UV- and salinity-induced oxidative effects in the marine diatom <Emphasis Type="Italic">Cylindrotheca closterium</Emphasis> during simulated emersion

The diatom Cylindrotheca closterium was exposed to transient light- and osmotic conditions as occur during its tidal emersion. The objective was to analyze how this simulated emersion contributes to the production of active oxygen species (AOS) and via this, to oxidative cell damage. Light- and salinity conditions were varied in factorial combination: low light (no UVB) or high light (unweighted UVB-dose rates of respectively 0.01; 0.07; 0.24; 1.03 W m⁻²) at normal (30 psu) or high salinity (60 psu). UVB (0.01–0.24 W m⁻²) and high salinity had a significant, negative effect on the photosynthetic efficiencies ΔF/F _m’ (steady-state quantum yield) and F _v/F _m (maximum yield). UVB at 1.03 W m⁻² (15 kJ m⁻² d⁻¹) almost arrested electron transport. At ecologically relevant UVB levels, i.e. below 0.24 W m⁻² (≈3.4 kJ m⁻² d⁻¹) with UVB:PAR<0.4:100 (PAR photosynthetically active radiation) only dynamic photoinhibition was observed (protection via heat dissipation). Non-photochemical quenching was positively correlated with the de-epoxidation of diadinoxanthin (DD) to diatoxanthin (DT). A decreasing ratio DT/(DD+DT) after 4 h of UVB at >0.07 W m⁻² and at 60 psu indicated a reversal of the diatom xanthophyll cycle (diminished photoprotection) which may be caused by an enhanced AOS production. Oxidative stress and -damage to C. closterium cells were assessed applying fluorescent indicator dyes, via confocal microscopy and quantitative image analysis. AOS production rates (cellular DCF fluorescence) were stimulated by UV, and were ~50% higher at 60 psu. AOS production decreased with an increasing pre-exposure (0–4 h) to normal UVB (0.24 W m⁻²), which indicated a stimulation of the antioxidative defence. Non-protein thiols (indicator CMF) and glutathione pools (HPLC-analyzed) decreased with UVB-dose rates (0.01–0.24 W m⁻²), most likely due to AOS-mediated thiol oxidation. Hypersalinity (60 psu) and UVB (0.01–0.24 W m⁻²) caused membrane depolarization (dye DIBAC₄(3)) and phospholipid hydrolysis (phospholipase A₂ dye: bis-BODIPY FL-C₁₁-PC). AOS production may have diminished the membrane polarity, and peroxidized the membrane lipids (HPLC-analyzed malondialdehyde) which enhanced PLA₂ activity. The dyes indicated an increased oxidative (lipid) damage at a 15% inhibition of photosynthesis in this diatom, at UVB levels and salinities that can be expected in situ during its periodic tidal emersion.     

Diurnal changes in photosynthesis of Antarctic fast ice algal communities determined by pulse amplitude modulation fluorometry

The aim of this project was to determine both the diurnal changes in photosynthetic activity of Antarctic sea ice algae and also the protective mechanisms they use to mitigate the effects of in situ UV radiation. Changes in the diurnal photosynthetic parameters of fast ice algal communities at McMurdo Sound were measured in situ, using a custom designed monitoring pulse amplitude modulation fluorometer. The sea ice microalgae were able to adapt rapidly to either increasing or decreasing ambient irradiances. ΔF/F_m' values were between 0.2 and 0.51, while E_k varied between 2.1 and 18 μmol photons m^-2 s^-1. ΔF/F_m', E_k, and relative electron transfer rate (rETR) all varied sequentially over the course of a day. rETR and E_k were highest at midday at the highest irradiances, when there was apparent midday down regulation of photosynthesis, while ΔF/F_m' was highest at midnight. The effects of natural UV radiation on sea ice were examined, but it was not possible to detect the effect of either UVB or UVA and UVB on photosynthesis. This was considered to be largely because of the large spatial and temporal heterogeneity of the under ice community, changing irradiances throughout the day and the relatively small change caused by UV.     

Irradiance-induced changes in the photobiology of <Emphasis Type="Italic">Halophila johnsonii</Emphasis>

The endangered seagrass Halophila johnsonii Eiseman, exhibits high-light adapted photophysiology consistent with its distribution in intertidal and shallow subtidal (0–3 m) coastal-lagoon habitats along 200 km of southeastern Florida. To examine the short-term responses of this seagrass to three controlled-irradiance treatments (PAR + UVA + UVB [full spectrum], PAR + UVA, and PAR only), greenhouse-acclimated plants were transferred to outdoor mesocosms during July–August 2002. Chlorophyll fluorescence, UV fluorescence, and samples for pigment extraction were collected in the greenhouse, prior to moving the plants outside and on days 1, 2, 3, 4, 6, 10, and 21 of the 24-day experiment. Typical of sun-adapted plants, effective quantum yields measured by pulse-amplitude modulated (PAM) fluorometry were relatively low in all treatments, ranging from 0.46 ± 0.09 (PAR only) to 0.58 ± 0.08 (PAR + UVA + UVB). In the PAR only treatments, there were strong effects on days 1 and 4, presumably because the irradiance in the greenhouse not only lacked all λ<400 nm, but also had low irradiance maxima (∼700 μmol photons m⁻² s⁻¹, compared with ∼1,500 μmol photons m⁻² s⁻¹ outside at midday). There were few treatment differences between PAR only and PAR + UVA treatments indicating little effect of UVA radiation on this species. Differences in effective quantum yields and relative electron transport rates between the PAR only and PAR + UVA + UVB treatments on day 4 indicated rapid acclimation to UVB radiation. Tissues of H. johnsonii contained compounds that absorbed strongly in the UV, with a λ_max at ∼345 nm (depending on the extraction solvent). Absorption peak maxima and minima changed over the course of the experiment but there were no significant light-treatment differences in any pigment parameters. Percent UV shield values, measured using a newly developed UVA PAM fluorometer, were highest the day after plants were transferred from the greenhouse to the outdoor mesocosms and declined significantly to pretreatment levels in all treatments by day 21. Percent UV shield exhibited a significant positive relationship with UV-absorbing pigment (UVP) absorbance, however, the absence of treatment effects suggests that the wavelengths inducing pigment synthesis must lie between 400 and 700 nm (PAR). The results indicate that H. johnsonii rapidly acclimates to high UVB and PAR which may largely explain its distribution in intertidal and shallow subtidal areas.     

Interactive effects of UV radiation and enhanced temperature on photosynthesis, phlorotannin induction and antioxidant activities of two sub-Antarctic brown algae

Lessonia nigrescens and Durvillaea antarctica, two large sub-Antarctic brown algae from the southern Chilean coast, were exposed to solar UV radiation in an outdoor system during a summer day (for 11 h) as well as to artificial UV radiation under controlled laboratory conditions at two temperatures (15 and 20 °C) for 72 h. Chlorophyll a fluorescence–based photoinhibition of photosynthesis was measured during the outdoor exposure, while electron transport rates, lipid peroxidation, antioxidant activity and content of phlorotannins were determined at different time intervals during the laboratory exposure. Under natural solar irradiances in summer, both species displayed well-developed dynamic photoinhibition: F _v/F _m values decreased by 70 % at noon coinciding with the levels of PAR >1,500 μmol m^?2 s^?1 and UV-B radiation >1 W m^?2 and recovered substantially in the afternoon. In treatments including UV radiation, recovery in D. antarctica started already during the highest irradiances at noon. The results from laboratory exposures revealed that (a) elevated temperature of 20 °C exacerbated the detrimental effects of UV radiation on photochemical parameters (F _v/F _m and ETR); (b) peroxidative damage measured as MDA formation occurred rapidly and was strongly correlated with the decrease in F _v/F _m, especially at elevated temperature of 20 °C; (c) the antioxidant activity and increases in soluble phlorotannins were positively correlated mainly in response to UV radiation; (d) phlorotannins were rapidly induced but strongly impaired at 20 °C. In general, short-term (2–6 h) exposures to enhanced UV radiation and temperature were effective to activate the photochemical and biochemical defenses against oxidative stress, and they continued operative during 72 h, a time span clearly exceeding the tidal or diurnal period. Furthermore, when algae were exposed to dim light and control temperature of 15 °C for 6 h, F _v/F _m increased and lipid peroxidation decreased, indicating consistently that algae retained their ability for recovery. D. antarctica was the most sensitive species to elevated temperature for prolonged periods in the laboratory. Although no conclusive evidence for the effect of the buoyancy of fronds was found, the interspecific discrepancies in thermo-sensitivity in the UV responses found in this study are consistent with various ecological and biogeographical differences described for these species.     

Distribution patterns of meiofauna associated with a sublittoral<Emphasis Type="Italic"> Laminaria</Emphasis> bed in the Cantabrian Sea (north-eastern Atlantic)

This study deals with meiofauna associated with a sublittoral population of the kelp Laminaria ochroleuca located on the northern coast of Spain. By sampling once a year over a 4-year period, we examined patterns of faunal distribution as a function of some environmental factors at the meso-scale level (depth, and exposure to waves and surge). We also examined the relationship between L. ochroleuca abundance (as dry weight biomass and number of plants per sampling quadrat) and abundance and diversity of meiofauna. Finally, we investigated patterns of within-plant distribution (algal frond vs. algal holdfast), using also the meiofauna of the adjacent bottom as a referent to estimate the level of "phytal dependence" of the meiofauna collected on L. ochroleuca. We found that the bulk of permanent meiofauna consisted of nematodes, copepods, mites, polychaetes, tanaids and ostracods, with copepods being predominant on the fronds of the alga and nematodes in the holdfasts. The temporary meiofauna consisted of juvenile amphipods, bivalves and gastropods, together with barnacle nauplii and cyprids. Abundance and major composition of meiofaunal taxa were unrelated to both depth and hydrodynamic exposure of the sampling quadrats. However, we detected significant qualitative and quantitative faunal differences as a function of microhabitat. All meiofaunal groups were more abundant in holdfast samples than in frond and bottom samples. The gross taxonomic composition of meiofauna in bottom samples was similar to that in holdfast samples, but substantially different from that of meiofauna associated with the fronds. The L. ochroleuca holdfasts, in which dense aggregations of meiofauna can occur, appear to function as ecotone between phytal and rocky-bottom microhabitats. All together, our results suggest that the distribution of meiofauna within the Laminaria bed is mostly affected by factors operating at the microhabitat level rather than the meso-scale level.Communicated by L. Hagerman, Helsingør     

A method for the assessment of light-induced oxidative stress in embryos of fucoid algae via confocal laserscan microscopy

A method was developed for measurement of active oxygen production in embryonic stages of the brown seaweed Fucus spiralis, using the label CM-DCFH-DA. Active oxygen species convert the label into the green fluorescent CM-DCF (exc/em 488/530 nm) that is detected via confocal laserscan microscopy and quantitative image analysis. Loading of the label did not harm the embryos; loading efficiency was age-independent, and the esterases needed for conversion to CM-DCFH were not inhibited by the effective UV dose (2 W m⁻²) applied in the experiments. After correction for daily variation of the laser power, and calibration with DCF standard solutions, this automated analysis of confocal images rendered active oxygen concentrations in fucoid embryos (μM DCF). An experiment was designed for the assessment of active oxygen production following irradiance stress in the light-sensitive embryos. Dim-light-acclimated, 1-, 2- and 4-day-old embryos were transferred for 60 min to defined high-light conditions (4π-irradiance 300 μmol photons m⁻² s⁻¹), optionally without UV radiation, including UVA, or including UVA plus UVB. PSII yield measurements (PAM fluorometer) were carried out in order to assess the degree of photoinhibition under these light conditions. The imposed light stress initially caused a rapid decrease of the PSII yields (Φ_P). With increasing embryo age, minimum Φ_P values attained under light stress remained higher. Consequently, electron transport rates (ETR) would increase with embryo age, i.e., with the development of their photosynthetic apparatus. Active oxygen production increased with ETR, and when UVB was included, relatively greater amounts of active oxygen were produced. A slow, second-phase decrease of Φ_P under light stress that was proportional to active oxygen production indicated that some photooxidative damage was caused, in particular during UVB exposure. Recovery from light stress was a rapid process in the absence of UVB; in such cases Φ_P was almost restored to the initial values within 60 min. The relative state of recovery of Φ_P was correlated with both the effective UV dose and active oxygen production rate (DCF). Recovery was slowest in embryos exposed for 60 min to an experimental UVB dose, which was representative of a situation at low tide, on a sunny day. The results suggest that active oxygen may cause an in situ inhibition of growth of the earliest life stages of F. spiralis. Received: 26 January 2000 / Accepted: 4 September 2000     

Effects of ultraviolet radiation on different life cycle stages of the south Pacific kelps, Lessonia nigrescens and Lessonia trabeculata (Laminariales, Phaeophyceae)

The effects of exposure to ultraviolet radiation (UVR), 280–400 nm, in different life histories and development stages of the kelps, Lessonia nigrescens and L. trabeculata, collected in the south-east Pacific coast (30°S) were evaluated in the laboratory. Germination and viability (motile zoospores, settled spores), diameter of the primary cell of the gametophytes, percentage of female gametophytes, fertility and sporophytes production were measured after exposure to three radiation treatments (PAR; PAR + UVA; PAR + UVA + UVB). The effects of UVR in young sporophytes (diploid stage) were evaluated as changes in maximal quantum yield of chlorophyll fluorescence of photosystem II (PSII) (F _v/F _m). A significant decrease in all variables was observed for the treatment that included UVB (PAR + UVA + UVB) after 2 and 4 h of exposure, in relation to the control. The motile spores were more sensitive to UVR exposure compared to settled spores and gametophytes, suggesting that along with an increase in ontogenetic development; there is an increase in the tolerance to UVR. In addition, it was observed that early stages of the intertidal L. nigrescens were more tolerant to UVR compared to the subtidal L. trabeculata. These results allow initially to infer that UVR may be regarded as an important environmental factor influencing the upper limit of distribution of these species, mainly through its detrimental effects on the early stages of the life cycle.     

Capability of dynamic photoinhibition in Arctic macroalgae is related to their depth distribution

The capability of several macroalgal species to protect photosynthesis from excessive irradiance by dynamic photoinhibition was investigated relative to their depth distribution in summer 1995 in the Kongsfjord (79°N; 12°E, Ny Ålesund, Spitsbergen, Norway). Photoinhibition of photosynthesis was induced by exposure of algae from different water depths to a high photon fluence rate of 500?μmol?m^?2?s^?1 for 2?h. Changes in optimal quantum yield (F _v/F _m) were measured during the inhibition phase. Recovery of photosynthesis was subsequently induced by dim white light (10?μmol?m^?2?s^?1) and observed as changes in the variable fluorescence. With a newly developed mathematical model different parameters of the response kinetics of inhibition and recovery were calculated and related to the depth distribution of each algal species. It is shown that two components with slow and fast reaction kinetics, respectively, are involved in photoinhibition and recovery of photosynthesis. Their possible molecular bases are discussed. The half-life time (τ) of the inhibition and recovery phases, i.e. the time necessary to reach half maximal response, is clearly related to the depth distribution of the investigated species. Algae collected close to the water surface show a fast reaction of both photoinhibition and recovery and, hence, have a low τ. With increasing depth the reactions become slower and τ increases. τ was highest in deep water algae. Further analysis of the reaction kinetics in Laminaria saccharina shows that the relative proportion of the two kinetics involved change with the collection depth. In contrast, a significant difference in the reaction rates of both kinetics was not observed.     

Photoinhibition and recovery of the kelp Laminaria saccharina at optimal and superoptimal temperatures

Effects of high irradiance on photosynthetic characteristics were examined in sporophytes of the kelp Laminaria saccharina Lamour. from 1992 to 1994. Exposure to high irradiance (700 mol photons m^-2s^-1) for 1 h at optimal temperature (12°C) caused a 40 to 60% decline in photosynthetic efficiency (alpha), quantum yield, and the ratio of variable to maximum chlorophyll fluorescence (F_v/F_m), an indicator of Photosystem II efficiency. Although the photoinhibition effects were partly attributable to protective mechanisms, a concurrent increase in minimal fluorescence (F_o) indicated damage to Photosystem II reaction centers. The magnitude of photoinhibition was proportional to irradiance and duration; however, F_v/F_m was significantly reduced after exposure to irradiances as low as 40 to 50 mol photons m^-2s^-1 for 1 h, or to 700 mol photons m^-2s^-1 for only 5 min. In contrast, photosynthetic capacity (P_max) was affected only at much higher irradiance. Superoptimal temperatures up to 24°C did not exacerbate high-light effects. At 25°C, however, alpha and P_max were more susceptible to photoinhibition than at lower temperatures. Recovery from photoinhibition was examined by following F_v/F_m and F_o for 24 h after exposure to high light. Recovery of F_v/F_m was fastest during the first 1 to 3 h, and slowed or ceased after 6 to 12 h, while recovery of F_o was relatively constant over 12 h. Dithiothreitol, which blocks formation of energy-dissipating xanthophylls, reduced both the initial rate and extent of recovery. Chloramphenicol, which blocks chloroplast-encoded protein synthesis, had little effect on initial rates of recovery, but stopped recovery after 3 h. Thus, L. saccharina appears to rely on the xanthophyll cycle to protect the photosynthetic apparatus, and reversal of this protective mechanism causes the rapid initial recovery in F_v/F_m. Longterm recovery depends on repair of damaged reaction centers. Both the rate and extent of recovery were temperature-dependent. The initial rate was higher at 18 to 22°C than at 12°C, but the extent of recovery over 24 h declined with increasing temperature. High temperatures, therefore, appear to enhance protective mechanisms, but disrupt repair processes. L. saccharina from Long Island Sound, an ecotype adapted to low light and high temperature, showed slightly but consistently greater effects of photoinhibition than plants from the Atlantic coast of Maine, but exhibited faster recovery at superoptimal temperatures.     

Inhibition of photosynthesis by ultraviolet radiation as a function of dose and dosage rate: Results for a marine diatom

The effects of ultraviolet radiation on phytoplankton are usually described as a function of dose (J m^–2, weighted appropriately). Experiments conducted in 1988 and 1989 on a marine diatom,Thalassiosira pseudonana (Clone 3H), demonstrate that during lightlimited photosynthesis in visible radiation, the inhibition of photosynthesis by supplemental ultraviolet radiation (principally UV-B: 280 to 320 nm) is a function of irradiance (W m^–2) as well as of dose: for equal doses of UV-B, a relatively short exposure to high UV-B irradiance is more damaging to photosynthesis than a longer exposure to lower irradiance. In fact, photoinhibition by UV-B is well described as a monotonic, nonlinear function of irradiance for time scales of 0.5 to 4 h. A nitrate-limited culture was about nine times more sensitive to UV-B than was a nutrient-replete culture, but the kinetics of photoinhibition were similar. These results have some bearing on efforts to describe the effects of ultraviolet radiation on marine primary productivity. Action spectra of photoinhibition by UV can be constructed, but they should only be used to describe photoinhibition for specified time scales. Vertical profiles of relative photoinhibition must be interpreted cautiously because photoinhibition by UV-B is likely to be a function of incubation time and results must therefore be interpreted in the context of vertical mixing.     

Colonisation of kelp imitations by epiphyte and holdfast fauna; a study of mobility patterns

The kelp Laminaria hyperborea provides a habitat for a diverse invertebrate community. Invertebrate abundance is particularly high in the kelp holdfasts and associated with epiphytes on the stipes. We investigated colonisation patterns of this fauna on artificial substrata (holdfast and epiphyte mimics) exposed for short time periods in the kelp forest in two seasons, summer and late autumn. Within 7 days, 99 taxa common on natural kelp substratum colonised the artificial substrata in large numbers, demonstrating high mobility and indicating that there may be a rapid exchange of individuals and species among algae. Gastropods and amphipods were the most abundant colonisers. In summer, a specific “epiphyte fauna” could be distinguished from a “holdfast fauna”, and species associated with epiphytes were significantly more abundant on the artificial substrata. Colonisation was more rapid in summer than in late autumn, which may be related to animal population density. The high dispersal rate can facilitate the interaction with other communities and with higher trophic levels in the kelp-forest food web. Published online: 13 August 2002     

Effects of light exposure on the retention of kleptoplastic photosynthetic activity in the sacoglossan mollusc <Emphasis Type="Italic">Elysia viridis</Emphasis>

The effects of light exposure on the photosynthetic activity of kleptoplasts were studied in the sacoglossan mollusc Elysia viridis. The photosynthetic activity of ingested chloroplasts was assessed in vivo by non-destructively measuring photophysiological parameters using pulse amplitude modulation (PAM) fluorometry. Animals kept under starvation were exposed to two contrasting light conditions, 30 μmol photons m⁻² s⁻¹ (low light, LL), and 140 μmol photons m⁻² s⁻¹ (high light, HL), and changes in photosynthetic activity were monitored by measuring the maximum quantum yield of photosystem II (PSII), F _v/F _m, the minimum fluorescence, F _o, related to chlorophyll a content, and by measuring rapid light-response curves (RLC) of relative electron transport rate (rETR). RLCs were characterised by the initial slope of the curve, α_RLC, related to efficiency of light capture, and the maximum rETR level, rETR_m,RLC, determined by the carbon-fixation metabolism. Starvation induced the decrease of all photophysiological parameters. However, the retention of photosynthetic activity (number of days for F _v/F _m > 0), as well as the rate and the patterns of its decrease over time, varied markedly with light exposure. Under HL conditions, a rapid, exponential decrease was observed for F _v/F _m, α_RLC and rETR_m,RLC, F _o not showing any consistent trend of variation, and retention times ranged between 6 and 15 days. These results suggested that the retention of chloroplast functionality is limited by photoinactivation of PSII reaction center protein D1. In contrast, under LL conditions, a slower decrease in all parameters was found, with retention times varying from 15 to 57 days. F _v/F _m, α_RLC and rETR_m,RLC exhibited a bi-phasic pattern composed by a long phase of slow decrease in values followed by a rapid decline, whilst F _o decayed exponentially. These results were interpreted as resulting from lower rates of D1 photoinactivation under low light and from the gradual decrease in carbon provided by photosynthesis due to reduction of functional photosynthetic units.     

Diurnal measurements of F v/F m used to improve productivity estimates in macroalgae

The ratio of variable fluorescence to maximal fluorescence (F _v/F _m) was measured during a night and day cycle in five different macroalgae growing in the littoral zone at the Swedish west coast; the green algae Ulva lactuca, Cladophora sp. and Enteromorpha flexousa, the red alga Ceramium nodulosum and the brown alga Fucus vesiculosus. All the green algae and C. nodulosum showed both diurnal fluctuations in F _v/F _m during days of high light intensities and a significant negative correlation between F _v/F _m and photon irradiance (PI). An attempt has been made to improve calculations of macroalgal net growth based on carbon fixation (α_net) considering this diurnal change in F _v/F _m. By assuming that the negative linear correlation between F _v/F _m and the maximum quantum yield for photosynthesis (Φ) is proportional to that between F _v/F _m and α_net, it was possible to include the daily variation of α_net due to photoinhibition. To compensate for the variation of F _v/F _m, a representative value for each day was obtained by weighting all values of F _v/F _m over the diurnal period in relation to total PI. For all algal species there was a fairly good agreement between this representative value and the F _v/F _m value measured around noon. As the daily representative F _v/F _m value showed a negative linear correlation with the daily mean PI, it was possible to correct α_net for differences of daily mean PI. Received: 10 March 1997 / Accepted: 25 August 1997     

Photosynthetic performance,oxidative damage and antioxidants in <Emphasis Type="Italic">Cylindrotheca closterium</Emphasis> in response to high irradiance,UVB radiation and salinity

Cylindrotheca closterium is a common marine diatom living in intertidal environments where it can be present both in the water column and on sediments, depending on the tidal regime. In the present work this diatom was employed to investigate the responses to desiccation and to increase in PAR and UVB intensity, as occurs during emersion. Under these circumstances, the production of active oxygen species (AOS) may be enhanced resulting in an oxidative stress. Stress responses in this species were measured by exposing it to normal (30) and double salinity (60), supplying light of low or high intensity for 12 h, in the latter case either without or with moderate dose rates of UVB. Pulse amplitude modulated fluorometry was used to measure Chl a autofluorescence (F ₀), an index of photosynthetic efficiency of PSII (F _v/F _m) and the relative electron transfer rate (rETR). The oxidative stress was evaluated by analysing GSH pools and SOD activity. It was observed that at double salinity and under low light, intracellular pools of reduced glutathione (GSH) were higher than under the two conditions of high light without and with UVB at both salinities. The antioxidative defence activity of superoxide dismutase (SOD) was far higher under hypersaline conditions. The oxidative damage was evaluated as protein and lipid damage. The results showed that it expressed itself mainly through protein peroxidation: at normal salinity relative protein carbonyl content was (a) twice as high as in cells grown at double salinity, and (b) three times as high under UVB. Total unsaturated lipid contents doubled under hypersalinity conditions. The lipid peroxidation marker malondialdehyde showed the strongest response to low light and UVB at salinity value of 60. Lipid peroxide content was significantly higher at salinity of 60 compared to normal salinity and was the highest under low light and high light with UVB. The simulated emersion condition of the diatom seems to lead to the establishment of a balance between damage and repair, expressed mainly as (a) oxidative protein damage at normal salinity, in particular due to UV radiation, (b) sufficient protection by SOD activity mainly under hypersaline conditions.     

Allocation of organic material and energy to the holdfast,stipe, and fronds inPostelsia palmaeformis (Phaeophyta: Laminariales) on the California coast

Postelsia palmaeformis were collected from the lower intertidal at Pigeon Point, California, USA, in May 1987, and the proximate composition and allocation of energy to the various body components were determined. The holdfast and stipe have a proximate composition (% dry weight) of ca. 40% ash, 5.3% protein, 1% lipid, 2% soluble carbohydrate, and 55% insoluble carbohydrate. The fronds have a proximate composition of ca. 25% ash, 6.5% protein, 2% lipid, 3% soluble carbohydrate, and 65% insoluble carbohydrate. The energetic level was ca. 12 kJ g^-1 dry wt and ca. 19 kJ g^-1 ash-free dry wt. The relative proportion of three plant components varied, comprising 26, 39, and 35% wet wt and 20, 42, and 38% kJ for the holdfast, stipe, and fronds, respectively. A plant with a basal stipe diameter of 33 mm contains 114 g wet wt and 266 kJ. The maximal density found in May 1987 was 826 plants, 49 301 g wet wt, and 106 157 kJ m^-2.P. palmaeformis differs in these characteristics from another intertidal pheophyte,Durvillaea antarctica, that is found in a high-energy intertidal zone.     

Effects of ultraviolet radiation on skin structure and ultrastructure in Caspian Sea Salmon,Salmo trutta caspius,during alevin stage

Ozone reduction enhances the amount of ultraviolet (UV) radiation reaching the aquatic ecosystems, which can induce adverse effects on fish. In order to study the UV radiation effects on skin structure and ultrastructure, the alevins of Salmo trutta caspius were exposed to two different doses of UV radiation (low doses of: 60 µW cm^?2 UVC; 100 µW cm^?2 UVB; and 40 µW cm^?2 UVA and high doses of: 90 µW cm^?2 UVC; 130 µW cm^?2 UVB; and 50 µW cm^?2 UVA) for 15 min once a day in dark condition. Alevins with no exposure to either solar light or UV exposure served as the control group. No mortality was observed in the control group, whereas all of the alevins died in the 9th day in both UV exposure groups. Light microscopic and scanning electron microscopic studies showed significant reduction in the epidermis thickness, number of skin mucous cells, and percentage of mucus secreting cells in both UV exposure groups. Sunburn cells, lifting epidermis from basal membrane, and disappearance of pavement cell microridges were also observed. Our results showed that UV radiation caused intensive skin cell lesions in the Caspian salmon alevins. Since the skin cells of trout alevins possesses essential roles in respiration, osmoregulation, excretion and defense, the observed damages to the skin could be attributed to their sudden mortality in the UV exposure.     

UV induces reactive oxygen species,damages sperm,and impairs fertilisation in the sea urchin <Emphasis Type="Italic">Anthocidaris crassispina</Emphasis>

We demonstrated that environmentally relevant levels of UVA and UVB can reduce sperm motility (UVA: by 38–58%; UVB: by 42–85%; P < 0.05) and subsequently fertilisation success (UVA: by 38–72%; UVB: by 91–98%; P < 0.05) of urchins (Anthocidaris crassispina) in a dose-dependent manner, implicating that recruitment of urchin populations might be reduced by UVR (ultraviolet radiation) prevailing in their natural habitats. Concomitantly, reactive oxygen species (ROS) production was enhanced by UVA and UVB in a dose-dependent manner (UVA: by 1.3-fold; UVB: by 6.6–7.3-fold; P < 0.05), and the increase in ROS resulted in an increase in lipid peroxidation (LPO) in urchin sperm (UVA: by 4.2–7.2-fold; UVB: by 2.3–2.7-fold; P < 0.05). This study demonstrated that ROS production and oxidative damages enhanced by UVR may account for the observed declines in sperm motility and fertilisation, and suggests that levels of UVR prevailing in the environment may pose a significant threat to the reproductive success of natural populations of urchins spawning in shallow waters.     

Photoinhibition of photosynthesis in a sun and a shade species of the red algal genus Porphyra

Gametophytes of two species of Porphyra collected around San Juan Island, Washington in 1986 and acclimated to low light conditions in culture showed different resistances to photoinhibition of photosynthesis. The intertidal species P. perforata J. Agardh exhibited photoinhibition at one-third the rate exhibited by the subtidal species P. nereocystis Anderson following treatments at 2000 mol photons m^-2 s^-1 under conditions of full hydration and optimal temperature. The greater resistance of P. perforata to photoinhibition could not be attributed to reduced photosynthetic pigment concentration, higher photosynthetic capacity, avoidance of light by chloroplast movement or to enhanced rates of photorespiration. Total carotenoid concentrations were similar in the two species. It is probable that the mechanisms of this resistance are operating at the level of the thylakoid membranes. Resistance to photoinhibition represents an adaptation of photosynthesis in P. perforata which may contribute to its persistance in the extreme environment of its intertidal habitat.     

Differences between native and invasive<Emphasis Type="Italic"> Caulerpa taxifolia</Emphasis>: a link between asexual fragmentation and abundance in invasive populations

The tropical and subtropical marine green alga Caulerpa taxifolia has invaded several temperate regions throughout the world, including southeastern Australia. In this study, I examined how invasive C. taxifolia from temperate southeastern Australia differed from native C. taxifolia from subtropical Moreton Bay, Australia, in the traits proposed as being important to its invasion success: thallus size and density, levels of asexual reproduction (fragmentation) and total biomass. Against the prediction of a large size for invasive C. taxifolia, native populations from Moreton Bay had larger stolons and fronds than invasive populations. However, invasive populations consistently had much higher densities of stolons, fronds and fragmented fronds; and a greater biomass compared to native populations. Average densities at invasive sites exceeded 4,700 stolons and 9,000 fronds/m² and were as high as 27,000 stolons and 95,000 fronds/m², which are the highest reported for C. taxifolia anywhere. Average densities of fragmented fronds at invasive sites were as high as 6,000/m² and up to 45% of all stolons at invasive sites could be directly linked to asexual recruitment via fragmented fronds. Importantly, at invasive locations there was a strong association between asexual reproduction and abundance demonstrated by positive correlations between the density of fragmented fronds and total biomass. These findings are the first to describe quantitative differences between native and invasive C. taxifolia and to demonstrate a link between the high levels of asexual reproduction and high abundance in invasive populations. Although the causes and consequences of high levels of asexual reproduction remain to be explored, this study suggests that changes in demographic and life-history traits during the invasion by C. taxifolia into temperate habitats may contribute to its success there.Communicated by M. S. Johnson, Crawley     

Photosynthesis and growth of red and green morphotypes of Kappaphycus alvarezii (Rhodophyta) from the Philippines

The effect of photosynthetic available radiation (PAR) levels, light quality, ultraviolet (UV) radiation, and temperature on photosynthesis, growth, and chlorophyll fluorescence was evaluated in red and green morphotypes of the rhodophyte Kappaphycus alvarezii (Doty) Doty under controlled conditions. Chlorophyll a and phycoerythrin (PE) levels were similar in the red and green morphotypes cultured under the same conditions, but phycocyanin (PC) and allophycocyanin (APC) levels were 2-fold greater in the green than in the red morphotype. Pigment characterization indicated that the overexpression of PC and APC masked the red pigmentation in the green morphotype. Maximum photosynthesis and photosynthetic efficiency were similar between the two morphotypes assayed at a wide temperature range, which was reflected in the similar growth rates observed in outdoor culture systems. In the green morphotype, photosynthetic efficiency increased 2-fold relative to the red morphotype when assayed with red light (λ > 600 nm), indicating that photosynthetic characteristics are modified as a result of pigment variation in these morphotypes. Such increase in photosynthetic efficiency in the green morphotype, however, did not result in greater growth rates when cultured under white light. Short exposure to high levels of solar radiation (UV-A + UV-B + PAR), and filtered solar radiation (UV-A + PAR or PAR) decreased effective quantum yield (ΔF/F _m′) in both morphotypes. The reduction of ΔF/F _m′ values in the red and green morphotypes was accounted for by high levels of PAR and not by the UV-A + UV-B + PAR and UV-A + PAR treatments. Photoinhibition caused by UV-A, UV-B, or PAR was completely reversed within 30 h after incubations. Recovery rates from photoinhibition, however, were significantly reduced in the green morphotype when incubated with UV-B radiation. The results here suggest that the overexpression of pigments do not necessarily increase photosynthesis and growth in these morphotypes. Received: 19 June 2000 / Accepted: 28 November 2000