Seasonal variation in ecophysiological patterns in macroalgae from an Arctic fjord. I. Sensitivity of photosynthesis to ultraviolet radiation

Marine macroalgae inhabiting Arctic coastal ecosystems are exposed to pronounced seasonal variations in the radiation regime, including harmful UVB radiation. This study presents the first data on the seasonal changes in the sensitivity of macroalgal photosynthesis towards UV exposure by comparing under-ice, clear-water and turbid-water conditions characteristic for late winter, spring and summer. Various brown (Laminaria saccharina, L. digitata, L. solidungula, Saccorhiza dermatodea, Desmarestia aculeata), red (Palmaria palmata, Devaleraea ramentacea) and one green macroalgal species (Monostroma aff. arcticum) were collected at the same water depth throughout the seasons in the Kongsfjord (Spitsbergen, Svalbard, Norway). Maximum quantum yield (F_v/F_m) and maximum photosynthetic electron transport rates (ETR_max) were determined immediately after collection, after 2 h exposure to artificial UV radiation and after 18 h recovery in dim white light. Photosynthesis of the studied species showed different responses depending on their morpho-functional and physiological characteristics, their life strategies, phenology and depth distribution. Within the genus Laminaria, maximum quantum yield of adult specimens of the deep-water species L. solidungula was most strongly UV sensitive. Adult L. saccharina exhibited a lower UV sensitivity than a 6-month-old specimen. Inhibition of photosynthesis after UV exposure remained at the same level throughout the study period, both in adult L. saccharina and S. dermatodea. However, adult specimens of L. saccharina collected in May showed partial recovery only, whereas photosynthesis of specimens from both species collected later recovered fully. D. aculeata exhibited a remarkable decrease of UV sensitivity during the study period. Photosynthesis of specimens collected under the ice was strongly inhibited by UV, but the degree of inhibition decreased during spring and summer. Concomitantly ETR_max values were low after UV exposure in specimens collected in June, but increased later in the season. P. palmata exhibited a relatively flexible response. Photosynthesis in specimens collected under the ice in June or in turbid water in July/August was relatively strongly inhibited; specimens collected during sunny periods and in clear water in spring showed a much lower degree of photoinhibition after UV exposure. The seasonal pattern of low/high ETR_max values in spring/summer is probably a characteristic of the life strategy of this species. The UV sensitivity of D. ramentacea exhibited a similar seasonal pattern. In M. aff. arcticum, UV sensitivity increased and ETR_max values decreased during the study period, reflecting the life strategy of this annual late winter/spring species. The physiological basis for the seasonal changes in UV sensitivity of photosynthesis is presented in a companion paper (this issue).     

Sensitivity and acclimation to UV radiation of zoospores from five species of Laminariales from the Arctic

Spores of five Laminariales from Arctic Spitsbergen were exposed in the laboratory to photosynthetically active radiation (PAR; 400–700 nm), PAR+UVA radiation (UVAR; 320–400 nm) and PAR+UVAR+UVB radiation (UVBR; 280–320 nm). Subsequently, germination was monitored over periods of 3, 6 and 9 days. The investigated species were the upper sublittoral Saccorhiza dermatodea, the upper to mid-sublittoral Alaria esculenta and Laminaria digitata, the mid-sublittoral L. saccharina and the lower sublittoral L. solidungula. The germination capacity decreased sharply after 16 h exposure to PAR+UVAR+UVBR in all species. However, S. dermatodea was able to recover from the damaging effects of UVBR. There was also a small increase in percentage germination of A. esculenta 6–9 days after the treatment. No recovery was evident in the other species. After 8 h exposure to PAR+UVA+UVB, L. digitata recovered completely, and L. saccharina and L. solidungula, partially. The only species susceptible to PAR+UVAR was L. solidungula. One prominent cytological feature of UVR-exposed spores was the enlargement of phenolic vesicles (physodes) (particularly seen in S. dermatodea and A. esculenta), which may have a protective function against UVR. Pilot experiments under natural irradiance conditions indicate that the PAR component of solar radiation exerts an additional stress. Overall the data show that zoospores of the species from the upper sublittoral are less sensitive to UVR or have the capacity to recover from UV stress in contrast to species from deeper waters, probably due to their UV protective and repair capabilities.Communicated by O. Kinne, Oldendorf/Luhe     

Effects of ultraviolet and photosynthetically active radiation on five seagrass species

Five seagrass species [Halophila ovalis (R.Br) Hook. f., Halodule uninervis (Forsk.) Aschers., Zostera capricorni Aschers., Cymodocea serrulata (R.Br) Aschers. (ed.) and Syringodium isoetifolium (Aschers.) Dandy] from Moreton Bay, Australia, were grown under increased (+25%) and ambient levels of ultraviolet (UV) radiation and photosynthetically active radiation (PAR), and various morphological and physiological responses were examined. Leaf fluorescence ratio (variable:maximum fluorescence) in conjunction with xanthophyll pigment content (violaxanthin, antheraxanthin and zeaxanthin) were used as a measure of photosynthetic efficiency. In addition, absorbance in the UV spectrum, chlorophyll content and chloroplast density were used as indicators of photosynthetic capacity. The seagrass species examined had varying degrees of sensitivity to UV radiation. Halophila ovalis and Halodule uninervis were the most sensitive species, exhibiting the largest decrease in photosynthetic efficiency and chloroplast density and the smallest increase in UV-blocking pigments in response to UV radiation. The more UV-tolerant species, Z. capricorni, C. serrulata and S. isoetifolium, were only significantly affected by increased levels of UV radiation, showing a gradual decline in photosynthetic efficiency and chloroplast density and the largest increases in UV-blocking pigment. UV sensitivity corresponded with leaf morphology, with thicker leaves (as in Z. capricorni, C. serrulata and S. isoetifolium) providing greater morphological protection for UV-sensitive organelles. Not all species were significantly affected by increasing PAR, with decreases in fluorescence ratio and increases in zeaxanthin content observed only in C. serrulata and S. isoetifolium. Sensitivity to PAR corresponded with morphological plasticity; species exhibiting a wide range of growth forms (e.g. Halophila ovalis, Halodule uninervis and Z. capricorni) were the least sensitive to increases in PAR. Seagrass depth-distributions in Moreton Bay appear to be influenced by species sensitivity to UV radiation and PAR, with other factors such as epiphytes, shading and nutrients also affecting species' tolerance. All species were affected to some degree by UV radiation, thus future changes in UV intensity may have repercussions on the distribution of seagrasses.     

Community responses to UV radiation. II. Effects of solar UVB on field-grown diatom assemblages of the Carribean

The effects of ambient solar UV on community productivity and structure were assessed during primary succession of benthic diatoms on artificial substrate in a coral reef of the Caribbean. Artificial substrates, partially enclosed by UV cutoff filters, were placed at a depth of 60 cm below surface. During the initial colonization stages, the assemblages exposed to the full solar spectrum had a mean productivity 43.4% lower than the assemblages exposed to PAR+UVA only. Some differences in species diversity of assemblages under the different UV treatments were also observed. Sensitive species to UVB exposure were Mastogloia angulata, M. ovata, M. paradoxa, Nitzschia longissima, Plagiogramma staurophorum, Rhopalodia musculus, and Surirella ovata. These UVB effects gradually diminished as succession proceeded; 5 to 6 weeks after the placement of the substrates in the water, no significant differences in productivity were observed between the different treatments, while after 6 weeks of growth, species diversity and evenness were higher, although not statistically significant, in the UVB-exposed assemblages. During the first 2 weeks of growth, the productivity under PAR+UVA was significantly lower than that under PAR only. Received: 12 July 1996 / Accepted: 20 November 1997     

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     

Biomass, photosynthesis and growth of Laminaria saccharina in a high-arctic fjord, NE Greenland

Biomass, photosynthesis and growth of the large, perennial brown alga Laminaria saccharina (L.) Lamour. were examined along a depth gradient in a high-arctic fjord, Young Sound, NE Greenland (74°18'N; 20°14'W), in order to evaluate how well the species is adapted to the extreme climatic conditions. The area is covered by up to 1.6-m-thick ice during 10 months of the year, and bottom water temperature is <0°C all year round. L. saccharina occurred from 2.5 m depth to a lower depth limit of about 20 m receiving 0.7% of surface irradiance. Specimen density and biomass were low, likely, because of heavy ice scouring in shallow water and intensive feeding activity from walruses in deeper areas. The largest specimens were >4 m long and older than 4 years. In contrast to temperate stands of L. saccharina, old leaf blades (2-3 years old) remained attached to the new blades. The old tissues maintained their photosynthetic capacity thereby contributing importantly to algal carbon balance. The photosynthetic characteristics of new tissues reflected a high capacity for adaptation to different light regimes. At low light under ice, or in deep water, the chlorophyll a content and photosynthetic efficiency (!) were high, while light compensation (E_c) and saturation (E_k) points were low. An E_c of 2.0 µmol photons m^-2 s^-1 under ice allowed photosynthesis to almost balance, and sometimes exceed, respiratory costs during the period with thick ice cover but high surface irradiance, from April through July. Rates of respiration were lower than usually found for macroalgae. Annual elongation rates of leaf blades (70-90 cm) were only slightly lower than for temperate L. saccharina, but specific growth rates (0.48-0.58 year^-1) were substantially lower, because the old blades remained attached. L. saccharina comprised between 5% and 10% of total macroalgal biomass in the area, and the annual contribution to primary production was only between 0.1 and 1.6 g C m^-2 year^-1.     

Effects of nutrients, herbivory, and depth on the macroalgal community in the rocky sublittoral

We studied the interacting roles of nutrient availability and herbivory in determining the macroalgal community in a rocky littoral environment. We conducted a factorial field experiment where we manipulated nutrient levels and herbivory at two sublittoral depths and measured macroalgal colonization and the following young assemblage during the growing season. At the community level, grazing reduced algal colonization, though the effect varied with depth and its interaction with nutrient availability varied in time. In shallow water, the total density of macroalgae increased in response to nutrient enrichment, but the ability of grazers to reduce macroalgal density also increased with the nutrient enrichment, and thus, the community could not escape from the top-down control. In deep water, the algal density was lower, except in July when nutrient enrichment caused a very dense algal growth. Grazing at the greater depth, though effective, was generally of smaller magnitude, and in July it could not limit algal recruitment and growth. Species richness peaked at the intermediate nutrient level in deep but not in shallow water during most of the growing season. Grazing had no effect on diversity of the algal community at either depth and only a minor effect on species richness at the greater depth. Opportunistic and ephemeral algae benefited from the nutrient enrichment but were also grazed to very low densities. Slowly growing and/or perennial species colonized poorly in the nutrient enriched treatments, and depending on the species, either suffered or indirectly benefited from herbivory. For all species, effects of nutrients on colonization depended on depth; usually both nutrient and herbivory effects were more pronounced at the shallow depth. We conclude that grazers are able to reduce macroalgae over a large range of nutrient availabilities, up to 12-fold nutrient enrichment in the current experiment, and that the sublittoral depth gradient generates variation in the algal community control exerted by both herbivory and nutrient availability. Thus temporal and spatial variability in both top-down and bottom-up control and in their interaction, especially along the depth gradient, may be crucially important for producer diversity and for the successional dynamic in a rocky sublittoral environment.     

Morpho-functional patterns of photosynthesis and UV sensitivity in the kelp <Emphasis Type="Italic">Lessonia nigrescens</Emphasis> (Laminariales,Phaeophyta)

The photosynthetic responses of the south Pacific kelp Lessonia nigrescens of the coast of Valdivia, Chile (40°S), were investigated by exposing its different thallus parts, fronds, stipes and holdfasts, to UV radiation in the laboratory. Biologically effective doses (BED_{photoinhibition300}) between 400 and 800 kJ m⁻² were required for a 40% inhibition in photosynthesis under UVA+UVB radiation. At BED_{photoinhibition300} close to 250 kJ m⁻² (in treatments without UVB), the inhibition of photosynthesis did not exceed 20%. These UV doses were in the range of current daily doses measured in Valdivia on cloudless summer days. In general, exposure to UVB for periods longer than 12 h reduced photosynthesis, measured as maximal quantum yield (F _v/F _m) and electron transport. The fronds were the most UV-sensitive section of this alga, coinciding with the highest pigments contents and carbon fixation. Evidence of a photodamage was also seen. After a 48 h exposure to PAR+UVA+UVB, a decrease of F _v/F _m in the fronds was close to 41%, while in the stipes and holdfasts it was 12 and 18%, respectively. Although the thalli from the different size classes showed marked differences in their morphology and morphometry, no obvious differences in the UV tolerance of the fronds were detected. The results indicated that the UV-related responses are integrated in the suite of morpho-functional adaptations of the alga. Although the fronds are spatially more exposed to solar radiation than basal structures (stipes and holdfast), due their high turnover rate they may compensate better detrimental effects of UV. In contrast, stipes and the holdfast are key support structures characterized by low replacement rates and designed to confer hydrodynamic resistance to drag forces.     

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.     

Growth and DNA damage in young Laminaria sporophytes exposed to ultraviolet radiation: implication for depth zonation of kelps on Helgoland (North Sea)

Growth as an integrative parameter of all physiological processes was measured in young sporophytes of temperate Laminaria digitata, Laminaria saccharina and Laminaria hyperborea exposed in the laboratory to irradiance consisting of either only photosynthetically active radiation (PAR) or to a spectrum including ultraviolet radiation (UVR) (PAR+UVA+UVB) by use of cut-off glass filters. Size increment was measured every 10 min over a period of 18–21 days using growth chambers with on-line video measuring technique. In the chamber, plants were grown at 10±2°C and 16:8 h light–dark cycles with 6 h additional UVR exposure in the middle of the light period. Tissue morphology and absorption spectra were measured in untreated young sporophytes while chlorophyll a content and DNA damage were measured in treated thalli at the end of the experiment. Sensitivity of growth under UVR was found to be related to the observed upper depth distribution limit of the upper sublittoral L. digitata, upper to mid sublittoral L. saccharina and lower sublittoral L. hyperborea. Tissue DNA damage is, however, dependent on thallus thickness which minimizes UVR effect where outer cell layers shade inner cells and provide longer pathlength for UVR. Exposure to UVR causes cellular, enzymatic and molecular damage. Presence of UV-absorbing compounds further reduces effective UVR from reaching physiological targets. The cost of producing higher amount of UV-absorbing compounds and effective DNA repair mechanism can, however, divert photosynthate at the expense of growth. Tissue chlorophyll a content was not significantly different between treatments suggesting a capacity for acclimation to moderate UVR fluence. Growth acclimation to repeated UVR exposure was observed within a period of 12 days while growth inhibition was observed after a longer UVR exposure period of 21 days. The results give further insight into the effects of UVR on the cellular level and show how ecological parameters such as the upper depth distribution limit are dependent on cellular processes.     

Seasonal and sex-specific variations in levels of photo-protecting mycosporine-like amino acids (MAAs) in soft corals

This study focusses on the nature and extent of variation in mycosporine-like amino acids (MAAs) in relation to annual cycles in solar radiation, seawater temperature, and reproduction in reef-flat populations of two soft coral species. The results show MAA tissue concentrations in shallow water colonies of Lobophytum compactum and Sinularia flexibilis to be significantly correlated to annual cycles in solar radiation (P<0.0006 and P<0.0005, respectively) and seawater temperature (P<0.0006 and P<0.0004, respectively). Evidence of seasonal cycles in MAA levels in the tissues of shallow-reef invertebrates positively correlating with annual cycles in solar radiation and temperature suggests that they are an integral component of the soft corals' biochemical defence system against high irradiance and/or temperature stress and thus bleaching. This is further corroborated by the higher production of MAAs in females than males prior to spawning (up to 67% and 56% for L. compactum and S. flexibilis, respectively), presumably to provide a high level of protection against irradiance stress for progeny.     

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.     

Light regime in an Arctic fjord: a study related to stratospheric ozone depletion as a basis for determination of UV effects on algal growth

Solar radiation as a primary abiotic factor affecting productivity of seaweeds was monitored in the Arctic Kongsfjord on Spitsbergen from 1996 to 1998. The radiation was measured in air and underwater, with special emphasis on the UV-B (ultraviolet B, 280–320 nm) radiation, which may increase under conditions of stratospheric ozone depletion. The recorded irradiances were related to ozone concentrations measured concurrently in the atmosphere above the Kongsfjord with a balloon-carried ozone probe and by TOMS satellite. For comparison, an ozone index (a spectroradiometrically determined irradiance of a wavelength dependent on ozone concentration, standardized to a non-affected wavelength) was used to indicate the total ozone concentration present in the atmosphere. Weather conditions and, hence, solar irradiance measured at ground level were seldom stable throughout the study. UV-B irradiation was clearly dependent on the actual ozone concentration in the atmosphere with a maximal fluence rate of downward irradiance of 0.27 W m⁻² on the ground and a maximal daily fluence (radiation exposure) of 23.3 kJ m⁻². To characterize the water body, the light transmittance, temperature and salinity were monitored at two different locations: (1) at a sheltered shallow-water bay and (2) at a wave-exposed, deep-water location within the Kongsfjord. During the clearest water conditions in spring, the vertical attenuation coefficient (K _d) for photosynthetically active radiation (PAR) was 0.12 m⁻¹ and for UV-B 0.34 m⁻¹. In spring, coinciding with low temperatures and clear water conditions, the harmful UV radiation penetrated deeply into the water column and the threshold irradiance negatively affecting primary plant productivity was still found at about 5–6 m depth. The water body in spring was characterized as a Jerlov coastal water type 1. With increasing temperature in summer, snow layers and glacier ice melted, resulting in a high discharge of turbid fresh water into the fjord. This caused a stratification in the optical features, the salinity and temperature of the water body. During melt-water input, a turbid freshwater layer was formed above the more dense sea water. Under these conditions, light attenuation was stronger than defined for a Jerlov coastal water type 9. Solar radiation was strongly attenuated in the first few metres of the water column. Consequently, organisms in deeper water are protected against harmful UV-B radiation. In the surface water, turbidity decreased when rising tide caused an advection of clearer oceanic water. In the course of the summer season, salinity continuously decreased and water temperature increased particularly in shallow water regions. The impact of global climate change on the radiation conditions under water and its effects on primary production of seaweeds are discussed, since organisms in the eulittoral and upper sublittoral zones are affected by UV radiation throughout the polar day. In clearer water conditions during spring, this may also apply to organisms inhabiting greater depths. Received: 20 June 2000 / Accepted: 17 October 2000     

Microhabitat segregation of the amphipod genus Gammarus (Crustacea: Amphipoda) in the Northern Baltic Sea

Closely related species may occupy very similar niches but are often found to diverge by one or more traits when they share the same habitat. Five indigenous and sympatric Gammarus species are characteristic for the Baltic rocky littoral ecosystem. Yet, the species-specific distribution of these sympatric gammarids has not been well studied in the northern Baltic Sea. This study was undertaken to assess the spatial distribution of gammarid amphipods along wave exposure and depth gradients to study whether they show segregation in their microhabitat use. We sampled 12 rocky sublittoral shores along a wave exposure gradient over a period of 5 years. Samples differed with respect to depth and macroalgal type. Three of the five gammarid species occurred mainly in different depth zones and among different macroalgae at the exposed shores. In contrast, on protected shores, where algal zonation is weaker, a link to zonation and macroalgal type was almost absent. Moreover, the microhabitat use was strongest during the reproductive seasons of the species. The observed microhabitats of the three gammarid species fit well to their species-specific mean body sizes.     

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.     

Spatio-temporal variation of hydroids and polychaetes associated with Cystoseira amentacea (Fucales: Phaeophyceae)

The temporal and spatial distribution of hydroids (sessile fauna) and polychaetes (vagile) on the brown alga Cystoseira amentacea (Fucales: Phaeophyceae) have been studied in the mid-littoral zone of the Apulian coast (Italy). Samplings were carried out in February and July 1997, at 1.5 m depth, at three sites (Gargano, Costa Merlata and Otranto), about 200 km apart from each other. Three replicates per site were collected for polychaetes, and another three, for hydroids. Samples yielded 142,426 polyps belonging to 20 hydroid species and 3,088 polychaete specimens belonging to 58 species. Seasonal variations in abundance and number of species were evident in hydroids. The only species probably exclusive of Cystoseira (the hydroid Clytia viridicans) was present only in the summer. Polychaetes showed marked seasonal differences only at the Gargano site. Multivariate techniques were used to compare, in both periods, assemblages within and among sites. Analysis of similarity testing revealed that, for both polychaetes and hydroids, assemblage structures significantly differed among sites and between seasons. Thus, the null hypothesis that the distribution in time and space of the epifauna on the same algal species is homogeneous over a wide geographical scale has been rejected, in spite of the homogeneity of the substrate. In both groups, quantitative differences of a few species seemed to be more important than qualitative ones in determining the spatial separation of sites, the species pool remaining rather consistent. A small-scale variability among replicates was also detected, especially at Gargano, for hydroids. Since the epifauna can allow better comparisons among sites than the actual distribution of the algal species, both sessile and vagile epiphytes of Cystoseira can be considered valid tools for evaluating environmental changes on coastal hard-bottom communities.     

Variation in UVB sensitivity of planula larvae of the coral Agaricia agaricites along a depth gradient

The effects of natural intensities of ultraviolet A (UVA, 320 to 400 nm) and B (UVB, 280 to 320 nm) radiation on planktonic planula larvae of the reef-building coral Agaricia agaricites (Linnaeus) were investigated through field experiments. Survival, chlorophyll concentrations, and solubilized protein concentrations were determined for larvae spawned from colonies at 3 and 24 m depth and subjected to one of three light regimes at 3, 10, or 24 m depth for 72 h: PAR (photosynthetically active radiation, 40- to 700 nm) only, PAR+UVA, or PAR+UVA+UVB. At 3 m depth, larvae in the PAR+UVA+UVB treatment showed lower survivorship than larvae exposed to either PAR alone or PAR+UVA. Within the PAR+UVA+UVB treatment at 3 m depth, larvae from colonies at 24 m depth suffered higher mortality than those from 3 m. Differences in survivorship between larvae originating from 3 and 24 m depth corresponded with tissue concentrations of UVB-protective mycosporine-like amino acids: larvae from 3 m had higher concentrations of mycosporine-glycine (_max=310nm) and palythine (_max=320nm) than those from 24 m depth. Chlorophyll concentrations were inversely correlated with PAR intensities, but variation in this parameter did not appear to be detrimental. These results show that sensitivity to high intensities of UVB radiation may affect survival of A. agaricites larvae in shallow reef-waters.     

Blue mussels, Mytilus edulis, at the edge of the range: population structure, growth and biomass along a salinity gradient in the north-eastern Baltic Sea

The blue mussel, Mytilus edulis, is the most conspicuous animal species in the northern Baltic rocky sublittoral. In the studied area the species lives at the margin of its salinity tolerance. Although dwarfed by the low-salinity conditions, blue mussels in the northern Baltic are very abundant and have a decisive role in the benthic and pelagic ecosystems. We studied abundance, size distribution, biomass and growth rate of blue mussels along a 270 km salinity gradient in the northern Baltic Sea. Samples (n=317, 1998-1999) from moderately exposed and exposed rocky shores at seven study areas were collected in the southern Archipelago Sea in the west and into the central Gulf of Finland in the east, where the species is becoming increasingly rare. The results show a marked decline in mean mussel size and biomass from the saline west to the less saline east. The growth rate also decreased with lower ambient salinity. However, abundance of small mussels was considerably higher in the central and eastern parts of the study area. Vertically, the highest biomass was recorded at intermediate depths (5 and 8 m), being lower at both shallower (3 m) and deeper bottoms (12 m). It is concluded that salinity is the foremost factor determining size structure and growth rate among populations within the area. The results suggest that predation further influences the population structure of blue mussels living at the edge of their range in the central Gulf of Finland ultimately set by their salinity tolerance.     

A comparative analysis of the photobiology of zooxanthellae and zoochlorellae symbiotic with the temperate clonal anemone Anthopleura elegantissima (Brandt)

The temperate anemone Anthopleura elegantissima hosts two phylogenetically different symbiotic microalgae, a dinoflagellate Symbiodinium (zooxanthellae, ZX) and a chlorophyte (zoochlorellae, ZC), throughout certain regions of its latitudinal range. Because of the broad intertidal and geographic range of this anemone, we examined the role of irradiance to ascertain which specific symbiotic parameters are affected and whether light intensity governs the observed distributions of natural populations of ZX and ZC. Irradiance appears to be a key factor in regulating both the photophysiology and metabolism of this alga-cnidarian association. Regardless of light intensity, algal densities remained stable for anemones harboring ZX or ZC, whereas the mitotic indices of ZX and ZC both varied directly with light intensity. The chlorophyll content of ZX remained fairly constant regardless of irradiance; in contrast, ZC chlorophyll content was inversely proportional to light intensity. Regardless of irradiance, the carotenoid content of both symbionts was constant; however, ZX carotenoid levels were higher than those of ZC. Net photosynthesis was directly related to light intensity for both algal symbionts and ZX photosynthetic rates were consistently higher than those of ZC. Similarly, the potential carbon contribution of ZX and ZC to animal respiration (CZAR) displayed a direct relationship with light intensity, peaking at 800 µmol·m^-2·s^-1, then subsequently declined. Lower ZX growth rates, coupled with higher photosynthetic rates and higher CZAR estimates, compared to ZC, suggest that the ZX should be the dominant symbiont as light intensity increases; this may explain the high densities of anemones in the field containing ZX where the levels of irradiance are naturally high. These results support the interpretation that irradiance is a significant environmental parameter that dictates the microhabitat and latitudinal distribution of the two symbiotic algal taxa. This is the second in a series of papers examining the physical parameters that influence the distribution of ZX- and ZC-bearing A. elegantissima.     

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.