphological effects were observed in Microcystis sp. In addition, in Nostoc sp. the increase in temperature counteracted the UVR impact on the reduction of the chain area. How these effects and mechanisms will affect the trophodynamics and production of aquatic ecosystems is still uncertain, but the specificity of the responses suggests that not all cyanobacteria would be equally benefited by temperature increases therefore affecting the balance and interaction among species in the water column.     

Antioxidative Responses of Two Marine Microalgae During Acclimation to Static and Fluctuating Natural UV Radiation

Photoacclimation properties were investigated in two marine microalgae exposed to four ambient irradiance conditions: static photosynthetically active radiation (PAR: 400–700 nm), static PAR + UVR (280–700 nm), dynamic PAR and dynamic PAR + UVR. High light acclimated cultures of Thalassiosira weissflogii and Dunaliella tertiolecta were exposed outdoors for a maximum of 7 days. Dynamic irradiance was established by computer controlled vertical movement of 2 L bottles in a water filled basin. Immediate (<24 h), short-term (1–3 days) and long-term (4–7 days) photoacclimation was followed for antioxidants (superoxide dismutase, ascorbate peroxidase and glutathione cycling), growth and pigment pools. Changes in UVR sensitivity during photoacclimation were monitored by measuring UVR-induced inhibition of carbon assimilation under standardized UV conditions using an indoor solar simulator. Both species showed immediate antioxidant responses due to their transfer to the outdoor conditions. Furthermore, upon outdoor exposure, carbon assimilation and growth rates were reduced in both species compared with initial conditions; however, these effects were most pronounced in D. tertiolecta . Outdoor UV exposure did not alter antioxidant levels when compared with PAR-only controls in both species. In contrast, growth was significantly affected in the static UVR cultures, concurrent with significantly enhanced UVR resistance. We conclude that antioxidants play a minor role in the reinforcement of natural UVR resistance in T. weissflogii and D. tertiolecta .     

Solar PAR and UV radiation affects the physiology and morphology of the cyanobacterium Anabaena sp. PCC 7120

Solar UV radiation (280-400 nm) may affect morphology of cyanobacteria, however, little has been evidenced on this aspect while their physiological responses were examined. We investigated the impacts of solar PAR and UVR on the growth, photosynthetic performance and morphology of the cyanobacterium Anabaena sp. PCC7120 while it was grown under three different solar radiation treatments: exposures to (a) constant low PAR (photosynthetic active radiation, 400-700 nm), (b) natural levels of solar radiation with and (c) without UV radiation (290-400 nm). When the cells were exposed to solar PAR or PAR+UVR, the photochemical efficiency was reduced by about 40% and 90%, respectively, on day one and recovered faster under the treatment without UVR over the following days. Solar UVR inhibited the growth up to 40%, reduced trichome length by up to 49% and depressed the differentiation of heterocysts. Negligible concentrations of UV-absorbing compounds were found even in the presence of UVR. During the first 2 d of exposure to natural levels of PAR, carotenoid concentrations increased but no prolonged increase was evident. Heterocyst formation was enhanced under elevated PAR levels that stimulated quantum yield and growth after an initial inhibition. Higher concentrations of carotenoids and a twofold increase in the carotenoid to chlorophyll a ratio provided protection from the high levels of solar PAR. Under radiation treatments with UVR the relatively greater decrease in chlorophyll a concentrations compared with the increase in carotenoids was responsible for the higher carotenoid: chlorophyll a ratio. Heterocyst formation was disrupted in the presence of solar UVR. However, the longer term impact of heterocyst disruption to the survival of Anabaena sp. requires further study.     

Variable fluorescence parameters in the filamentous Patagonian rhodophytes, Callithamnion gaudichaudii and Ceramium sp. under solar radiation

The filamentous rhodophytes Callithamnion gaudichaudi Agardh and Ceramium sp. were utilized to study the effects of solar radiation (PAR, 400-700 nm, UV-B, 280-315 nm and UV-A, 315-400 nm) on the photosynthetic performance in situ in Patagonia waters (Argentina). A pulse amplitude modulated (PAM) fluorometer was used to determine the fluorescence parameters. The two species grew in different habitats in the eulittoral: Ceramium sp. was found only in rock pools while C. gaudichaudii grew on exposed rocks and fell dry during low tide. Both species differed in their fluorescence parameters and their sensitivity to solar radiation exposure. The photosynthetic quantum yield had its lowest values at noon, but it recovered in the afternoon/evening hours, when irradiances were lower. PAR (irradiance of about 400 W m(-2) at noon) was responsible for most of the decrease in the yield on clear days, especially in Ceramium sp., but UVR (280-400 nm) also accounted for a significant decrease. Fluence rate response curves indicated that both species were adapted to low fluence rates and showed a pronounced non-photochemical quenching at intermediate and higher irradiances. Both species showed a rapid adaptation during measurement of fast induction kinetics but differed significantly in their fluorescence components. All photosynthetic pigments were bleached after 8 h exposure to solar radiation over a full day. Strong absorption in the UV-A range, most likely due to mycosporine-like amino acids, was detected in both strains. The pronounced sensitivity to solar radiation in situ and the recovery capacity of these two filamentous Rhodophyte species, as well as the presence of protective compounds, suggests that these algae have the ability to adapt to the relatively high radiation levels and changes in irradiance found in the Patagonia waters.     

Dynamics of potentially protective compounds in Rhodophyta species from Patagonia (Argentina) exposed to solar radiation

The impact of solar radiation upon potentially protective compounds (i.e., UV-absorbing compounds and carotenoids) was assessed in four Rhodophyte species from Patagonia (i.e., Ceramium sp. Lyngbye, Corallina officinalis Linnaeus, Callithamnion gaudichaudii Agardh and Porphyra columbina Montagne) during short-term (i.e., 46 h) experiments. Algae were exposed to solar radiation under two treatments (PAR only: 400-700 nm, and PAR+UVR: 280-700 nm) and sub-samples were taken every 3 h (or longer periods at night) to determine the spectral absorption characteristics and concentration of UV-absorbing compounds, carotenoids and photosynthetic pigments. Except for C. gaudichaudii which displayed a decrease in chl-a concentration throughout the experiment, photosynthetic pigments had small variations in all species. UV-absorbing compounds concentration had species-specific responses: Ceramium sp. was the only species in which UV-absorbing compounds concentration varied as a function of solar irradiance, with maximum values around local noon. In C. officinalis and P. columbina UV-absorbing compounds concentration increased as compared to that of chl-a; in Ceramium sp. and C. gaudichaudii, however, there was no relationship between UV-absorbing compounds content and chl-a concentration. Carotenoids, on the other hand, did co-vary with chl-a in all species. Our data suggest that, with the exception of C. gaudichaudii, the differential responses of UV-absorbing compounds concentrations are more associated to the previous light history of the algae (i.e., in turn due to their position in the intertidal zone) rather than to the radiation treatment imposed to the samples. Based on our results, the variable impact of solar radiation upon productivity (and eventually biodiversity) of macroalgae from Patagonia might consequently differentially affect higher trophic levels of the aquatic food web.     

Phlorotannin production and lipid oxidation as a potential protective function against high photosynthetically active and UV radiation in gametophytes of Alaria esculenta (Alariales, Phaeophyceae)

Radiation damage can inter alia result in lipid peroxidation of macroalgal cell membranes. To prevent photo-oxidation within the cells, photoprotective substances such as phlorotannins are synthesized. In the present study, changes in total fatty acids (FA), FA composition and intra/extracellular phlorotannin contents were determined by gas chromatography and the Folin-Ciocalteu method to investigate the photoprotective potential of phlorotannins to prevent lipid peroxidation. Alaria esculenta juveniles (Phaeophyceae) were exposed over 20 days to high/low photosynthetically active radiation (PAR) in combination with UV radiation (UVR) in the treatments: PAB (low/high PAR + UV-B + UV-A), PA (low/high PAR + UV-A) or low/high PAR only. While extracellular phlorotannins increased after 10 days, intracellular phlorotannins increased with exposure time and PA and decreased under PAB. Interactive effects of time:radiation wavebands, time:PAR dose as well as radiation wavebands:PAR dose were observed. Low FA contents were detected in the PA and PAB treatments; interactive effects were observed between time:high PAR and PAB:high PAR. Total FA contents were correlated to extra/intracellular phlorotannin contents. Our results suggest that phlorotannins might play a role in intra/extracellular protection by absorption and oxidation processes. Changes in FA content/composition upon UVR and high PAR might be considered as an adaptive mechanism of the A. esculenta juveniles subjected to variations in solar irradiance.     

Effects of nitrogen source and UV radiation on the growth, chlorophyll fluorescence and fatty acid composition of Phaeodactylum tricornutum and Chaetoceros muelleri (Bacillariophyceae)

Cultures of the marine diatoms Phaeodactylum tricornutum and Chaetoceros muelleri were grown in f/2 medium supplied with either nitrate (N-Nt), ammonium (N-Am) or urea (N-Ur) as the nitrogen (N) source at the same final N concentration (0.88 mM). Exponential growth phase cultures of the two diatoms were exposed to four different light regimes for 2 days: (UVAR) PAR (60 micromol quanta m-2 s-1) plus 8.22 W m-2 (unweighted) UVAR; (high UVBR) PAR (60 micromol quanta m-2 s-1) plus 1.04 W m-2 (unweighted) UVBR plus 13.73 W m-2 (unweighted) UVAR; (low UVBR) PAR (60 micromol quanta m-2 s-1) plus 0.19 W m-2 (unweighted) UVBR plus 2.76 W m-2 (unweighted) UVAR and (PAR) PAR (60 micromol quanta m-2 s-1) alone (control). No significant effects of N source on the growth rates of the two diatoms were detected. The maximum effective quantum yield of PSII, PhiPSIIe-max, and the initial slope of the light curve, alpha, of P. tricornutum and C. muelleri were all inhibited, whereas Ik was somewhat increased, as a consequence of 2 days of exposure to all the UVR treatments. Multiple factor ANOVA revealed that all the major fatty acids, both in P. tricornutum and C. muelleri, were influenced more strongly by N source than by UVR. The composition of saturated fatty acids (SFA), monounsaturated fatty acids (MUFA) and polyunsaturated fatty acids (PUFA) in P. tricornutum and C. muelleri exhibited almost the same pattern of variation with N source and UVR. The maximum value of SFA was found in the N-Am treatment, that of MUFA in the N-Nt treatment and for PUFA in the N-Ur treatment irrespective of the UV radiation. On the other hand, the impact of UVR resulted in an increase of PUFA and a reduction of SFA both in P. tricornutum and C. muelleri under all N sources.     

Differential Responses of Growth and Photochemical Performance of Marine Diatoms to Ocean Warming and High Light Irradiance

Diatoms have relatively high biomass in mid- to high-latitude oceans, which is also the most sensitive region to climate change. Photoautotrophs are thus predicted to become exposed to both higher temperatures and increased solar irradiance. In this study, we examined the consequences of such changes for the growth and photo-physiology of two diatoms by mimicking the scenarios that correspond to present day and that predicted for the end of this century. Elevated light induced higher rates of damage to photosystem II (PSII) that significantly reduced photochemical yields of both diatoms. Treatments including UV radiation induced ~ 50% inhibition of PSII under present PAR levels. Generally, warming alleviated UVR inhibition, resulting in higher photochemical yields, and faster recovery during dim light exposure. Therefore, concurrent increase of irradiance and temperature mitigated UV inhibition of PSII by 8–15%. The growth was stimulated by warming under PAR treatment, while less stimulation, or even decreased growth rates were found under the PAR + UVR treatment. Results suggest that ocean warming could fully offset the inhibition of high light on PSII. However, under the latter higher UVR stress scenario, the energetic expenditure required by the diatoms to repair damage could lead to their lower overall growth in future oceans.     

Interaction of UV radiation and inorganic carbon supply in the inhibition of photosynthesis: spectral and temporal responses of two marine picoplankters

The effect of ultraviolet radiation (UVR) on inhibition of photosynthesis was studied in two species of marine picoplankton with different carbon concentration mechanisms: Nannochloropsis gaditana Lubian possesses a bicarbonate uptake system and Nannochloris atomus Butcher a CO2 active transport system. Biological weighting functions (BWFs) for inhibition of photosynthesis by UVR and photosynthesis vs irradiance (PI) curves for photosynthetically active radiation (PAR) were estimated for both species grown with an enriched CO2 supply (high dissolved inorganic carbon [DIC]: 1% CO2 in air) and in atmospheric CO2 levels (low DIC: 0.03% CO2). The response to UVR and PAR exposures was different in each species depending on the DIC treatment. Under PAR exposure, rates of maximum photosynthesis were similar between treatments in N. gaditana. However, the cultures growing in high DIC had lower sensitivity to UVR than the low DIC cultures. In contrast, N. atomus had higher rates of photosynthesis under PAR exposure with high DIC, but the BWFs were not significantly different between treatments. The results suggest that one or more processes in N. gaditana associated with HCO3- transport are target(s) for UV photodamage because there was relatively less UV inhibition of the high DIC-grown cultures in which inorganic carbon fixation is supplied by passive CO2 diffusion. Time courses of photochemical efficiency in PAR, during UV exposure and during subsequent recovery in PAR, were determined using a pulse amplitude modulated fluorometer. The results were consistent with the BWFs. In all time courses, a steady state was obtained after an initial decrease, consistent with a dynamic balance between damage and repair as found for other phytoplankton. However, the relationship of response to exposure showed a steep decline in activity that is consistent with a constant rate of repair. A novel feature of a model developed from a constant repair rate is an explicit threshold for photosynthetic response to UV.     

Hydrogen peroxide formation and decay in iron-rich geothermal waters: the relative roles of abiotic and biotic mechanisms

Hydrogen peroxide (H2O2) is widely distributed in surface waters where the primary photochemical formation pathway involves the interaction between dissolved organic carbon (DOC) and ultraviolet radiation (UVR). In laboratory studies using iron-rich water from Yellow-stone's Chocolate Pots spring, H2O2 formation depended on sample treatment (unfiltered, < 0.2 micron filtered, autoclaved) prior to irradiation, suggesting several formation pathways. Similar H2O2 formation in filtered and unfiltered water indicates that it is primarily soluble material that is responsible for H2O2 formation. H2O2 formation with soluble material probably includes only photochemical reactions with DOC and/or metals. Greater H2O2 formation in unfiltered and filtered water than in autoclaved water suggests that the agent(s) involved in H2O2 formation is (are) not stable at high temperatures and pressures and degrade to nonphotoreactive species. Such unstable agents may include DOC and/or dissolved complexes of iron or other metals. UVR absorbance occurs across the UV spectrum and, though slightly greater in the UVA range (320-400 nm), is similar to that of other surface waters. Increased UVR absorbance after autoclaving suggested degradation or alteration of some components, which in turn affected H2O2 formation. The spectral region used for irradiation affected net formation and yield. H2O2 formation in water irradiated with UVA radiation was 2.5-3 times that formed in water irradiated with UVB radiation (280-320 nm) in experiments using artificial light sources. Apparent quantum yields comparable to those reported by others could not be calculated because the instrumental designs are not the same. However, approximate quantum yields were calculated for these experiments but should be viewed with caution. Quantum yields were higher in these experiments (0.0040 mol H2O2 per mol photon at 310 nm and 0.0012 mol H2O2 per mol photon at 350 nm) than values reported by other researchers (< 0.0007 mol H2O2 per mol photon at 300 nm and < 0.0005 H2O2 per mol photon at 340 nm; [Scully, N. M., D. R. S. Lean, D. J. McQueen and W. J. Cooper (1996) Limnol. Oceanogr. 41, 540-548]). In natural solar source experiments, H2O2 formation was greater in experiments with UVA and photosynthetically active radiation (PAR; 400-700 nm) than with PAR alone or with UVB, UVA and PAR. However, H2O2 capacity (nM H2O2 W-1 h-1 m2) was greatest with UVB radiation and lowest with PAR radiation. Source regions could not be studied separately. Dark decay of H2O2 occurred via two mechanisms. The main mechanism responsible for H2O2 decay involved particulate matter (probably microorganisms), whereas a secondary mechanism involved soluble matter (i.e. DOC, metal ions and other dissolved species involved in Fenton reactions).     

Solar UV Irradiances Modulate Effects of Ocean Acidification on the Coccolithophorid Emiliania huxleyi

Emiliania huxleyi, the most abundant coccolithophorid in the oceans, is naturally exposed to solar UV radiation (UVR, 280–400 nm) in addition to photosynthetically active radiation (PAR). We investigated the physiological responses of E. huxleyi to the present day and elevated CO₂ (390 vs 1000 μatm; with pH_NBS 8.20 vs 7.86) under indoor constant PAR and fluctuating solar radiation with or without UVR. Enrichment of CO₂ stimulated the production rate of particulate organic carbon (POC) under constant PAR, but led to unchanged POC production under incident fluctuating solar radiation. The production rates of particulate inorganic carbon (PIC) as well as PIC/POC ratios were reduced under the elevated CO₂, ocean acidification (OA) condition, regardless of PAR levels, and the presence of UVR. However, moderate levels of UVR increased PIC production rates and PIC/POC ratios. OA treatment interacted with UVR to influence the alga's physiological performance, leading to reduced specific growth rate in the presence of UVA (315–400 nm) and decreased quantum yield, along with enhanced nonphotochemical quenching, with addition of UVB (280–315 nm). The results clearly indicate that UV radiation needs to be invoked as a key stressor when considering the impacts of ocean acidification on E. huxleyi.     

Impact of solar ultraviolet radiation on marine phytoplankton of Patagonia, Argentina

Patagonia area is located in close proximity to the Antarctic ozone "hole" and thus receives enhanced ultraviolet B (UV-B) radiation (280-315 nm) in addition to the normal levels of ultraviolet A (UV-A; 315-400 nm) and photosynthetically available radiation (PAR; 400-700 nm). In marine ecosystems of Patagonia, normal ultraviolet radiation (UVR) levels affect phytoplankton assemblages during the three phases of the annual succession: (1) prebloom season (late summer-fall), (2) bloom season (winter-early spring) and (3) postbloom season (late spring-summer). Small-size cells characterize the pre- and postbloom communities, which have a relatively high photosynthetic inhibition because of high UVR levels during those seasons. During the bloom, characterized by microplankton diatoms, photosynthetic inhibition is low because of the low UVR levels reaching the earth's surface during winter; this community, however, is more sensitive to UV-B when inhibition is normalized by irradiance (i.e. biological weighting functions). In situ studies have shown that UVR significantly affects not only photosynthesis but also the DNA molecule, but these negative effects are rapidly reduced in the water column because of the differential attenuation of solar radiation. UVR also affects photosynthesis versus irradiance (P vs E) parameters of some natural phytoplankton assemblages (i.e. during the pre- but not during the postbloom season). However, there is a significant temporal variability of P vs E parameters, which are influenced by the nutrient status of cells and taxonomic composition; taxonomic composition is in turn associated with the stratification conditions (e.g. wind speed and duration). In Patagonia, wind speed is one of the most important variables that conditions the development of the winter bloom by regulating the depth of the upper mixed layer (UML) and hence the mean irradiance received by cells. Studies on the interactive effects of UVR and mixing show that responses of phytoplankton vary according to the taxonomic composition and cell structure of assemblages; therefore cells use UVR if >90% of the euphotic zone is being mixed. In fact, cell size plays a very important role when estimating the impact of UVR on phytoplankton, with large cells being more sensitive when determining photosynthesis inhibition, whereas small cells are more sensitive to DNA damage. Finally, in long-term experiments, it was determined that UVR can shape the diatom community structure in some assemblages of coastal waters, but it is virtually unknown how these changes affect the trophodynamics of marine systems. Future studies should consider the combined effects of UVR on both phytoplankton and grazers to establish potential changes in biodiversity of the area.     

Motility of Daphnia spinulata as affected by solar radiation throughout an annual cycle in mid-latitudes of Patagonia

During an annual cycle, Daphnia spinulata collected from a plankton community of Patagonia was exposed outdoors to assess the impact of recently received solar radiation on motility (i.e. swimming speed and gravitaxis). Individual values of these parameters were obtained by video recordings and image analysis at different time intervals during the day. Initial swimming speed varied throughout the year, and changes in speed during exposure were not significantly affected by any waveband used in our experimental design (i.e. PAB, 280-700 nm; PA, 320-700 nm; and P, 400-700 nm). Overall, most of the individuals swam downwards, regardless of the radiation treatment imposed to the samples. We found that multifactor interactions (i.e. not a single parameter explained more than 40% of the observed variability) explained most of our observations on motility parameters. These factors include not only solar radiation, but other physical (underwater radiation field and wind intensity) and biological parameters (food availability, presence of predators and congeners). Our findings indicate that the plankton dynamics in the study site is likely to be governed by a sum of factors which must be taken into account when considering solar radiation effects on aquatic ecosystems.     

Daily variation in cellular content of UV-absorbing compounds mycosporine-like amino acids in the marine dinoflagellate Scrippsiella sweeneyae

Sudden exposure experiments to high PAR (photosynthetically available radiation) or high PAR+UVR (ultraviolet radiation) were conducted for the marine dinoflagellate Scrippsiella sweeneyae acclimated to either low PAR or high PAR to determine the induction of cellular mycosporine-like amino acid (MAA) in relation to photosynthesis status. When the exposure to high PAR (30.8 Wm(-2)) was provided at different time in the light period for S. sweeneyae acclimated to low PAR (7.7 Wm(-2)) which suppressed photosynthesis, S. sweeneyae could enhance the induction of MAA but it only occurred in the first half of the light period. When UVR exposure was provided for the culture acclimated to high PAR which enhanced photosynthesis, cellular MAA content did not increase during the entire light period, but displayed daily variation similar to the control for two and half days. Daily variation of cellular MAA content did not synchronized with that of cell volume and cellular chlorophyll a content. The individual MAAs also revealed similar daily variations with different phase, which increased for a few hours in the beginning of the light period, except for cellular palythine content. Thus the total cellular MAA content revealed daily variation with changing the relative composition within a few hours. As one of the biological protective strategies against harmful UVR in sunlight, the daily vertical migration in the bloom forming dinoflagellates might be accompanied by the daily variation of cellular MAA content for a photosynthesis at daytime.     

Variability of bio-optical factors influencing the seasonal attenuation of ultraviolet radiation in temperate coastal waters of Japan

The study identifies the relative contribution of various bio-optical factors to the total attenuation of ultraviolet radiation (UVR) wavelengths and photosynthetically active radiation (PAR) in temperate coastal waters of Japan by surveying the physical properties of the water column, UVR and PAR penetration, and the absorption characteristics of dissolved and particulate material. Spectral absorbance properties of pigment (aph), detritus (ad) and chromophoric dissolved organic material (aCDOM) displayed both seasonal and wavelength specific variability. On an annual basis, absorbance by aCDOM was the highest absorbing fraction (47-59%) for the UVR wavelengths measured (305, 320, 340 and 380 nm) but decreased (32%) at 450 nm. Contribution of pigments to total absorbance was highest (40-60%) during a spring bloom for both UVR and PAR. A large variability (C.V. > 42%) for annual average attenuation coefficients (Kd[lambda]) at respective wavelengths observed suggests that the spectral composition of the water column changes throughout the year in this region. A significant relationship was observed between Kd(lambda) and aCDOM at 305, 320, 340 and 380 nm only (P < 0.01) but not for 450 nm (PAR) indicating the role of CDOM in regulating variations in Kd(lambda), particularly in the UVR range. The slope S, obtained from a natural-log plot of the absorption coefficient of CDOM against wavelength, ranged between 0.014 and 0.036 nm-1 annually (average = 0.020 +/- 0.007, C.V. = 35%) and suggests seasonal changes in the origin of CDOM between terrestrial (low S) and biogenous (high S) CDOM.     

Photoacclimation to long-term ultraviolet radiation exposure of natural sub-Antarctic phytoplankton communities: Fixed surface incubations versus mixed mesocosms

Solar UVB radiation (280-320 nm) is known to have detrimental effects on marine phytoplankton. Associated with the seasonal ozone hole in Antarctica, stratospheric ozone depletion occasionally influences the sub-Antarctic (Beagle Channel, Argentina) region, enhancing levels of UVB. The primary objective of this work was to study the effects of several (i.e. 6-10) days of exposure to UVB on the taxonomic composition and photosynthetic inhibition of local phytoplankton communities. For different light treatments, fixed-depth incubations placed in an outdoors water tank were compared with incubations in 1900 L mesocosms, where vertical mixing was present. Phytoplankton growth was inhibited by UV radiation (UVR) in fixed-depth experiments but not in the mixed mesocosms. Under fixed and mixed conditions alike, photosynthesis was significantly inhibited by UVB at the beginning of the experiment but no longer after several days of exposure, suggesting that cells had acclimated to radiation conditions. There was a change in species composition in response to UVR exposure in both experiments, which likely explained acclimation. In the community exposed to fixed conditions this change was from a phytoflagellate-dominated assemblage to a community with high relative abundance of diatoms after 6 days of exposure. UVA was responsible for most of the observed growth inhibition; however, the reduction in photosynthesis was produced by UVB. The reasons behind this variability in responses to UVR are associated with species-specific sensitivity and acclimation, and the previous light history of cells. In the community exposed in mesocosms, an assemblage codominated by phytoflagellates and diatoms was observed at the beginning of the experiments. After 10 days of exposure, green algae (Eutreptiella sp.) had increased, and phytoflagellates were the dominant group. The synthesis of mycosporine-like amino acids (MAAs), antioxidant enzymes and photosynthetic antenna pigments, in relation to repair and protection processes, may explain the reduced inhibition of both growth and photosynthesis that was observed in the phytoplankton community after several days of exposure. For environments such as the Beagle Channel seasonally exposed to the ozone hole, the results obtained from the fixed-depth experiments show that species can cope with UVR by means of MAA synthesis, while mixing would primarily promote a change in species composition and defense strategies.