121 Influence of Benthic vs. Pelagic Seeding on Coastal Diatom Bloom Development

Diatom blooms may be initiated by cells that have survived in the plankton or germinated resting stages from the sediments. However, it is not well understood how these different inocula contribute to bloom development. We followed diatom community development in twenty‐liter microcosms given different inocula. Surface sediment and phytoplankton were collected in Gullmar Fjord, Sweden. Replicate microcosms were then dosed with local sediment and/or plankton and incubated in situ in Gullmar Fjord. We also followed the concurrent development of the phytoplankton community in the fjord. Experiments run in both spring and fall 2002 showed that bloom development in the microcosms was significantly faster when seeded by planktonic cells. However, addition of sediment may have stimulated planktonic growth and also provided additional propagules. The type of inoculum used strongly influenced the diatom composition in the microcosms. Sediment additions, through germination of resting stages, resulted in communities dominated by Detonula confervacea and Thalassiosira minima in spring, and Skeletonema costatum in fall. Planktonic inocula resulted in blooms of T. nordenskioeldii and Chaetoceros debilis in spring, and S. costatum and several Chaetoceros spp. in fall. Microcosms dosed with both plankton and sediment showed a mixed species assemblage. Comparisons between the microcosms and fjord phytoplankton suggest an important role for benthic seeding of diatom blooms.     

Linking the planktonic and benthic habitat: genetic structure of the marine diatom Skeletonema marinoi

Dormant life stages are important strategies for many aquatic organisms. The formation of resting stages will provide a refuge from unfavourable conditions in the water column, and their successive accumulation in the benthos will constitute a genetic reservoir for future planktonic populations. We have determined the genetic structure of a common bloom‐forming diatom, Skeletonema marinoi, in the sediment and the plankton during spring, summer and autumn two subsequent years (2007–2009) in Gullmar Fjord on the Swedish west coast. Eight polymorphic microsatellite loci were used to assess the level of genetic differentiation and the respective gene diversity of the two different habitats. We also determined the degree of genetic differentiation between the seed banks inside the fjord and the open sea. The results indicate that Gullmar Fjord has one dominant endogenous population of S. marinoi, which is genetically differentiated from the open sea population. The fjord population is encountered in the plankton and in the sediment. Shifts from the dominant population can happen, and in our study, two genetically differentiated plankton populations, displaying reduced genetic diversity, occurred in September 2007 and 2008. Based on our results, we suggest that sill fjords maintain local long‐lived and well‐adapted protist populations, which continuously shift between the planktonic and benthic habitats. Intermittently, short‐lived and mainly asexually reproducing populations can replace the dominant population in the water column, without influencing the genetic structure of the benthic seed bank.     

Competing phytoplankton undermines allelopathy of a bloom-forming dinoflagellate

Biotic interactions in the plankton can be both complex and dynamic. Competition among phytoplankton is often chemically mediated, but no studies have considered whether allelopathic compounds are modified by biotic interactions. Here, we show that compounds exuded during Karenia brevis blooms were allelopathic to the cosmopolitan diatom Skeletonema costatum, but that bloom allelopathy varied dramatically among collections and years. We investigated several possible causes of this variability and found that neither bloom density nor concentrations of water-borne brevetoxins correlated with allelopathic potency. However, when we directly tested whether the presence of competing phytoplankton influenced bloom allelopathy, we found that S. costatum reduced the growth-inhibiting effects of bloom exudates, suggesting that S. costatum has a mechanism for undermining K. brevis allelopathy. Additional laboratory experiments indicated that inducible changes to K. brevis allelopathy were restricted to two diatoms among five sensitive phytoplankton species, whereas five other species were constitutively resistant to K. brevis allelopathy. Our results suggest that competitors differ in their responses to phytoplankton allelopathy, with S. costatum exhibiting a previously undescribed method of resistance that may influence community structure and alter bloom dynamics.     

Marine pelagic rotifers and tintinnids - important trophic links in the spring plankton community of the Gullmar Fjord, Sweden

During the winter and spring of 1981, the dynamics of a marinepelagic rotifer, Synchaeta vorax Rousselet and tintinnids ofthe genus Tintinnopsis, were studied in the Gullmar Fjord onthe Swedish west coast. Although present in February and March,it was not until early April when the surface water temperatureexceeded 6?C that the rotifer population started to grow rapidly.Thereafter, the population increased exponentially for 6 weeks.Estimates of consumption of phytoplankton by rotifers and tintinnidsin their most dynamic phase approximated to that of the phyto-planktonproduction. Lack of data on the amount of detritus availableand the number of young rotifers passing the 90 µm netare however obvious limitations in such calculations. Nevertheless,the results indicate that the tintinnids and the rotifers arehighly important components in the area's spring plankton community.Through high turnover rates, they are able to utilize effectivelythe spring phytoplankton bloom and thus serve as a link betweenthe primary and the larger secondary producers.     

Seasonal and annual variability in the phytoplankton community of the Raunefjord,west coast of Norway from 2001–2006

Changes in phytoplankton community composition potentially affect the entire marine food web. Because of seasonal cycles and inter-annual variations in species composition, long-term monitoring, covering many sequential years, is required to establish a baseline study and to reveal long-term trends. The current study describes the phytoplankton biomass variations and species composition in relation to hydrographic and meteorological conditions in the Raunefjord, western Norway, over a 6-year period from 2001 to 2006. The extent of inflow or upwelling in the fjord varied from year to year and resulted in pronounced differences in water column stability. The annual phytoplankton community succession showed some repeated seasonal patterns, but also high variability between years. Two to four diatom blooms were observed per year, and the spring blooms occurring before water column stratification in March were dominated by Skeletonema marinoi and Chaetoceros socialis, and other Chaetoceros and Thalassiosira spp. Blooms of the haptophytes Phaeocystis pouchetii and Emiliania huxleyi were irregular and in some years totally absent. Although E. huxleyi was present all year round it appeared in bloom concentrations only in 2003, when the summer was warm and the water column characterized by high surface temperatures and pronounced stratification. The annual average abundance of both diatoms and flagellates increased during the six years. Despite the high variation from year to year, our investigation provides valuable knowledge about annual phytoplankton community patterns in the region, and can be used as a reference to detect possible future changes.     

Relationship Between Periodic Resuspension Events and Planktonic Diatom Community Structure in Lake Michigan: A Field and Laboratory Investigation

Lake Michigan provides an ideal location for comparing episodic physical forcing events (storms) on phytoplankton processes and the more persistent seasonal variability of phytoplankton communities. This is due to the duration and extent of the highly turbid, recurrent coastal plume (RCP) in the lake during the winter/early spring. Although the RCP can coincide with the initiation of the basin wide spring diatom bloom, linkages between the duration and intensity of the plume and the prominent role of light availability in regulating Lake Michigan phytoplankton growth during the spring isothermal period have been postulated, but not verified. The concurrent physical and biological events provide a novel opportunity to examine phenomena associated with the RCP affecting the distribution and abundance of species in Lake Michigan's planktonic diatom flora. In this study, planktonic diatom assemblages from pre, post and active spring plume events were examined from stations along Lake Michigan's southern shoreline. Species abundance changed rapidly during storm events. Sediment resuspension via storm activity created a suboptimal growth environment. Post‐storm event diatom phytoplankton communities were floristically distinct from prestorm event communities, with resting cell‐forming taxa playing a significant role in these community‐restructuring periods. Laboratory simulations of resuspension events using Lake Michigan sediments were conducted under a variety of environmental conditions. Parameters varied included day length, temperature, and silica. The resulting assemblages were quantitatively counted. A statistically significant relationship was identified between day length and the vegetative growth of many resting cell‐forming diatom species. When day length was calculated for poststorm event field data, it revealed a high correlation between poststorm event communities and those predicted by the laboratory simulations. Timing of storm events and latitudinal position of the aquatic system are then important elements for consideration when predicting diatom phytoplankton community structure due to the relationship these factors have with day length.     

SEASONAL VARIABILITY IN POLYUNSATURATED ALDEHYDE PRODUCTION POTENTIAL AMONG STRAINS OF SKELETONEMA MARINOI (BACILLARIOPHYCEAE)1

The cosmopolitan bloom‐forming diatom Skeletonema marinoi Sarno et Zingone is known to produce toxic polyunsaturated aldehydes (PUAs) in response to cell damage that can affect a diverse suite of organisms, including grazing species and competitor plankton species. The production of PUAs in nine different S. marinoi strains isolated at three different times of the year (spring, summer, and autumn) was assessed in relation to the predominant conditions at the time of isolation from Gullmar Fjord, Skagerrak. During the initial stages of growth, PUA production potential of S. marinoi was generally the highest in summer strains, although there was a substantial variation among strains isolated at the same time. Spring strains, however, showed a strong capacity for increased PUA production potential in later stage cultures with diminishing nutrient levels, reaching amounts similar to those observed in summer strains. In contrast, PUA production potentials of summer and autumn strains did not change significantly from the original values. There is negligible grazing pressure during the spring bloom in Gullmar Fjord, but a potential for high competition for resources, such as nutrients, toward the later stages of the bloom. In contrast, grazing pressure is much greater during summer and autumn, and there may also be nutrient limitation at this time. The PUA production potentials of S. marinoi appear to reflect the ecological conditions at the time of isolation with higher production potentials in strains isolated when conditions were likely to be less beneficial for survival.     

The effects of temperature increases on a temperate phytoplankton community — A mesocosm climate change scenario

Prior to the spring bloom in 2003 and 2004, batch temperature experiments of approximately 3 weeks' duration were carried out in land-based mesocosms in at the Espeland field station (Norway), with temperatures on average increased ~ 2.7-3 °C (T1) and ~ 5.2-5.6 °C (T2) above in situ fjord temperature (RM). The development in the chlorophyll concentrations showed an earlier bloom as a response to increased temperatures but the carbon biomass showed that the warmest treatment yielded the lowest biomass. This study indicates that a part of the relationship between temperature and spring bloom timing stems from a temperature-induced change in phytoplankton algal physiology (the efficiency of photosystem II, F_v/F_m, and growth rates, µ_max), i.e. a direct temperature effect. Data analysis performed on microscope identified and quantified species did not show a significant temperature influence on phytoplankton community composition. However, the HPLC data indicated that temperature changes of as little as 3 °C influence the community composition. In particular, these data showed that peridinin-containing dinoflagellates only increased in abundance in the heated mesocosms and that a prasinophycean bloom, which was undetected in the microscope analyses, occurred prior to the blooms of all other phytoplankton classes in all treatments. The microscope analyses did reveal a temperature effect on individual species distribution patterns. Thalassionema nitzschioides was more abundant in the warm treatments and, in the warmest treatment, the spring bloom forming Skeletonema marinoi comprised a smaller proportion of the diatom community than in the other treatments.     

南麂列岛海洋自然保护区浮游植物的种类多样性及其生态分布

于2006年5月至2007年2月之间,对南麂列岛海域的浮游植物类群进行了4个季节的调查,分析了该海域浮游植物的种类组成、优势种类、群落结构以及水平分布等特征参数的季节变化。共鉴定浮游植物80种,隶属于4个门,硅藻种类最多,甲藻其次。浮游植物可划分为3个生态类群,以广温类群为主。春季和夏季分别以三角棘原甲藻和中肋骨条藻为绝对优势种,秋冬季的优势种类组成多样化。共鉴定57种赤潮生物,占浮游植物种类数的71.25%。调查期间,三角棘原甲藻和中肋骨条藻分别于春季和夏季形成赤潮。浮游植物的物种丰富度呈现春、夏、秋、冬递减的趋势。浮游植物细胞丰度的年平均值为1.03×106cells/L,春夏季显著高于秋冬季。春季和夏季时,浮游植物高值区集中在南麂岛西北近岸海域;秋季和冬季时,浮游植物高值区相对集中在南麂岛东南近岸海域。浮游植物群落的多样性指数(H')以秋季最高,冬季最低。春季的三角棘原甲藻赤潮期间,水体中N/P值显著升高;夏季的中肋骨条藻赤潮期间,水体中N/P值显著降低。     

Warming accelerates termination of a phytoplankton spring bloom by fungal parasites

Climate change is expected to favour infectious diseases across ecosystems worldwide. In freshwater and marine environments, parasites play a crucial role in controlling plankton population dynamics. Infection of phytoplankton populations will cause a transfer of carbon and nutrients into parasites, which may change the type of food available for higher trophic levels. Some phytoplankton species are inedible to zooplankton, and the termination of their population by parasites may liberate otherwise unavailable carbon and nutrients. Phytoplankton spring blooms often consist of large diatoms inedible for zooplankton, but the zoospores of their fungal parasites may serve as a food source for this higher trophic level. Here, we investigated the impact of warming on the fungal infection of a natural phytoplankton spring bloom and followed the response of a zooplankton community. Experiments were performed in ca. 1000 L indoor mesocosms exposed to a controlled seasonal temperature cycle and a warm (+4 °C) treatment in the period from March to June 2014. The spring bloom was dominated by the diatom Synedra. At the peak of infection over 40% of the Synedra population was infected by a fungal parasite (i.e. a chytrid) in both treatments. Warming did not affect the onset of the Synedra bloom, but accelerated its termination. Peak population density of Synedra tended to be lower in the warm treatments. Furthermore, Synedra carbon: phosphorus stoichiometry increased during the bloom, particularly in the control treatments. This indicates enhanced phosphorus limitation in the control treatments, which may have constrained chytrid development. Timing of the rotifer Keratella advanced in the warm treatments and closely followed chytrid infections. The chytrids' zoospores may thus have served as an alternative food source to Keratella. Our study thus emphasizes the importance of incorporating not only nutrient limitation and grazing, but also parasitism in understanding the response of plankton communities towards global warming.     

PELAGIC AND BENTHIC ENVIRONMENTAL CONTROLS ON THE SPATIAL DISTRIBUTION OF A VIABLE DIATOM PROPAGULE BANK ON THE SWEDISH WEST COAST1

The distribution of viable diatom resting stages in sediments on the Swedish west coast was assessed by the most probable number (MPN) culture technique. Multivariate analyses correlated benthic and pelagic environmental factors to the observed spatial variations in the size and taxonomic composition of the propagule bank. Viable diatom resting stages were plentiful (0.2–4.8 million cells·g ^{? 1} 1 Received 11 September 2001. Accepted 12 June 2002.
dry weight) and were dominated by the genera Skeletonema, Detonula, Chaetoceros, and Thalassiosira. Size of the propagule bank was primarily related to planktonic biomass (measured as chl a) and was highest in the Orust‐Tjörn fjord system. Species composition in this fjord system was dominated by D. confervacea (Cleve) Gran and T. nordenskioeldii Cleve in contrast to stations on the outer coast, which contained more cells of T. minima Gaarder, Asterionellopsis glacialis (Castracane) Round, and Leptocylindrus danicus Cleve. These taxonomic variations were principally influenced by deep water oxygen concentrations and water column stability. Benthic resting cells of S. costatum (Greville) Cleve were abundant all along the coast but showed reduced viability in low oxygen environments. Calculations based on MPN values estimated that resuspension of sediment could provide a sizable inoculum to the plankton, although the development of planktonic blooms will also depend on forces of hydrography and weather. Although benthic resting stages may not be absolutely necessary for survival of all diatoms, these cells may be important in determining species cycles, succession, and the spatial distribution of diatoms.     

Earlier onset of the spring phytoplankton bloom in lakes of the temperate zone in a warmer climate

The decoupling of trophic interactions is potentially one of the most severe consequences of climate warming. In lakes and oceans the timing of phytoplankton blooms affects competition within the plankton community as well as food–web interactions with zooplankton and fish. Using Upper Lake Constance as an example, we present a model‐based analysis that predicts that in a future warmer climate, the onset of the spring phytoplankton bloom will occur earlier in the year than it does at present. This is a result of the earlier occurrence of the transition from strong to weak vertical mixing in spring, and of the associated earlier onset of stratification. According to our simulations a shift in the timing of phytoplankton growth resulting from a consistently warmer climate will exceed that resulting from a single unusually warm year. The numerical simulations are complemented by a statistical analysis of long‐term data from Upper Lake Constance which demonstrates that oligotrophication has a negligible effect on the timing of phytoplankton growth in spring and that an early onset of the spring phytoplankton bloom is associated with high air temperatures and low wind speeds.     

1996年德国石荷西部沿岸水域浮游植物种群变动及其整齐圆筛藻赤潮

1996年早春至初夏,德国石荷州西海岸水域尤其是在易白河河附近海区发生了数次赤潮事件,河口区的高叶绿素浓度总是与低盐、高营养盐浓度的分布区吻合。从早春3月至5月底,发生了持续时间很长的大规模赤潮事件,其引发赤潮原因种为整齐圆筛藻,6月份对浮游植物生物量的贡献主要来自几种硅藻如微小细柱藻、聚生角毛藻、远洋角毛灌以及属于鞭毛藻类的棕囊藻群体,7月中旬,自养的纤毛虫类Myrionecta rubra (Mesodinium rubrum）达到数量高峰。从7月下旬至8月中旬,以梭形角藻、叉状角藻占优势的甲藻类几科贡献了全部的浮游植物生物量,8月中旬至9月上旬为甲藻和硅藻等类别的混合期,9月中旬以后,硅藻重新占据浮激植物 数量的绝对优势,主要优势种为浮动弯角藻、复瓦根管藻等,该海域浮游植物种群的演替反映了温带海域的特征。在早期发生的大规模赤潮期间,整齐圆筛藻的细胞密度达到2180cells dm^-3,与之相对应的生物量为758.1μg C dm^-3,占浮游植物生物量的94.2%,整齐圆筛藻的爆发性增殖所产生的大量油膜覆盖了黑尔果兰岛和东Frisian群岛周围的海域,这一现象有史以来第一次被卫星所观测到。     

Assimilation of Diazotrophic Nitrogen into Pelagic Food Webs

The fate of diazotrophic nitrogen (N_D) fixed by planktonic cyanobacteria in pelagic food webs remains unresolved, particularly for toxic cyanophytes that are selectively avoided by most herbivorous zooplankton. Current theory suggests that N_D fixed during cyanobacterial blooms can enter planktonic food webs contemporaneously with peak bloom biomass via direct grazing of zooplankton on cyanobacteria or via the uptake of bioavailable N_D (exuded from viable cyanobacterial cells) by palatable phytoplankton or microbial consortia. Alternatively, N_D can enter planktonic food webs post-bloom following the remineralization of bloom detritus. Although the relative contribution of these processes to planktonic nutrient cycles is unknown, we hypothesized that assimilation of bioavailable N_D (e.g., nitrate, ammonium) by palatable phytoplankton and subsequent grazing by zooplankton (either during or after the cyanobacterial bloom) would be the primary pathway by which N_D was incorporated into the planktonic food web. Instead, in situ stable isotope measurements and grazing experiments clearly documented that the assimilation of N_D by zooplankton outpaced assimilation by palatable phytoplankton during a bloom of toxic Nodularia spumigena Mertens. We identified two distinct temporal phases in the trophic transfer of N_D from N. spumigena to the plankton community. The first phase was a highly dynamic transfer of N_D to zooplankton with rates that covaried with bloom biomass while bypassing other phytoplankton taxa; a trophic transfer that we infer was routed through bloom-associated bacteria. The second phase was a slowly accelerating assimilation of the dissolved-N_D pool by phytoplankton that was decoupled from contemporaneous variability in N. spumigena concentrations. These findings provide empirical evidence that N_D can be assimilated and transferred rapidly throughout natural plankton communities and yield insights into the specific processes underlying the propagation of N_D through pelagic food webs.     

Size‐spectrum based differential response of phytoplankton to nutrient and iron‐organic matter combinations in microcosm experiments in a Chilean Patagonian Fjord

The Patagonian fjords have been recognized as a major region of relatively high primary productivity systems during spring–summer bloom periods, where iron‐organic matter forms may be essential complexes involved in key growth processes connected to the carbon and nitrogen cycles. We used two dissolved organic matter (DOM) types, marine polysaccharide and siderophore, as a model to understand how they affect the bioavailability of Fe to phytoplankton and bacteria and to assess their ecological role in fjord systems. A 10‐day microcosm study was performed in the Comau Fjord during summer conditions (March 2012). Pico‐, nano‐, and microphytoplankton abundance, total chlorophyll‐a and bacteria abundance, and bacterial secondary production estimates were analyzed in five treatments: (i) control (no additions), (ii) only nutrients (NUT: PO₄, NO₃, Si), (iii) nutrients + Fe(II), (iv) polysaccharide (natural diatoms extracted: 1–3 beta Glucan), and (v) Hexandentate Desferroxiamine B (DFB, siderophore). Our results showed that while DFB reduced Fe bioavailability for almost all phytoplankton assemblages in the fjord, polysaccharide did not have effects on the iron bioavailability. At Nutrients + Fe and Polysaccharide treatments, chlorophyll‐a concentration abruptly increased from 0.9 to 20 mg m^?3 during the first 4–6 days of the experimental period. Remarkably, at the Nutrients + Fe treatment, the development of the bloom was accompanied by markedly high abundances of Synechococcus, picoeukaryotes, and autotrophic nanoflagellates within the first 4 days of the experiment. Our study indicated that small plankton (phytoplankton <20 μm and bacteria) were the first to respond to dissolved Nutrients + Fe compared to large sized micro‐phytoplankton cells (>20 μm). This could be at least partially attributed to biological utilization of Fe (2 to 3 nM) by <20 μm phytoplankton and bacteria through the interaction with organic ligands released by bacteria that eventually could increase solubility of the Fe dissolved fraction thus having a positive effect on the small‐sized phytoplankton community.     

Ctenophore-zooplankton-phytoplankton interactions in Narragansett Bay, Rhode Island, USA, during 1972-1977

Plankton dynamics at a station in lower Narragansett Bay, RIare compared for six summer and fall seasons, 1972–1977.In four of these years, initiation of the summer pulse of thectenophore Mnemiopsis leidyi was accompanied by a rapid declinein zooplankton abundance and a summer phytoplankton bloom. Terminationof the phytoplankton bloom coincided with depleted ctenophoreabundance and increased zooplankton biomass in two of the years.Yearly variations in the summer abundance of the diatom Skeletonemacostatum were positively related to the magnitude of the ctenophorepulse. The magnitude of ctenophore population was related tothe zooplankton biomass present at the start of the pulse. Theserelationships, the timing and magnitude of the plankton eventssuggest that M. leidyi regulated summer zooplankton and phytoplanktondynamics. Ctenophores may control phytoplankton blooms indirectlythrough their predation on herbivorous zooplankton and directlyby the nutrient excretion accompanying such grazing. This evidencethat a planktonic carnivore two trophic steps removed from thephytoplankton regulates the latter's dynamics in NarragansettBay is analogous to reported regulation of benthic algal (kelp)dynamics by the sea otter, lobster and various crabs throughtheir predation on herbivorous sea urchins. The factors responsiblefor the seasonal decrease in ctenophores remain unresolved;ctenophore predators on Mnemiopsis are absent in NarragansettBay. Infection by the vermiform larval anemone, Edwardsia lineata,grazing by the butterfish, Peprilus triacanthus, and changesin food availability, temperature and salinity likewise do notexplain this disappearance.     

Future Climate Scenarios for a Coastal Productive Planktonic Food Web Resulting in Microplankton Phenology Changes and Decreased Trophic Transfer Efficiency

We studied the effects of future climate change scenarios on plankton communities of a Norwegian fjord using a mesocosm approach. After the spring bloom, natural plankton were enclosed and treated in duplicates with inorganic nutrients elevated to pre-bloom conditions (N, P, Si; eutrophication), lowering of 0.4 pH units (acidification), and rising 3°C temperature (warming). All nutrient-amended treatments resulted in phytoplankton blooms dominated by chain-forming diatoms, and reached 13–16 μg chlorophyll (chl) a l⁻¹. In the control mesocosms, chl a remained below 1 μg l⁻¹. Acidification and warming had contrasting effects on the phenology and bloom-dynamics of autotrophic and heterotrophic microplankton. Bacillariophyceae, prymnesiophyceae, cryptophyta, and Protoperidinium spp. peaked earlier at higher temperature and lower pH. Chlorophyta showed lower peak abundances with acidification, but higher peak abundances with increased temperature. The peak magnitude of autotrophic dinophyceae and ciliates was, on the other hand, lowered with combined warming and acidification. Over time, the plankton communities shifted from autotrophic phytoplankton blooms to a more heterotrophic system in all mesocosms, especially in the control unaltered mesocosms. The development of mass balance and proportion of heterotrophic/autotrophic biomass predict a shift towards a more autotrophic community and less-efficient food web transfer when temperature, nutrients and acidification are combined in a future climate-change scenario. We suggest that this result may be related to a lower food quality for microzooplankton under acidification and warming scenarios and to an increase of catabolic processes compared to anabolic ones at higher temperatures.     

Impact of increased river discharge on the phytoplankton community in the outer Oslofjord,Norway

An extraordinary flood event occurred in SouthernNorway in May–June 1995. The impact of fresh waterdischarge from the river Glomma on phytoplanktonspecies composition and on primary production duringthe flood was studied. The phytoplankton community wasdominated by species usually found in the fjord, thediatom Skeletonema costatum dominated thecommunity together with cyanobacteria. Potentiallyharmful species (e.g. Chrysochromulinapolylepis) were found in low cell densities. No largeincrease in nutrient concentrations were found.Primary production and uptake rates of nitrate andammonium were very low in low-salinity water near theriver outlet. Nutrients from the river water wereefficiently utilized in the open part of the fjord.The flood trigged a phytoplankton bloom dominated byharmless species commonly observed in the fjordplankton. This revised version was published online in July 2006 with corrections to the Cover Date.     

AN ECOLOGICAL BASIS FOR EXTRACELLULAR CARBONIC ANHYDRASE IN MARINE UNICELLULAR ALGAE

Extracellular carbonic anhydrase (CA_e) is expressed by many, but not all, autotrophic species of aquatic unicellular protists. We measured CA_e activities in unicellular marine algae characteristic of either high nutrient spring, fall, and winter blooms or low nutrient summer populations to provide ecological/evolutionary information about the enzyme. Highest activities occurred in spring bloom and opportunistic diatoms exposed to long photoperiods (16 h) when pH was highest and CO₂ was lowest. Lower activities were recorded for a fall-bloom diatom exposed to the long photoperiod, and lowest values were found under all culture conditions for one diatom and a number of flagellated species typical of summer low nutrient environments. Other potential sources of variance in measurements of CA_e activity were examined. Maximum activities of CA_e were recorded for the diatom, Skeletonema costatum (Greville) Cleve, during late exponential phase of growth and within 8 h of the beginning of the photoperiod. We concluded that ecological factors are important in determining CA_e activities in marine unicellular protists. Potential functions of CA_e in the metabolism of marine unicellular algae are discussed.     

Distribution of diatom surface sediment assemblages within Effingham Inlet, a temperate fjord on the west coast of Vancouver Island (Canada)

Twenty-nine surface sediment samples from Effingham Inlet, a small fjord on the west coast of Vancouver Island, British Columbia, were analyzed for diatoms. This fjord has been selected for paleoceanographic investigation due to the presence of laminated sediments resulting from the dysoxic to anoxic bottom water conditions in the inner and outer basins of the inlet. Distributional patterns of the diatom microflora reflected proximity to littoral regions, phytoplankton production, and marine influence from outside the fjord. Principal components and cluster analyses of the microflora established four diatom assemblages with a clear separation between the inner and outer basin diatom floras. Inner basin stations were characterized by elevated absolute abundance with assemblages dominated by spring–early summer bloom taxa including Skeletonema costatum, Thalassiosira nordenskioeldii, and Thalassiosira pacifica. Chaetoceros spp. resting spores were abundant throughout Effingham Inlet, with the exception of the stations closest to the fjord head. Stations located in the outer basin and towards the fjord mouth had relatively lower absolute abundance yet showed a higher relative and absolute abundance of Thalassionema nitzschioides, Rhizosolenia setigera, Coscinodiscus radiatus, Ditylum brightwellii, Odontella longicruris, and Paralia sulcata in relation to the inner basin. Many of these latter taxa are often associated with late summer and autumn conditions in fjords along coastal British Columbia. Oceanographic data for Effingham Inlet suggest that increased offshore penetration is more likely to occur from summer to early fall, with a more restricted offshore influence in the inner basin. Diatom surface sediment assemblages in Effingham Inlet appear to reflect incursions of offshore waters into the fjord. Absolute abundance estimates and the preservation of lightly silicified taxa suggest excellent preservation of fossil material in the sediments of the predominantly anoxic inner basin. Preservation in the outer basin is reduced, reflecting more frequent recharge by oxygenated waters spilling over the outer sill into the basin. Our findings suggest the inner basin should be an optimal site for reconstruction of diatom production, with records from the outer basin providing more consistent information about offshore influence and coastal upwelling conditions over the Holocene. Estimates of diatom abundance within the inner basin sediment may serve as a good proxy of production, although proxy tracers of bottom water conditions and sedimentological analyses must be coupled to the diatom record to ensure depositional conditions were not influencing valve preservation or abundance. Our results suggest that fjords can serve as good environments for paleoceanographic reconstructions of both inshore and offshore conditions although careful site selection and understanding of processes affecting the microfossil record are essential.